D. L. DEONU A Systematic and
Ecological Study
of Nearctic Hydrellia
Diptera: Ephydridae)
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 68
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SMITHSONIAN CONTRIBUTIONS TO
ZOOLOGY
NUMBER 68
A Systematic and
Ecological Study
of Nearctic Hydrellia
(Diptera: Ephydridae)
SMITHSONIAN INSTITUTION PRESS
CITY OF WASHINGTON
1971
ABSTRACT
Deonier, D. L. A Systematic and Ecological Study of Nearctic Hydrellia (Diptera:
Ephydridae). Smithsonian Contributions to Zoology, 68:1-147. 1971.?The adults
of 57 species of Hydrellia are described for the Nearctic Region. Adults of Hydrellia
are semiaquatic and the larvae are leafminers in aquatic and semiaquatic plants.
Twenty-one of the described species are new, and one represents a new geographic
distribution record. Some or all of the immature instars of 18 species are described.
The male terminalia and certain other critical characters are figured for the adults
of 52 species. The larval feeding apparatus of 16 species and the puparia of 17
species are illustrated. Geographic distribution data are given for all species. New
and previously recorded host plants are listed for several species, and other biolog-
ical information such as habitats and behavior is given for most of the species. The
general morphology of the genus, including morphology of the adult feeding appara-
tus and gut and internal genitalia, is discussed and illustrated. The general ecology
of Hydrellia is discussed in regard to ecological role and distribution, parasitological
data, dispersal and zoogeography, behavior, and environmental tolerance. Addi-
tional ecological data are included in checklists of known host plants and known
hymenopterous parasites of Hydrellia. Literature pertinent to the genus is reviewed.
Official publication date is handstamped in a limited number of initial copies and is recorded
in the Institution's annual report, Smithsonian Year.
UNITED STATES GOVERNMENT PRINTING OFFICE
WASHINGTON : 1971
For sale by the Superintendent of Documents, U.S. Government Printing Office
Washington. D.C. 20402 - Price $1.75 (paper cover)
Contents
Page
Introduction 1
Review of Literature 1
Acknowledgments and Repositories 3
Methods and Materials 4
Morphology 6
Glossary 6
External Morphology of Adult 8
Internal Morphology of Adult 13
Morphology of Immatures 15
Ecology 18
Ecological Role and Distribution 18
Parasitological Data 19
Dispersal and Zoogeography 20
Behavior 20
Environmental Tolerance 23
Host Plants and Parasitic Wasps 24
Systematic Treatment 24
Classification 24
Key to Adult Hydrellia 26
Male Terminalia Key to Hydrellia 29
Key to Third-Instar Larvae of some Hydrellia 33
Key to Puparia of some Hydrellia 34
Species Descriptions 36
Checklist of Known Host Plants 106
Checklist of Known Parasitic Wasps 113
Literature Cited 118
Localities of Specimens Illustrated 125
Abbreviations Used on Illustrations 126
Illustrations 128

D. L. Deonier A Systematic and
Ecological Study
of Nearctic Hydrellia
(Diptera: Ephydridae)
Introduction
The Ephydridae constitute a medium-size family
of acalyptrate Diptera. They lead several different
modes of life, mostly in aquatic habitats. Hydrellia
and Lemnaphila are the only known genera of
aquatic leaf-mining ephydrids.
At least two species of Hydrellia?H. griseola and
H. ischiaca?are economically important. In 1953,
H. griseola destroyed from 10 to 20 percent of Cali-
fornia's rice crop, with an estimated loss of $16
million. The same species caused heavy losses of rice
in California in 1922. It has damaged rice in Japan
in the last two decades. Ulljeborg (1861) first re-
corded the pest status of H. griseola. He reported
the fairly widespread damage of barley, oats, and
timothy grass by that species in southern and south-
eastern Sweden during the summer of 1860. After
this, several authors reported outbreaks of H. griseola
in various places in the Palaearctic Region, including
Egypt. Balachowsky and Mesnil (1935) reported
50 percent infestation of barley by H. griseola in
northern Europe. Hydrellia ischiaca attacks wild
rice, which is now a minor crop in Minnesota. Most
of the known larvae of other Hydrellia species feed
mainly in plants of Potamogetonaceae, Alismataceae,
and Hydrocharitaceae.
There are 185 available specific names in Hydrellia
other than my new species. Of these, perhaps 133
have existing holotypes, and perhaps 132 of the 185
have Palaearctic type-localities. Possibly 113 of the
D. L. Deonier, Department of Zoology and Physiology,
Miami University, Oxford, Ohio 45056.
available names are valid. These 113 species are dis-
tributed as follows: 55 Palaearctic, 35 Nearctic, 1
Holarctic, 2 Oriental, 12 Australian, 2 Ethiopian,
and 6 Neotropical. This essentially cosmopolitan
generic distribution has caused some interest in the
dispersal center. Some data possibly indicate this
was in the northern temperate zone.
I started this research with the following objec-
tives: (1) to describe, redescribe, and construct keys
to the adults and as many immature stages as pos-
sible of Nearctic species; (2) to describe the life
cycles and behavior of as many as possible of the
Nearctic species; (3) to present a brief morphology
of the genus.
Review of Literature
The taxonomic literature on Hydrellia dates from
1813, when Fallen described Notiphila griseola and
several congeneric species from southern Sweden.
In 1830, Robineau-Desvoidy erected the new genus
Hydrellia. Macquart (1835), Zetterstedt (1846),
Walker (1856), Loew (1860), Schiner (1864),
Brischke (1883), Gobert (1887), Kowarz (1894),
and Becker (18%, 1903) made many of the initial
contributions to the taxonomy of Palaearctic species.
Strobl (1904), Griinberg (1910), Becker (1919),
Collin (1928), Frey (1933), de Meijere (1939),
Goetghebuer (1942), Grensted (1944), Kloet and
Hincks (1945), and Tsacas (1959, 1960) added some
descriptions of new species, but primarily they pre-
sented reviews and new distribution records. Dahl
(1967) presented a zoogeographical synopsis of
1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
eleven European species and (1968) described three
new species along with new records from eastern
Siberia. Becker's (1896, 1926) synoptic keys to and
descriptions of Palaearctic adult Hydrellia and Hen-
nig's (1943) specific key to larval Hydrellia including
Palaearctic species contributed most to the taxonomy
of Palaearctic species of Hydrellia. The recent re-
vision by Dahl (1964) of the Stenhammar and
Zetterstedt collections eliminated much confusion
and synonymy in Palaearctic Hydrellia.
Loew (1861, 1862) described the first new species
of Nearctic Hydrellia. Loew (1872), Osten Sacken
(1878), Becker (1896), Aldrich (1905), Jones
(1906), and Coquillett (1910a) presented short sec-
tions on Nearctic Hydrellia. Very little additional
taxonomic study of the genus was made until 1915,
when Cresson started describing new species. Johann-
sen (1935) has reviewed and presented a key to the
known immature instars of Nearctic species. Hennig
(1943) has summarized the literature on immature
Hydrellia and presented a specific key to all known
immature instars, including the few in the Nearctic.
Cresson's studies culminated in a key (Cresson,
1944b) to the adults of the 35 Nearctic species then
known. Subsequent to this, Berg (1949, 1950) con-
tributed to the biology and taxonomy, including a
key, of the immature instars of six species; Hennig
(1952) presented morphological interpretations of
immature Hydrellia; Wirth and Stone (1956) in-
cluded a key to California species and a Nearctic
generic key; Grigarick (1959) redescribed the life-
cycle stages of H. griseola; Deonier (1964) presented
keys to the adults of species of Iowa and adjacent
states; and Wirth (1965) cataloged many Nearctic
species.
Cresson (1918, 1947a) described and presented
keys to the six species of Hydrellia known from the
Neotropical Region. Wirth (1968) cataloged these,
and included data on their distribution. Wirth
(1969) reported H. vulgaris Cresson from the Gala-
pagos Islands.
Little study has been made of Hydrellia in the
Oriental Region. Cresson (1948) listed only two
species, H. latipalpis Cresson and H. luteipes Cresson,
from the Oriental Region in his Indo-Australian
synopsis. The latter species is known only from
Formosa, which is more or less transitional between
the Oriental and Palaearctic Regions.
Of Hydrellia in the Australian Region, Coquillett
(1903), Tonnoir and Malloch (1926), Cresson
(1948), and Harrison (1959) published on some or
all of the twelve known indigenous species. The
relatively small amount of attention given to Orien-
tal and Australian Hydrellia is matched only for the
Ethiopian Hydrellia. Cresson (1932, 1947b) listed
two species, both from the Cape of Good Hope.
The works of the following authors provide some
framework for a more searching morphological study
of adult Hydrellia. Becker (1896, 1926), Griinberg
(1910), Cresson (1918), and Wilke (1924) dealt
mainly with chaetotaxy, but Cresson did briefly dis-
cuss facial contour and the supposed absence of
vibrissae in Ephydridae. Frey (1921) discussed the
mouthparts of H. obscura and the cibaria of other
ephydrids. Seguy (1934) illustrated the head and
wing of H. griseola, and Scotland (1940) illustrated
by photographs the proboscis, antenna, and wing of
the closely related genus Lemnaphila.
Hering (1950) briefly described and illustrated the
excised male terminalia of H. xenophaga and H.
nigricans. Kato (1955)'and Kuwayama (1955) de-
scribed and illustrated several aspects of the external
morphology of adult H. griseola. Hennig (1958)
discussed the external morphology of the head and
the thoracic chaetotaxy of several ephydrid repre-
sentatives including H. griseola. With phylogenetic
interpretations as a primary objective, Hennig made
an important contribution here, especially in propos-
ing the presence of vibrissae in Ephydridae. Dahl
(1959) illustrated the adult mouthparts, female
abdomen, and hind tarsus of H. griseola. Grigarick
(1959) illustrated the male and female abdomina,
the wing, and the mesonotum of H. griseola. Harrison
(1959) illustrated the head, wing, and portions of
the male terminalia of five New Zealand species of
Hydrellia. Dahl (1964) illustrated the male ter-
minalia of 17 species of Hydrellia from the Sten-
hammar and Zetterstedt collections. Deonier (1964)
illustrated the head of H. harti and the chaetotaxy
and sclerite nomenclature of the head and thorax
exemplified in Ephydra riparia. Dahl (1968) illus-
trated parts of the male and female genitalia for
three new species of Hydrellia in eastern Siberia.
Sturtevant (1925, 1926) made the only contribu-
tions to the internal morphology of Hydrellia in his
survey of spermathecae in Acalyptratae. Bolwig
(1940, 1941) described and illustrated the internal
genitalia and mouthparts of Scatophila unicornis.
NUMBER 6 8
These studies helped in understanding the mor-
phology of these structures in Hydrellia.
Several authors contributed morphological data
on the immatures of Hydrellia: von Frauenfeld
(1866) described the gross aspects of the metamor-
phosis of H. albilabris; Stein (1867) summarized
briefly the life cycle of H. griseola; Gercke (1879,
1882, 1889) briefly described and illustrated the
puparia and feeding apparatus of H. mutata and
H. fulviceps; and Marchal (1903) figured in gross
aspect the third-instar larva of H. ranunculi.
Though concerned with Ephydra riparia, an in-
vestigation by Tragardh (1903) so lucidly illustrated
the larval feeding apparatus, gut, musculature, and
tracheal system that it formed a basis for anatomical
study of larval Hydrellia. Brocher (1910) examined
the gross morphological, ecological, and physiolog-
ical aspects of the tracheal system' of H. mutata.
Keilin (1915) also studied the metapneustic tracheal
system of larval Hydrellia and illustrated the larval
feeding apparatus. Malloch (1915) briefly described
the larva and illustrated the puparium of H. griseola
(as H. scapularis). Ping (1921), with his morpho-
logical descriptions and illustrations, especially of
the feeding-apparatus musculature of the larvae of
Ephydra riparia (as E. subopaca), provided a very
good basis for such work in Hydrellia. Likewise,
Schiitte (1921), in discovering the phenomenon of
seasonal (summer and winter) forms of the puparia
of Hydromyza livens, provided the starting point for
a similar investigation in Hydrellia which could
answer several ecological questions.
Wilke (1924) and Schoyen (1930) described the
gross morphology of the immature instars of H.
griseola. Collin (1928) illustrated the habitus of the
third-instar larva of H. nasturtii. Johannsen (1935)
briefly described the gross morphology of immature
Hydrellia. Hennig (1943) summarized the known
morphology of immature Hydrellia. Berg (1950)
contributed much to the external morphological
data on immature Hydrellia by describing and illus-
trating most immature instars of six species. Seguy
(1950) referred briefly to some aspects of morpholog-
ical interest in immature Hydrellia. Hering (1951)
commented generally on larval respiration and the
puparial operculum. In 1952, Hennig discussed and
illustrated much of the morphological data on im-
mature Hydrellia. Lange et al. (1953) presented
some significant photographs of the life-cycle stages
of H. griseola. Kato (1955), Kuwayama (1955), and
Grigarick (1959) showed detailed figures of the egg,
feeding apparatus, larval body, puparium, spiracular
peritremes, spinulosity, and setulosity of H. griseola.
Many authors have contributed ecological data on
Hydrellia, and the following have made major con-
tributions: Stormer and Kleine (1911), Sorauer and
Reh (1913), Linnaniemi (1913), Hendel (1926),
and Kreuter (1927) reported some host plants of
Hydrellia, principally of H. griseola; DeOng (1922)
investigated the phenology of H. griseola and its
damage to domestic rice; Hering (1924, 1937, 1951,
1957) contributed much to host-plant data and host
damage of Palaearctic species of Hydrellia; Bala-
chowsky and Mesnil (1935) reported a host-plant
list and host damage for H. griseola; Thompson
(1943) listed some hymenopterans parasitic on a few
species of Hydrellia; Wahlgren (1947) published on
the species of Hydrellia mining in Stratiotes aloides;
Berg (1949, 1950) recorded some larval behavior,
oviposition behavior, and some host-plant species of
six species of Hydrellia; Laurence (1952) described
some entomophagous behavior of H. griseola; Lange
et al. (1953) discussed the biology and control of
H. griseola in California; Grigarick (1959) studied
the ecology of H. griseola in California rice fields;
and Burghele (1959a, 1959b) and Fulmek (1962)
recorded several hymenopterans parasitic in species
of Hydrellia in the Palaearctic Region.
Acknowledgments and Repositories
I acknowledge the following sources of financial
assistance for this research project: General Biolog-
ical Supply House, Inc. (Turtox Scholarship) ; the
National Science Foundation (summer grants at the
Gulf Coast Research Laboratory, Ocean Springs,
Mississippi, 1962; the University of Minnesota Bio-
logical Station, Lake Itasca, Minnesota, 1963; and
the Highlands Biological Station, Highlands, North
Carolina, 1968); the Tennessee Academy of Sciences
(summer grant at the Reelfoot Lake Biological Sta-
tion, Tennessee, 1962); the University of Minnesota
Agricultural Experiment Station (grant for research
on wild rice pests, 1967) ; and Miami University
(grant within United States College Work-Study
Program).
I am grateful to the following individuals: Dr.
C. F. W. Muesebeck of the United States National
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Museum of Natural History for determination of the
parasitic Hymenoptera; Dr. Jean Laffoon of Iowa
State University for initial supervision of the research;
Mr. E. A. Fiser of the Miami University Computer
Center for analyzing certain data; my wife Mary
Deonier, Miss Mary Wolf, and Miss Donna Mutchler
for manuscript typing and proofreading.
The following institutions and individuals loaned
material to me for this research (abbreviations of
those serving as paratype repositories are included
in parentheses) : Academy of Natural Sciences of
Philadelphia (ANSP), American Museum of Nat-
ural History (AMNH), California Academy of
Sciences (CAS), University of California at Davis
(UCD), Canadian National Collection (CNC), Car-
negie Museum, Chicago Natural History Museum,
Cornell University (CU), Illinois State Natural
History Survey, Iowa State University (ISC), Uni-
versity of Kansas (UKC), Kansas State University,
University of Massachusetts (UMM), University of
Michigan, University of Minnesota (UMC), Uni-
versity of Missouri, Museum of Comparative Zoology,
Harvard University (MCZ), University of Nebraska,
Oregon State University, United States National
Museum of Natural History (USNM), Utah State
University, Washington State University (WSC),
Dr. C. O. Berg of Cornell University, Mr. Kenneth
Goeden of the Oregon State Department of Agricul-
ture (KG), Dr. W. W. Judd of the University of
Western Ontario, Mr. George Steyskal of the United
States Department of Agriculture, and Dr. M. R.
Wheeler of the University of Texas (MRW).
My own paratype holdings are labeled (DLD)
under the pertinent new species.
Methods and Materials
COLLECTING OF ADULTS.?I collected adults by
sweeping emergent vegetation, by lowering a killing
tube over them on floating vegetation, and by a
lighted-receptacle method. In the last method, I
inserted the bottom of an open jar into the recessed
lens of a flashlight and beamed the light to floating
leaves. Adults of Hydrellia, Lispe, Hydromyza,
Donaciinae, and a few others flew readily into the
lighted jar. After several adults were in the jar, I
closed it and replaced it with another. This method
failed with H. bilobijera, H. trichaeta, and H. cras-
sipes in North Carolina. Altogether, it was tried on
about ten species.
As a survey method for adults, I used a device
invented and described by Grigarick (1959). As
modified for this project, the device consisted of a
circular aluminum pan 3 cm deep and 25 cm in
diameter inserted in a hole of similar dimensions cut
centrally in a square piece of styrofoam 50 by 3 cm.
Holes in the corners of the styrofoam float accom-
modated dowels run vertically through them and
into the lake bottom to anchor the device. A deter-
gent solution in the pan constituted the actual trap
mechanism. The detergent destroyed the effective-
ness of the tarsal hydrofuge setae so that flies landing
on the solution sank immediately. In these traps I
caught specimens of Donaciinae, Gerridae, Veliidae,
Hydrometridae, Hydrophilidae, Tridactylidae, Col-
lembola, Hymenoptera, and several dipterous fami-
lies in addition to those of Hydrellia.
I made most of my behavioral observations while
collecting. After observing the behavior of the indi-
vidual, I collected it with a killing tube or live-
capture tube. I tried to keep some captured adults
alive in the laboratory long enough to make addi-
tional observations or for inseminated females to
oviposit.
PREPARATION OF ADULTS.?I killed and preserved
in 80 percent ethanol specimens intended for anatom-
ical and food-habits studies. I used a hydration
series on these specimens before dissecting or section-
ing them. I hydrated some specimens just before
dissection and some just after fixation in Bouin's
solution or dioxane in preparation for serial sec-
tioning.
To clear terminalia, I placed them in a hot, 10
percent potassium hydroxide solution for about 3
minutes and then added two to ten drops of 30
percent hydrogen peroxide over and around the
floating terminalia. By closely watching the termi-
nalia, I could ascertain when they were sufficiently
bleached and desclerotized. The time required for
this combined clearing and bleaching process varied
with the condition of the terminalia and with the
species. The average time was about 3.5 minutes?
3 minutes in hot caustic followed by a few seconds
in the mixture of peroxide and caustic. Terminalia
not quickly removed after the critical time limit often
were overcleared and not usable. Apparently, the
oxygen released from the hydrogen peroxide acts
as the bleaching agent. After passing the terminalia
through glacial acetic acid, distilled water, and
NUMBER 6 8
absolute ethanol, I placed them in a small drop of
glycerol in a depression slide for microscopic study.
I used an ocular grid and squared paper to draw to
scale the terminalia and other adult structures.
Except for the structures that require a bilateral or
whole view for proper interpretation, I show only
the left half of bilaterally symmetrical structures in
most drawings. I drew most of the adult structures
with the aid of a compound microscope.
COLLECTING OF IMMATURES.?In collecting the
immatures of Hydrellia, I first searched for eggs on
or near potential host plants in the habitats of the
adults. When I discovered eggs I placed the entire
plant on which they were laid into a plastic bag.
After searching for eggs, I sampled the potential host
plants in the locality for larvae and puparia. This
was a random process in regard to larvae, for they
can be found in situ most readily in the laboratory
with strong direct illumination and a stereomicro-
scope. In sampling some submergent plants in
depths greater than 1.3 meters, I used a pike pole
or employed a scuba diver. In studying the over-
wintering of some Hydrellia, I used an ice chisel to
cut through the ice to obtain samples of some host
plants.
In the laboratory, I examined all parts of each
plant for immature Hydrellia by holding the plant
part before a strong light. When there was evidence
of mining, but no larvae apparent, I examined the
tissue with a stereomicroscope.
REARING.?When I discovered an egg or larva,
I removed the plant part in or on which it was
situated and placed the part in a small culture dish
with tap water. I isolated each puparium together
with some of the surrounding plant part in a 75-ml
test tube containing a small amount of tap water
and loosely plugged with cotton.
I observed and recorded behavior (larval eclosion,
molting, and ecdysis, mining activity, pupariation,
adult emergence, etc.) and associated morphological
changes daily. During these microscopic examina-
tions, I would occasionally find a first- or second-
instar larva previously undetected. In almost all
cases I could distinguish between it and the original.
This difficulty in detecting first-instar and some
second-instar larvae existed because of their small
size, translucency, the greater depth and shorter
length of the mine. Because of this difficulty, I
kept all plant material examined in screen-covered
aquaria. I examined all of this plant material for
immature Hydrellia periodically for two weeks.
I prepared voucher specimens of each plant spe-
cies examined for Hydrellia and also of some species
collected for ecological indicators. Because of the
large number of plants of many species examined,
I could not preserve each plant found to be a host
of Hydrellia. These specimens are on deposit at the
Iowa State University Herbarium.
Since temperature control was unavailable for the
rearing work, I attempted only to record the tem-
perature of the water in which specimens were
reared. During the summer of 1963 at the University
of Minnesota Biological Station and at most times
thereafter to the present, I used a circular thermo-
graph with sensor element kept in a beaker of tap
water for recording daily temperature fluctuations
in the laboratory. I changed the tap water around
the sensor each time I changed the tap water in
the rearing vessels.
PREPARATION OF IMMATURES.?For species of
Hydrellia for which I had collected sufficient speci-
mens of each of the immature instars, I preserved
some of each instar, but for several species I had so
few specimens that I allowed all to develop to the
adult instar.
I preserved some Hydrellia eggs in 80 percent
ethanol and some between filter-paper strips moist-
ened with AFA and placed in small shell vials
(about 5-ml capacity). I plugged the vials with
cotton and immersed themf in either ethanol or AFA
in large museum jars. I preserved the larvae and
puparia by the same method, after killing them in
hot water. To insure good specimens for anatomical
and food-habits studies, I used AFA, but it had the
one deleterious effect of bleaching the chlorophylls
and carotenoids in the larval gut contents.
For stereomicroscopic study, I placed larvae in
80 percent ethanol in depression slides. To study
the same material with a compound microscope, I
had only to add a few drops of glycerol to the
ethanol in the concavity. I used this method for
studying, drawing, and photographing eggs, larvae,
and puparia.
To store egg choria and puparia from which adults
had been reared, I placed them in small drops of
glycerol in the bottom of cork-stoppered, glass micro-
vials and placed them on the pin holding the cor-
responding adult. This method afforded a flexibility
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
in studying specimens that was not afforded in the
use of permanent glass-slide mounts; also, it kept
most of the life-cycle stages together.
Because of the microscopic size of some structures,
e.g., setulae, spinules, and some other cuticular proc-
esses, I did not risk clearing immature specimens
with potassium hydroxide. Clearing was necessary
only infrequently, and then it usually involved parts
of puparia, for which a very mild agent such as
methyl salicylate sufficed.
I used the same procedures and materials for
drawing structures of immatures as I did for adults,
except that I made the habitus drawings of puparia
initially with a stereomicroscope and then filled in
some details under a compound microscope.
Morphology
Glossary
New structures, configurations, and indices encoun-
tered in a taxonomic study must be named. Since
zoological terminology is already burdened with
much ambiguity, I constructed the following defini-
tions and explanations of structural and index
names. In these, all distance measurements are
straight-line and uniplanar on preserved specimens
unless otherwise specified.
TERMINOLOGY FOR ADULTS
A-index: The quotient of the subcranial breadth divided
by the anteclypeal breadth. (Figure 11.)
Anteclypeal breadth (acb): The maximum transverse
distance between the outer edges of the paraclypeal
phragmata of the anteclypeus. (Figure 11.)
Anteocellar distance (aod) : The distance between the
anterior margin of the median ocellus and the upper
edge of the ptilinal fissure along the frontal midline.
(Figure 11.)
Anterior pronto-orbital (afr): The anterior seta of the
two setae commonly prominent in each fronto-orbital
area. (Figure 1.)
Apicodorsal antennal (a3ap): A spinous seta on the
dorsal apex of antennal segment 2.
Aristal rays (arr): All of the trichoid projections of each
arista including the apical one. (Figure 1.)
B-index: The quotient of the maximum anteroposterior
extent of the fused surstyli divided by the length of the
sclerotized portion of the cercus as measured in ventral
view. (Figure 4.)
Basal coxal (be) : A seta inserted laterally on the mid
coxa. (Figure 6.)
Basal end of costa (bee): The enlarged basal portion of
the costa proper adjacent to the humeral plate. (Figure
135.)
Body length: The distance between the most prominent
part of the face and the posterior end of the abdomen.
It is measured in lateral view and as if the head and
abdomen were aligned horizontally.
C-index: The quotient of the midline anteroposterior
extent of sternum 5 divided by the projected length of
the postgonite uncus.
Color: The descriptions of color apply to views perpendi-
cular to the sclerite concerned unless otherwise stated.
Color designations follow the ISCC-NBS method (Kelly
and Judd, 1955).
Copulobus (cl): One of a pair of posterior lobes, or
projections, of male sternum 5. (Figures 4, 136?138.)
Costal section I: The distance between the distal edges
of crossvein h and Rt apex. (Figure 135.)
Costal section II: The distance between the distal edges
of Rj and R2 + 3 apices. (Figure 135.)
Costal section III: The distance between the distal edges
of R2 + 3 and R4 + 5 apices. (Figure 135.)
Costal section IV: The distance between the distal edges
of R4 + 5 and M t + 2 apices. (Figure 135.)
Costal section V: The distance between the distal edge
of Mj + 2 apex and the proximal edge of M3+CU1 apex.
(Figure 135.)
Epistomal breadth: The transverse distance between the
pair of primary facial rows at the level of the epistoma,
measured from the inner edge of the setal sockets. (Figure
1.)
Epistomal index: The quotient of the epistomal breadth
divided by the minimum interocular distance .(Figure 1.)
Frontal vitta {mj): The median quadrangular area of
the frons on which are situated the ocellar triangle and
the ocellar and postocellar setae. (Figure 1.)
Fronto-orbital area: The narrow lateral section of each
parafrontale parallel and contiguous to the compound
eye on which the fronto-orbital setae are inserted. This
usage differs from that of some other authors. (Figure 1.)
Interfractural costal: A seta inserted on the costa between
the two costal "fractures." (Figure 135.)
Laterotergite (It): The metapleuron of other terminology.
(Figure 6.)
M1 + g index: The quotient of the distance between the
distal edge of the junction of crossvein m and Mj + 2
divided by the distance between the distal edge of the
m and M t + 2 junction and the distal edge of the r-m
and M 1 + 2 junction. (Figure 135.)
Mesanepimeron (aem^) : Part of the pteropleuron of other
terminology. (Figure 6.)
Mesanepisternum (aes2): The mesopleuron of other ter-
minology. (Figure 6.)
Mesofacial height (mfh): The distance between the outer
edge of the subcranial cavity and the lower edge of the
ptilinal fissure along the midline of the face. (Figure 11.)
Mesofacial index: The quotient of the mesofacial height
divided by the minimum interocular distance.
Mesokatepisternum (kes2): The sternopleuron of other
terminology. (Figure 6.)
NUMBER 6 8
Metapleuron {mp): The hypopleuron of other terminol-
ogy. (Figure 6.)
Minimum interocular distance: The minimum transverse
distance between the compound eyes in the area of the
face. (Figures 1, 11, 139.)
Ocular height (vod): The maximum distance between
the upper and lower edges of the compound eye. The
line of measurement is not quite vertical in most species.
(Figure 9.)
Ocular index: The quotient of the nearly vertical ocular
height divided by the subocular height. (Figure 9.)
Parafrontale (If): The region of the frons between the
frontal vitta and the upper edge of the compound eye.
(Figure 11.)
Postdorsocentral (pdc): A seta inserted in the dorsocentral
line posterior to the transverse sulcus. (Figure 6.)
Posterior fronto-orbital (pfr) : The posterior seta of the
two setae commonly prominent in each fronto-orbital
area. (Figure 1.)
Postgonite (pog): One of a pair of curved-, fingerlike
projections of the gonal arch on each side of the disti-
phallus. In terminalia preparations, the postgonite usually
appears to be anterior to the pregonite. (Figures 4, 10,
13, 136, 137.)
Postgonite uncus (pu) : The apical section of the postgo-
nite which is often hooklike or clawlike and usually dis-
tinctly more heavily sclerotized than the remainder of
the postgonite. (Figures 10, 13.)
Postocular: One of the setae inserted in a row posterior
to and more or less parallel with the posterior edge of
each compound eye. (Figure 9.)
Pre dor so central (adc): A seta inserted in the dorsocentral
line anterior to the transverse sulcus. (Figure 6.)
Pregonite (prg): One of a pair of bifurcate setose pro-
jections of the gonal arch on each side of the basiphallus.
In terminalia preparations, the pregonite usually appears
to be posterior to the postgonite. (Figures 4, 10, 13.)
Primary facial (pfa): One of the longer facial setae in-
serted in a row on each side of the face parallel and
medial to the ptilinal fissure. The primary facial row
is parallel to the outer edge of the obscured epistomal
sulcus. (Figure 1.)
Secondary facial (sfa): One of the shorter facial setae
that are often in a row parallel and lateral to the primary
facial row.
Subcranial breadth (scb): The maximum transverse dis-
tance between the outer lateral edges of the subcranial
cavity. (Figure 11.)
Subocular height (soh): The minimum distance between
the lower edge of each compound eye and the outer
lateral edge of the subcranial cavity. (Figure 9.)
Vertex breadth (vb): The distance between the upper
edges of the compound eyes at the level of the lateral
ocelli. (Figure 11.)
Vertex index: The quotient of the vertex breadth divided
by the anteocellar distance. (Figure 11.)
Wing length: The distance between the apex of the tegula
and the wing tip. (Figure 135.)
TERMINOLOGY FOR IMMATURES
Anal-plate index: The quotient of the transverse extent
of the anal plate divided by the midline anteroposterior
extent of the anal plate. (Figure 107.)
Bifurcation index: The quotient of the longitudinal dis-
tance between the level of the phragmatal bifurcation
and the posterior end of the ventral phragmatal ramus
divided by the minimum distance between the end of the
dorsal phragmatal ramus and the upper edge of the
ventral phragmatal ramus. (Figure 85.)
Clypeal arch: The area of inclination in the frontoclypeus
just anterior to the cheliform spot. (Figure 85.)
Cly'peal-arch index: The dorsoventral extent of the fronto-
clypeus at the level of the anterior edge of the cheliform
spot divided by the dorsoventral extent of the fronto-
clypeus at the level of the anterior edge of the labial
gland orifice. (Figure 85.)
Early pupa: The pupa as it appears prior to the time
when its compound eyes become apparent.
Egg length: The distance between the ends of the egg as
measured in dorsal view. (Figures 75, 77, 78, 125.)
Frontoclypeal length: The distance between the anterior
edge of the frontoclypeus and the posterior end of the
ventral phragmatal ramus as measured in lateral view.
(Figures 84, 75.)
Late pupa: The pupa as it appears after the time when
its compound eyes become apparent.
Larval length: The distance between the anterior edge
of the head lobe and the posterior end of the spiracular
peritreme measured with the larva outstretched. (Figure
124.)
Maximum egg breadth: The maximum transverse extent
of the egg as measured in dorsal view. (Figures 75?78.)
Maximum larval breadth: The maximum transverse ex-
tent of the larva as measured in dorsal view.
Maximum mouth-hook base thickness: The maximum
thickness of the articulated end of the mouth-hook as
measured in lateral view. (Figure 85.)
Maximum mouth-hook beak thickness: The maximum
thickness of the free (or beak) end of the mouth-hook
just distal to the enlarged base as measured in lateral
view. (Figure 85.)
Minimum puparial breadth: The minimum transverse
extent of abdominal segment 8 as measured in dorsal
view anterior to the tracheospiracular siphon. (Figure
107.)
Phragmatal index: The quotient of the longitudinal dis-
tance between the anterior edge of the frontoclypeus and
the level of the phragmatal bifurcation divided by the
longitudinal distance between the level of the phragmatal
bifurcation and the posterior end of the ventral phrag-
matal ramus. (Figure 85.)
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Puparial length: The distance between the anterior pro-
thoracic margin of the puparium and the posterior end
of the spiracular peritreme measured as if the puparium
were outstretched. (Figure 107.)
Ventral frontoclypeal index: The quotient of the distance
between the anterior edge of the frontoclypeus and the
anterior edge of the labial gland orifice divided by the
dorsoventral extent of the frontoclypeus midway between
the anterior edge of the frontoclypeus and the anterior
edge of the labial gland orifice. (Figure 85.)
External Morphology of Adult
HEAD.?There are several controversies concerning
the morphology of the schizophoran head. The
major of these is centered on the clypeus and frons.
Snodgrass (1935, pp. 317, 322, 323), in elucidating
the morphology of the clypeus, stated:
In the higher Diptera, the median part of the clypeus
becomes an independent sclerite, but the dilator muscles
of the pump retain their attachments upon it. . . . The
inverted V?shaped plate of the anterior wall of the rostrum
(Fig. 174, C, D, dp) bears upon its lateral arms the
origins of the dilator muscles of the cibarial pump (D, 3).
There can be little question, therefore, that this sclerite
represents at least the median part of the clypeus in the
head of Tabanus (Fig. 171 B, dp). . . . The attachment
of the dilator muscles of the cibarial pump on the arms of
the V-shaped rostral plate, however, clearly demonstrates
the clypeal origin of this sclerite, and confirmatory evidence
of its homology with the median clypeal region in Tabanus
is seen in the fact that a pair of labral muscle? (Fig 174 D,
2) take their origin on its dorsal part. The smaller sclerite
above the V-shaped clypeal plate (C, e) is either a part
of the clypeus or a secondary sclerotization hinging the
latter to the lower margin of the face.
In his discussion, Snodgrass did not commit himself
on the morphology of the region of the head capsule
between the subcranial margin, antennal sockets,
and the compound eyes, but by emphasizing that the
hinged anterior plate of the basiproboscis is homolo-
gous with the median clypeal region in Tabanus he
inferred that some part of the clypeus remains above
the subcranial margin. In a later paper Snodgrass
(1944, p. 70) withdraw this inference by stating that
"In the Cyclorrhapha and some of the Brachycera,
the median, muscle-bearing plate of the clypeus be-
comes isolated by a membranization of the surround-
ing clypeal area, and is thus flexible on its hinge
with the frons." Between 1935 and 1944, Snodgrass
decided that the muscoid homologue of the tabanid
median clypeal plate was hinged "with the frons"
and not "to the lower margin of the face." In 1953
he omitted any discussion of the partial membran-
ization of the clypeus and the boundaries of the
frons in explanations of the evolution of the dip-
terous cibarium and proboscis.
Snodgrass never explained the fate of the epis-
tomal (frontoclypeal) sulcus, but since, by the
accepted definition, the epistomal sulcus is the ex-
ternal cuticular furrow, or groove, between the ante-
rior tentorial pits which delimits clypeus and frons,
I must assume that in the Snodgrass view the ante-
rior tentorial pits and epistomal sulcus disappeared
completely by membranization in the evolution of
the schizophoran head capsule.
Kim and Cook (1966, p. 79) stated that "The
clypeus (tormae of Peterson (1916) ) lies below the
ventral margin of the prefrons (Figures 1, 2, 10, 11)
between the labrum and the frons and is anteriorly
delimited by the membrane above the labrum and
posteriorly by the membranous frontoclypeal suture."
These authors apparently failed to answer the ques-
tion of how the frontoclypeal (epistomal) suture or
sulcus, which they interpreted to be the basiproboscis
membrane between anteclypeus and head capsule
proper, became entirely disconnected from its origin
in the anterior tentorial pits. They stated: "The
anterior tentorial pit is indistinct externally in
sphaerocerids but is situated at the upper lateral
angle of the prefrons below the eye and above the
frontal ridge" (Kim and Cook, 1966, p. 79).
Frick (1952) and Downes (1958) showed some
evidence of the epistomal sulcus in Agromyzidae and
Sarcophagidae respectively. Frick (1952) labeled
what he considered the anterior tentorial pits in the
apparent epistomal sulcus in Agromyzidae. My in-
vestigation of the problem in Hydrellia indicated the
validity of the interpretations of Frick and Downes.
As shown in Figures 1, 11, 139, the epistomal, or
frontoclypeal, sulcus extends from the lateral sub-
cranial margins dorsad along the primary facial rows
as faint internal cuticular thickenings. These thick-
enings were visible only after clearing, and they were
not connected above in the two species studied
closely. The epistomal sulcus is incomplete in several
species of insects. The ptilinal fissure apparently
extends ventrolaterad from its conspicuous supra-
antennal arc to the subocular genal regions. These
paraocular extensions delimit the lateral facial areas
next to the orbits as the parafacialia. Downes (1958)
labeled the facial areas between the extensions of the
NUMBER 6 8
ptilinal fissure and the epistomal sulcus as facial
ridges, while, for some reason, Frick (1952) labeled
the same areas parafacial regions. Frick did not
label the areas contiguous to the eyes and set off by
the ptilinal fissure extensions.
Bolwig (1941) illustrated the upper extremities of
the epistomal sulcus in Scatophila unicornis Czerny
(Ephydridae), but he interpreted them as dorsal
tentorial pits. It is possible that Bolwig (1941, p. 3)
was nearer to the truth than Snodgrass, Frick, or
Downes when he stated "that those extending from
the impression beneath the antennae to the mouth
opening [subcranial cavity] (d.t.) are homologous
with the dorsal arms of the tentorium. The lateral
thickenings (ant.t.) of the edge of the mouth open-
ing are then supposed to be homologous with the
anterior arms of the tentorium, while the thicken-
ings stretching from the occipital foramen down-
wards (pt.) to the mouth opening are supposed to
be homologous with the posterior arms of the ten-
torium." This concept of continuous sulci from
dorsal tentorial pits through anterior tentorial pits
to posterior tentorial pits can be visualized in illus-
trations by Bonhag (1951, figs. 1-4). For Bolwig's
interpretation to obtain, the anterior tentorial pits
need have shifted posteroventrad only a short dis-
tance. It is perhaps impractical with our present
knowledge to attempt to distinguish pit and sulcus.
In the classical view, the thickening between dorsal
and anterior tentorial pits would be the epistomal
sulcus, while the thickening between anterior and
posterior tentorial pits would be the subgenal sulcus
(in part, hypostomal sulcus).
In my synthesis of several concepts, I have at-
tempted to present a working scheme very similar to
that used in modern calypterate taxonomy. In this
collation, the question of the limits of postclypeus
and the frons was considered moot and relatively
unimportant because tradition has favored such
terms as mesofacial plate, medifacies, medifacial
plate, and facial plate and because specialization in
cibarial muscles has obscured ordinary divisions. I
have called the anterior edge of the subcranial cavity
between the extremities of the epistomal sulcus the
epistoma (Figures 1, 11, 139). There is considerable
precedent for this designation, and the term is exten-
sively used in chaetotaxy. The area superior to this
epistoma between the indistinct arms of the epis-
tomal sulcus I have called the facial plate (Figures
1, 11, 139). The ptilinal fissure forms the dorsal and
lateral limits of the face. Frick (1952) called this
area the mesofacial plate and placed the antennal
sockets as the upper limit. Downes (1958) termed a
similar area in Sarcophagidae the facial plate. The
antennal sockets may be considered to demarcate a
facial subregion above and between them and the
ptilinal fissure called the frontal lunule.
The cuticular area bounded by the lower edge of
the compound eye, the lateral subcranial margin, the
lower angle of the face, and the posterolateral post-
genal inflection is commonly called the gena. Since,
however, the gena is rather ill-defined, I have used
the term subocular height in defining the ocular
index (Figure 9). The term defined is identical to
the genal height illustrated by Dahl (1959), but it
avoids the ambiguity of the gena concept, for the-
oretically much of the immediate postocular area is
also part of the gena.
Above and posterior to the ptilinal fissure is the
frontal vitta, bordered on each side by an indistinct
parafrontale, or genovertical plate. It often is conven-
ient to distinguish a narrow paraocular section of the
parafrontale as the fronto-orbital area. The frontal
vitta and the parafrontalia constitute the frons. Frick
(1952) distinguished only the frontal vitta, and
lateral to it a genovertical plate, which is apparently
synonymous with the parafrontale. At the vertex in
the frontal vitta, the ocellar triangle protrudes only
slightly in Hydrellia. Posteroventrad of the ocellar
triangle, the frons transcends with the occiput, i.e.,
the occipital sulcus is absent. The incomplete post-
occipital sulcus only partially defines the postocciput
with its inconspicuous occipital condyles.
One can distinguish three major divisions of the
proboscis: the basiproboscis extending from the sub-
cranial margin to and including the bases of the
maxillary palpi, the mediproboscis extending from
the bases of the maxillary palpi to the distal end of
the hypopharynx, and the distiproboscis extending
from the distal end of the hypopharynx to the ex-
tremities of the labella (Figures 1, 134, 139). In the
basiproboscis, the conjunctiva (membrane) encloses
the cibarium, the paraclypeal phragmata, and the
stipes. Only the anteclypeus, or rostral plate, and the
maxillary palpi are exposed (Figures 134, 139, 140).
In the mediproboscis, the sclerotized prementum is
the prominent structure. Anteromedially from the
prementum is the labial gutter containing the hypo-
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
pharynx and the labrum. Enclosed within the con-
junctiva basal to the labrum are the labral apodemes
and the trachea-like siphon connecting the labial
gutter with the cibarium. The distiproboscis con-
sists essentially of the labella. Each labellum is par-
tially supported above by a labellar sclerite and con-
sists of seven canaliculi which radiate from the
margin of the prestomum. Apparently, the canaliculi
terminate in the prestomum posterior to the apparent
sclerotized prestomal margin. Each canaliculus has a
row of several canalicular teeth.
The schizophoran cibarium is homologous with
the generalized orthopteroid cibarium, i.e., it is
the chamber between the connected epipharyngeal
clypeal wall and sitophore (anterior and posterior
cibarial walls). Since in Diptera the chamber has a
pumping function, it is stabilized either by antagon-
istic muscles as in Nematocera and many Brachycera
or by inflections of the lateral borders of the ante-
clypeus, called paraclypeal phragmata.
Within the cibarium, inserted on the anterior, or
epipharyngeal, wall are small trichoid sensilla: paired
clusters of three near the distal end of the cibarium
and a biseriate row running dorsolaterally from each
of these initial clusters to the end of the anteclypeus
(Figures 1, 140-142). Frey (1921) apparently first
illustrated these cibarial sensilla in Hydrellia and
other ephydrid genera, but he simply called them
setae as did Bolwig (1941). I closely examined and
counted these cibarial sensilla only in several speci-
mens of H. griseola, but I confirmed their presence
in several congeners and in representatives of all
dipterous suborders. The number and arrangement
of the cibarial sensilla apparently varies somewhat in
Hydrellia, for Frey (1921, p. 137) stated: "Die obere
Pharynxwand hat 2 Reihen von 5-6 Borsten, wozu
vorn jederseits eine Gruppe von 3 kiirzeren, dichter
gestellten kommt." This concerned H. obscura
Meigen. In H. griseola, I found four pairs of sensilla
in the two long rows.
Regarding the function of the cibarial sensilla, I
offer the following hypothesis. The cibarium is the
first and primary ingestion pump, and among the
Schizophora the sole pump. This being the case,
proprioceptors are necessary to maintain cibarial
rhythm. The cibarial pump must perform the initial
ingestive suction and in Schizophora the subsequent
expulsion from the cibarium into the esophagus. To
perform these two progressive functions, the shape
of the cibarium must change rhythmically, for no
valve has been demonstrated distal to the cibarium
to prevent backflow from the contracting cibarium.
My hypothesis is that during the initial ingestive
phase the cibarium dilates progressively proximad,
and during the subsequent emptying it constricts
progressively proximad, and that the cibarial sensilla
coordinate or govern these rhythmic changes. As
proprioceptors, these sensilla enable the cibarium to
act as a valve as well as a pumping chamber. As
they touch the opposite wall, impulses pass from
their neurones. In this way, regurgitative loss of
cibarial contents is reduced.
Before conducting this investigation, I thought the
high rates of proboscis protraction and retraction
observed in Parydra (Ephydridae) was evidence that
pinching of the siphon connecting the food meatus
and cibarium prevented regurgitative cibarial loss.
The absence of such repeated flexion in many
Schizophora, including Hydrellia, controverted this
interpretation.
Bonhag (1951) illustrated and described a func-
tional mouth (and valve) just distal to the cibarium
operated by muscles termed the labral compressors.
If this functional mouth exists in the Schizophora,
it can only be the trachea-like siphon of the food
meatus. I did not find the unpaired labral com-
pressors in the preparations of H. griseola nor did I
find the median labral process on which they would
be inserted.
The cranial chaetotaxy differs very little from that
common in Acalyptratae. The conspicuous macro-
chaetae are the genal, inner and outer verticals, and
postocellar. The ocellar, anterior and posterior
fronto-orbitals, apicodorsals of antennal segment 2,
postocular, and primary facial setae are less con-
spicuous. In most species, smaller secondary facials
occur, usually lateral to the primary facial row.
According to Hennig (1958), the vibrissa is pres-
ent but indistinguishable in Ephydridae. Also,
according to Hennig, the postocellar has replaced
the postvertical at least in prominence and approxi-
mate location.
THORAX.?My presentation of the gross external
morphology of the thorax proper of Hydrellia is
essentially an interpretative synthesis of the contri-
butions of Osten Sacken (1881), Curran (1934),
Snodgrass (1935), Comstock (1940), Crampton
(1942), Bonhag (1949), and Downes (1955).
NUMBER 6 8 11
The prothorax consists of the greatly reduced
antepronotum as part of the border of the pro-
thoracic foramen, the postpronotum, the propleura,
the prosternum, and the fore legs (Figure 6). The
prosternum consists of a small presternum and a
considerably larger probasisternum with a median
sulcus and a prosternellum, the defining sternacostal
sulcus of which is continuous with the median
probasisternal sulcus. I am uncertain about the
nature of an apparent sclerite just above and pos-
terior to pleurocoxal condyle 1 (Figure 6) , but it is
similar to the proepimeron illustrated by Crampton
(1942) andBonhag (1949).
The anterior spiracle?or, as I have called it, the
prothoracic spiracle according to Keilin's (1944)
analysis?is situated in a small fossa inferior to the
postpronotum and posterior to the superior termina-
tion of the propleural sulcus.
The mesonotum is divided by the transverse and
scutoscutellar sulci and the postscutellar suture into
the notopleuron, mesoscutum, mesoscutellum, and
mesopostscutellum respectively. The mediotergite
and the laterotergites are the ordinary discernible
sclerites of the mesopostscutellum, but often an
anatergite and katatergite can be distinguished as
constituents of the laterotergite. The expanded
mesopleura, each divided by the mesopleural sulcus
into the mesepisternum and the mesepimeron (Fig-
ure 6) , the wings, and the mid legs are the remaining
main constituents of the mesothorax.
The dorsopleural (or notopleural) sulcus sepa-
rates the notopleuron and the mesopleuron. At the
anterior end of this sulcus is a slightly distinguish-
able sclerotization, which may represent part of the
mesoprescutum. Bonhag (1949) illustrated the
mesoprescutum as a pair of small triangular lobes
interposed between the evident notopleura and
mesopleura; however, Snodgrass (1935) illustrated
the notopleura as the prescutal lobes.
The large mesepisternum is partially divided by
the incomplete mesanepistemal sulcus into the
mesanepisternum and the mesokatepisternum. The
posterior part of the mesanepisternum is traversed
by a secondary suture, or membranous cleft. A con-
spicuous sclerite occupies the upper corner of this
cleft. Crampton (1942) showed a sclerite which
resembles this one except for its apparent fusion with
the posterior mesanepistemal lobe isolated by the
secondary cleft. Crampton called this sclerite the
anterior basalare. Downes (1955) showed two
sclerites, basalarites A and B situated in the upper
part of this cleft. Basalarite A is separated and occu-
pies the uppermost part of the cleft. Basalarite B is
connected by a narrow neck to the isolated
mesanepistemal lobe. Bonhag (1949) showed two
separate sclerites occupying this cleft in Tabanus.
The anterior sclerite called the basalare extends down
to the mesokatepisternum. Bonhag did not name
the small, second sclerite in the uppermost part of
the cleft. The arrangement of sclerites in the secon-
dary cleft and the course of the mesopleural sulcus
in Hydrellia approximates most closely Downes'
illustration except that the basalar ampulla has
evidently been displaced posteriad. The subalar
ampulla is a distinct crescentic sclerotization in the
lower wing area.
The mesepimeron is distinctly represented only by
the mesanepimeron. The mesokatepirneron is prob-
ably represented by the small sclerite contiguous to
the anterior edge of the metathoracic spiracular
peritreme. It is possible that this small sclerite is
part of the metapleuron, as may be inferred from
Crampton's illustration. According to Crampton,
the mesokatepimeron is usually indistinguishably
fused with the large meron (meropleurite). Downes
showed the mesokatepimeron to be slightly distin-
guished by the coxopleural streak from the upper
meral margin. The laterotergite extends down
farther in Hydrellia than shown by Crampton,
Bonhag, and Downes. This fact and the partial
demarcation of a small triangular lobe anterior to
the metathoracic peritreme illustrated by Bonhag led
me to emphasize the first-mentioned interpretation
of the location of the mesokatepimeron.
The precoxale (anterior to and above the mid
coxa) of the mesokatepisternum is distinguished as
a narrow glossy sclerotization. The posterior meral
margin is similarly distinguished.
The apparent ventral extension of the meso-
katepisternum is considered by Snodgrass (1935) a
composite of the sternum, precoxale, and episternum.
Basically, Bonhag differed little or not at all from
this view.
Of the metathorax, only thoracic phragma 2, the
greatly narrowed metapleura, the halteres, and the
hind legs remain distinct (Figure 6) . The meta-
pleuron consists of a metathoracic precoxale, appar-
ently defined by the lower part of the sclerotized
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
posterior meral margin and a continuation of this
heavy sclerotization posteriad of coxa 3, the metepis-
ternum, which is sharply narrowed and apparently
divided by a transverse sulcus near the posterior
meral lobe, and the very linear metepimeron behind
the indistinct metapleural sulcus.
The wing-vein nomenclature is modified from the
Comstock-Needham system as illustrated by Downes
(1955). Because of taxonomic importance, I added
the designation "basal end of the costa" (Figure
135). The halter has three apparent divisions: the
knoblike capitellum, the pedicel, and the basal
scabellum.
My thoracic chaetotactic nomenclature is modi-
fied from Osten Sacken (1881), Curran (1934), and
Cornstock (1940). The main modifications are in
the pleural setae, which are named according to
the sclerite on which they are inserted, e.g., post-
pronotals, propleural, mesanepisternals, mesokatepis-
ternal(s), and basicoxal(s). On the mesonotum,
the dorsocentrals and acrostichals anterior to the
incomplete transverse sulcus are termed predor-
socentrals and preacrostichals, while those posterior
to this point are called the postdorsocentrals and
postacrostichals. The prescutellar macrochaeta is
often considered a dorsocentral, though it is inserted
between the dorsocentral and acrostichal lines.
There is one large postalar macrochaeta and a small
one inserted midway on the postalar bridge. More or
less in line with the large postalar and the prescutel-
lar is one large interalar (corrected from "intraalar"
by Sturtevant and Wheeler, 1954). Above the meso-
pleural wing process are two small supra-alar setae.
There are two notopleural macrochaetae and one lat-
eral macrochaeta just anterior to the notopleural
apex. Inserted on the scutellar margin are the basal
scutellar, intermediate scutellar, and the apical scu-
tellar.
The predominant features of the wing are the two
costal fractures or discontinuities, one just proximal
to Ri apex and one slightly distal to crossvein h.
Although the whole costa is setose, the setae on the
enlarged basal end of the costa and the setae between
the costal fractures, the anterior and the dorsal inter-
fractural costals, are of main taxonomic importance
(Figure 135).
ABDOMEN.?Except for dimensional and color
characters, secondary sex characters in Hydrellia
appear to be limited to the abdomen, especially the
postabdomen. Only Hering (1950), Dahl (1959,
1964), Grigarick (1959), and Harrison (1959) have
studied the abdomen other than superficially.
It is convenient and traditional to distinguish two
abdominal regions?the preabdomen and the post-
abdomen. According to Steyskal's (1957) hypothet-
ical illustration, the division in the male abdomen
should be between terga 6 and 7. Unfortunately,
these two terga are obliterated in male Hydrellia so
that, from a practical view, I placed the division
just posterior to tergum 5 in both sexes. Thus, in my
discussion, the first five segments (except sternum 5)
constitute the preabdomen and the remainder con-
stitute the postabdomen. The male sternum 5 must
also be considered part of the terminalia (Figures
4,5,136-138) in Hydrellia.
Tergum 1 is partially fused with tergum 2 and is
so short that it is very much obscured in dorsal view
in its normal relation to the thorax. This condition
led some earlier workers to assume there were four
instead of five preabdominal terga. This assumption
was enhanced by the slight displacement posteriad
of sternum 1 and the highly modified form of sternum
5 in the male (Figure 4) . Although there is some
assumption in simple sequential identification of the
sterna, the condition of the female sternum in H.
griseola as shown by Grigarick (1959, pi. 1, fig. 3)
seems to support this course.
I have illustrated the male postabdomen in Fig-
ures 4, 5, 10, 12, 13, 136-138. Snodgrass (1957)
believed that his analysis of the evolution and
homologies of external male genitalia could be
applied to all insect groups, but he failed to do this
for the higher Diptera. Insufficient morphological
knowledge has precluded the homologizing of the
parts of the terminalia with paramere or mesomere.
For this reason, I selected a system of terminalia
nomenclature synthesized from Crampton (1944),
van Emden and Hennig (1956), and Tuxen (1956).
I think this system is more appropriate to the situa-
tion in Hydrellia than that of Crampton (1944),
Wirth (1948),orSteyskal (1957).
I have considered the terminal tergum a syntergum
of segments 9 and 10. Steyskal (1957) and previous
authors have called this tergum the epandrium.
There is little need for this name in Hydrellia because
the tergum is only occasionally important taxonom-
ically, and it seems to be only slightly involved in
copulation. The surstyli have fused to a variable
NUMBER 6 8 13
extent, forming a ventral flap partially covering the
phallus (often most of the basal half). Crampton
(1944) and Steyskal (1957) used the term surstyli,
while Dahl (1959) called the two structures together
the hypandrium. I have called the entire intromit-
tent structure the phallus, partly because I was
uncertain how much of it represented the true
aedeagus and partly because phallus was a con-
venient suffix (basiphallus and distiphallus). I
needed these terms because of the taxonomic impor-
tance of the common division of the phallus into a
somewhat bulbous, heavily sclerotized basal part
(basiphallus) and a usually narrower, lightly sclero-
tized distal part (distiphallus). The basiphallus may
be the actual phallobase and the distiphallus?the
aedeagus of Snodgrass (1935, 1959). Phase micro-
scopy confirms the presence of a duct (presumably
the ejaculatory duct) passing through the length of
the distiphallus.
The phallus is suspended from the posterior ends
of a structure I have called the gonal arch (after
Tuxen, 1956, and Phillip Clausen, of the University
of Minnesota, in litt., 1964). Crampton (1944) and
Steyskal (1957) did not discuss or illustrate any
comparable structure. Tuxen (1956) illustrated a
structure in Aedes called the proctiger, which is
somewhat similar to the gonal arch. The anterior
ends of the gonal arch articulate with or are con-
tinuous with a median phallapodeme, or aedeagal
apodeme, and with sternum 5. Projecting from the
gonal arch at sternum 5 are two pairs of sclerites or
lobes called gonites (after Tuxen, 1956). The larger
pair are considered the postgonites. The smaller,
often inconspicuous pair with setose, bifurcated tips
are considered the pregonites despite their usual pos-
terior termination. In many species, the pregonites
appear to have a more definite articulation with the
phallapodeme. I have called the more heavily sclero-
tized, clawlike end of the postgonite the postgonite
uncus. This specific designation was needed because
of its taxonomic significance.
Sternum 5 varies specifically in shape. Usually,
the general impression is a horseshoe shape. The
posterior lobes are so significant taxonomically that
I have designated them copulobi after Crampton
(1944).
Although almost certainly additional unillustrated
muscles function in phallic movement, I have anal-
yzed the process in the following manner. The
phallapodeme and the gonal arch are pulled anteriad
by contraction of phallic depressors inserted at their
anterior ends and originating on or near sternum 4
(Figure 12). This contraction and phallapodeme
movement has two effects: (1) it lowers, or de-
presses, the phallus as a result of leverage between
the superior condyle of the basiphallic socket and
the posterior end of the phallapodeme; and (2) it
lowers the postgonites, which in most cases are
attached or closely appressed to the anterior end of
the phallapodeme. (I surmise that the copulobi are
simultaneously depressed by the same or associated
muscles. When depressed, the bilobate sternum 5
would fit the female postabdomen just anterior to
the cerci like a saddle). The postgonites either
titillate some female terminalia or hold the cerci erect
to permit coitus. This hypothesis is supported by the
fact that in several species the distiphallus in repose
projects anteriorly over the free posterior margin of
sternum 5, and by the angle, or attitude, of the
mounted male in several species. Phallic elevation
is effected by the phallic levators levering the pos-
terior end of the phallapodeme against the inferior
condyle of the basiphallic socket and by the phallic
depressors and associated muscles relaxing synchron-
ously. Once elevated into the genital pouch, or
cubiculum (after Crampton, 1944), the phallus is
apparently retained by the gonites, sternum 5, or by
both of these structures.
I have illustrated parts of the female terminalia of
a few species in Figures 2, 67-72. Grigarick (1959)
accurately illustrated the female abdomen in ventro-
lateral view. Dahl (1959) illustrated only a portion
of the female abdomen of H. griseola. Wirth (1948)
and Tuxen (1956) referred to the cerci as anal lobes
and valvulae mediales respectively. This is under-
standable, since in female Hydrellia and many other
Diptera segments 9 and 10 have been obliterated so
that tergum and sternum 8 are immediately anterior
to the cerci. Sternum 8 is often called the subgenital,
or pregenital, plate.
Internal Morphology of Adult
GUT.?I have illustrated most of the gut of H.
griseola in Figures 1, 3. These illustrations seem also
to apply fairly well for H. tibialis, H. bilobifera,
H. harti, and H. columbata. In gross anatomy, the
foregut, or stomodeum, is a uniform tube from the
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
dorsal cornua of the paraclypeal phragmata to the
vicinity of the cardia near the level of thoracic
phragma 2. Here there is a diverticulum from the
esophagus, the crop meatus, leading to the crop
proper. Distended, the crop occupies nearly the ven-
tral half of the abdominal cavity. The midgut starts
with the cardia, which contains the stomodeal valve.
From this structure to a level about midlength of the
crop, the midgut is a linear tube with the labial
gland ducts and labial, or salivary, glands parallel
and contiguous. At about midlength of the crop the
midgut is bent sharply dorsad and coiled three or
four times. The pyloric valve and most of the ileum
are similarly coiled. The site of evagination of the
Malpighian tubules is obscured by the gut coils.
In H. griseola and H. tibialis the Malpighian tubules
branch from two stems, or bases, arising near the
pyloric valve. Two tubules run anteriad and two
posteriad, both pairs parallel and close to the gut.
The rectal valve and rectum with its rectal glands
are the conspicuous parts of the hindgut, or proc-
todeum. The anus is located between the cerci.
The inner lines apparently representing the intima
of the midgut and hindgut in Figure 3 may actually
represent the peritrophic membrane.
INTERNAL GENITALIA.?The internal genitalia of
H. bilobifera illustrated in Figures 2, 14 are similar
in outline to those of H. griseola, H. tibialis, and
H. columbata. The internal male genitalia consist
of testes, vasa deferentia, pouchlike enlargements of
the vasa deferentia called seminal vesicles, a median,
unpaired ejaculatory duct, and accessory glands.
The testis and the initial portion of the vas deferens
are suspended and covered by a thin peritoneal
sheath. Although this sheath partially obscured the
testicular components, I observed a germarium of
spermatogonia in the apex anterior to the compactly
coiled spermatic tubule. Snodgrass (1959, p. 78)
stated that "Each testis, however, appears in its
entirety to be a single testicular tube. The same is
true of the testes in other Diptera." In the posterior
end of the testis and in the seminal vesicle I saw
what appeared to be packets of spermatids or
spermatozoa.
The accessory gland appeared dense and had a
reniform contour. The accessory gland ducts joined
the ejaculatory duct where it is convoluted anteriad
between the testes. Posteriorly the ejaculatory duct
is looped over the rectum as a result of pupal circum-
version. Between the rectum and the phallapodeme,
the ejaculatory duct is noticeably dilated for some
distance before it passes into the posterodorsal region
of the basiphallus. I located the apparent gonopore
but, as Snodgrass (1935) inferred, the gonoduct
probably opens into an endophallus, the external
opening of which is the apical phallotreme instead
of the true gonopore.
The internal female genitalia (Figures 2, 133)
consist of ovaries, lateral oviducts, common oviduct,
genital chamber, median spermatheca, two lateral
spermathecae, and two accessory glands. Each ovary
is composed of several polytrophic ovarioles. Each of
these consists of a short terminal filament, a long egg
tube, and a very short pedicel connecting with the
lateral oviduct. Histologically, the egg tube is com-
posed of a very short germarium of oogonia and a
long vitellarium of follicles. Each immature follicle
contains cystocytes, or follicle cells, trophocytes, and
an oocyte. The basal follicle contains a large oocyte,
complete with yolk and chorion. Covering each
ovariole is an epithelial sheath which seems to be
continuous with the terminal filament. The expanded
receptacular area of fusion of pedicels and lateral
oviduct is the calyx. I found a peritoneal sheath
over the ovary as shown by Snodgrass (1935) in
Rhagoletis pomonella and by Miller (1950) in
Drosophila. It is very delicate and is apparently the
ovarian suspensorium, or suspensory ligament.
Sturtevant (1925, 1926) first illustrated and de-
scribed the spermathecae and accessory glands of
Hydrellia in his comparative morphological study of
these structures in Aschiza and Schizophora Acalyp-
tratae. To the lateral spermathecae, which arise
dorsally from the junction of common oviduct and
genital chamber, Sturtevant was inclined to ascribe
a glandular function. He illustrated the accessory
glands, or parovaria, as arising just posterior to the
lateral spermathecae and showed them subequal to
the lateral spermathecae in H. griseola. Sturtevant
(1926), unlike Snodgrass (1935) and Imms (1957),
distinguished the median spermatheca as basically
different from the lateral spermathecae and called it
the ventral receptacle.
Regarding the function of the ventral receptacle,
Sturtevant (1926, p. 11) stated: "Sperm have been
found in this organ [median spermatheca] in
Dimecoenia, Discocerina, Hydrellia, llythea, and
Philygria. In no case in this family [Ephydridae]
NUMBER 6 8 15
have any sperm been found in any other part of the
female reproductive system." He did not ascribe a
specific function to the accessory glands, but I sur-
mise that they or the lateral spermathecae secrete a
lubricant and adhesive for the egg as it passes from
the genital chamber through the vulva. The bases
for this supposition are several observations of eggs
protruding from the vulva ventral to the cerci and
the fact that eggs are cemented to the oviposition
substrate.
Morphology of Immatures
EGO.?Hydrellia eggs exhibit some variability in
shape, condition of the chorion, and in size. The
size range of known eggs is 0.40 by 0.12 to 0.71 by
0.22 mm. In dorsal view the eggs are usually sub-
fusiform, or cucumiform, with fairly symmetric con-
tours except on the ends (Figures 75, 77, 78). In
lateral view, their shape is often boatlike, with the
ends upcurved almost symmetrically (Figures 73, 74,
125). The micropylar end is often somewhat more
acute than the opposite end. In all known eggs of
Hydrellia the chorion is corrugated or rugulose, with
alternate longitudinal ridges and channels. It is not
known if these ridges, or cristulae, have an adaptive
value or are simply incidental follicular impressions.
It is possible that the ridges serve as egg guides to
maintain some necessary orientation of the egg in
the common oviduct and genital chamber or that
they help maintain chorion shape. The identification
of the micropylar protuberance is presumptive, since
Berg (1950), Kato (1955), and Grigarick (1959)
offered no conclusive proof such as direct observa-
tion of sperm entry or sperm tracing.
The nature of the globose, lacunose, pluglike struc-
ture on the end opposite the micropylar protuberance
is still unknown. Whether the chorion is unilaminate
or multilaminate is also unknown.
LARVA.?Basically, the three mobile larval instars
differ little morphologically. The most conspicuous
developmental changes occur in the feeding appara-
tus and the spiracular peritreme. Von Frauenfeld
(1866), Stein (1867), Gercke (1879, 1882, 1889),
Brocher (1910), Keilin (1915, 1944), Hennig (1943,
1952), Berg (1950), Kato (1955), Nye (1958), and
Grigarick (1959) contributed to the description and
analysis of these changes.
The first instar appears setulose and warty and is
about 0.35-0.75 by 0.10-0.15 mm when newly
eclosed. It changes rapidly so that at the end of the
stadium it is similar to depictions in Figures 124
and 127 and measures about 1.00-2.25 by 0.18-0.25
mm. I have illustrated the changes in the feeding
apparatus in Figures 93, 103, and 105 for the three
mobile larval instars of H. spinicornis Cresson. The
conspicuous differences are in size, color, and clypeal
arch contour. In Figures 81, 84, and 85 I have illus-
trated the morphological nomenclature of the feed-
ing apparatus, or the so-called cephalopharyngeal
skeleton. I collated the nomenclature of the feeding
apparatus from Berg (1950), Hollande et al. (1951),
Frick (1952), Snodgrass (1953), Downes (1955), and
Allen (1957a, 1957b). From paired symmetrical parts
as shown by Berg (1950) in Notiphila loewi Cresson,
the feeding apparatus has evolved in Hydrellia to
the single, partially fused state. The two mouth-
hooks have become entirely fused, the posterior or
proximal ends of the H-shaped sclerite (called
hypostomal sclerite by Berg, 1950) and the anterior
ends of the paraclypeal phragmata have become
fused, and the cross piece lost in all species studied.
The paraclypeal phragmata are closely apposed and
are united dorsally by the frontoclypeal plate
anteriad to the bottom of the clypeal arch (area of
inclination near the usually distinct cheliform spot).
Ventrally, they are united by the sitophore, or
hypopharyngeal plate.
Each paraclypeal phragma is bifurcated pos-
teriorly into dorsal and ventral phragmatal rami
between which the cibarial dilator muscles can be
seen stretching from the dorsal rami to the epipharyn-
geal clypeal wall (dorsal cibarial wall). Anterior to
each phragmatal bifurcation is a protuberance,
which, because of its shape, Berg (1950) termed the
cheliform spot. From my dissections, it is evidently a
process for muscle insertion. Supported by unillus-
trated observations of feeding mechanics and some
dissections, I hypothesize that the fossae of the cheli-
form spots serve as insertions for protractor muscles
of the feeding apparatus, as does the prominent
clypeal arch, or dorsal angle of Allen (1957a), illus-
trated by Berg (1950) in Notiphila, Hollande et al.
(1951), Frick (1952), Snodgrass (1953), and Downes
(1955). Retractor muscles insert on the rami of the
paraclypeal phragmata and rotators on the phrag-
mata below the cheliform spots.
The primary factor underlying the reduction of
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
the clypeal arch and probably the entire stream-
lining of the feeding apparatus in Hydrellia has
been selective pressure for manipulative flexibility
and for an anterior end capable of extreme flattening
during feeding. The selecting factor, or at least the
factor compatible with this anatomical modification,
has been leaf mining?feeding internally in very thin
submergent leaves such as in most Potamogeton.
Metapneustism and perhaps some kind of cuticular
respiration opened the way to obtaining oxygen from
the aerenchyma of certain plants, and this in turn
provided the opportunity to move deeper into plant
tissues. But before this opportunity could be utilized,
manipulative flexibility of the feeding mechanism
had to be increased greatly to allow for feeding in
very tight places. Much of the increase in rotatability
of "head" and feeding apparatus has been due to the
reductive fusion of the two mouth-hooks and of the
sclerites of the paraclypeal phragmata and to the
overall compression of the feeding mechanism. All
of these modifications have enabled the larva to
rotate the head lobe and feeding apparatus 90 de-
grees either way from the vertical attitude.
Anteroventrally on the feeding mechanism are
two projections which Berg (1950) called ventral
projections. Hollande et al. (1951), Frick (1952),
Snodgrass (1953), and Allen (1957a) showed the
larger, posterior one of these to be the aperture of
the common labial gland (salivary) duct. After
several dissections, I could concur with the findings
of these authors. I have called the anterior projection
the labial sclerite, although there is some reason to
think it is a remnant of the H-shaped sclerite and is
involved with the mouth-hook depressor apodeme in
depression of the mouth-hook.
It became obvious after making several dissections
that the labrum, contrary to Snodgrass's (1953)
illustration, is noticeably anterior to the hase of the
mouth-hook. This is obvious from Figures 81 and
84, where the fusion of sclerites can be seen to elimi-
nate any space for the labrum in the position shown
by Snodgrass. Nye (1958) showed the labrum as
bilobate, with a lobe on each side and above the beak
of the mouth-hook in H. incana.
The tracheospiracular system of Hydrellia is meta-
pneustic, but Keilin (1915, 1944) demonstrated a
vestigial, closed prothoracic spiracular atrium. Ac-
cording to Hennig (1943, 1952) there appears to
be a trend toward functional metapneustism in the
Ephydridae. Developmentally, the caudal tracheo-
spiracular siphon changes considerably between the
first two larval instars of several species at least. I
have illustrated (Figure 80) a spiracular condition
which may be representative of the first-instar larvae
of many species of Hydrellia. The peritreme is con-
ical with a terminal aciculous spine, the spiracular
atrium narrow, and the secondary atrial orifice medi-
ally located. The spiracular peritreme is probably
nonretractile. On abdominal segment 8 are supra-
spiracular and subspiracular protuberances: the
supraspiracular protuberance has one long spinous
seta, and the subspiracular one several shorter setae.
I did not discover the function of these setae and
spines, but they may possibly aid in locomotion and
anchorage.
Although Figure 79 is of a different species than
Figure 80, it is representative of the change after the
first molt. The conical peritreme is larger and less
aciculate, and the secondary atrial orifice is dorsally
situated. The spiracular atrium and the primary
tracheal orifice at the junction of atrium and the
main longitudinal trachea are discernible (Figures
79, 83, 127). In species where supraspiracular pro-
tuberances and conspicuous spinous processes occur
in the second-instar larvae, these are lost at the
second molt.
According to Hennig (1943, 1952), the third-
instar spiracular peritreme has three secondary atrial
orifices; however, since he did not make an extensive
survey, this may not be a generic character. Grigarick
(1959) illustrated one secondary atrial orifice per
peritreme in the second instar as in Figures 79, 83.
Keilin (1915) did not specify the instar but he illus-
trated a single, dorsal secondary atrial orifice in each
peritreme in H. modesta Loew. Kato (1955) also
showed a single, dorsal secondary orifice.
In the third-instar larvae, the increased promi-
nence of the creeping welts makes it possible to
discern the eleven apparent segments and a head
lobe. I have shown the head lobe bearing apparently
three-segmented antennae (Figure 81). Immediately
posterior to the head lobe, the dorsal and ventral
head folds of Snodgrass (1953) can be seen. I have
followed Hollande et al. (1951) in calling the setu-
lose ventral fold the postlabial pad. Berg (1950)
called it the postoral tuft. Although Nye (1958)
gave it no specific designation, he showed i* as the
anteromedial part of the prothoracic venter, as did
NUMBER 6 8 17
Hollande et al. (1951) in a nonephydrid species.
Nye considered the postlabial pad a constituent of
the facial mask, along with the pair of two-segmented
antennae, anterior cephalic papillae, a pair of frontal
papillae, maxillary palps, and two labral lobes.
Also Nye definitely showed two-segmented antennae,
whereas Berg (1950) and Kato (1955) illustrated
three-segmented antennae. Grigarick (1959, p. 10)
stated, "Antennae (Fig. 11) two-segmented, set on
a small enlargement which may be a third seg-
ment . . . ." Obviously, the question will be difficult
to resolve even with dissection because of the small
size of the structure.
Keilin's (1944) concept that all dipterous larvae
have three thoracic and eight apparent abdominal
segments can be applied to Hydrellia, but with some
difficulty, for all known larvae exhibit a postanal
extension of abdominal segment 8 in the third instar
and puparium, which appears to be a ninth segment.
According to Nye (1958), tracheation and innerva-
tion prove it is not a true segment.
As inferred by Hinton (1955), the creeping welts
are homologous with prolegs. These creeping welts
and specifically variable portions of the segments are
covered by microcuticular processes called spinules
by Berg (1950) and Hollande et al. (1951), loco-
motory spinules by Hinton (1955), tubercles by Allen
(1957b), and spiculi by Nye (1958) and Grigarick
(1959). Because the term is more descriptively flex-
ible than either tubercles or spiculi, I have called
these microcuticular processes spinules. They are
seldom truly needle-shaped, or spiculiform, nor are
they often truly tuberculiform in Hydrellia. Accord-
ing to Hinton (1955), the large locomotory spinules
are similar in origin, structure, and function to the
crochets of most dipterous prolegs.
Allen (1957b) and Nye (1958) placed consider-
able significance on tubercular (or spicular) zones
and the number of processes and rows in each. But
as Berg (1950) illustrated, and as I have shown in
Figures 107-123, in the puparia the shape and size
of creeping-welt spinular zone in relation to the
lateral spinular pattern is the more readily usable
taxonomic character in Hydrellia. Anterior to each
creeping welt, Berg (1950), Grigarick (1959), and
I found a transverse row of six setulae. Keilin (1915)
and Nye (1958) showed only four setulae in this
location in H. incana. Both of the latter also showed
no cheliform spot on the feeding apparatus. Perhaps
it was rudimentary or so uniformly sclerotized as to
be indistinguishable. The spinulosity and setulosity
of the third-instar larva often are less distinct than
in the puparium because the cuticle of the former is
more translucent than that of the latter. The trans-
lucency is such that the living larva in situ appears
green or yellow, depending upon the combined effect
of labial gland color and aliment color. In some
species, the fat body appears to retain some plant
pigments.
PUPARIUM.?Although it is actually part of the
third instar, the puparium is a distinct morpholog-
ical phase. The changes involved in the formation
of this uniquely economic "cocoon" are physiolog-
ically complex. Keilin (1944) used the term pupario-
genesis to describe this formation. An equally satis-
factory term and one more readily usable verbally
is pupariation. In Hydrellia, the third-instar larva
becomes quiescent and opaque just as at the first
two molts, and within a few hours the cuticle has
contracted lengthwise, expanded transversely, and
hardened. For two species I found dimensional
changes from third-instar larva to puparium to be
from 5.53 by 0.60 mm to 4.00 by 1.06 mm and 6.50
by 1.22 mm to 4.75 by 1.60 mm. The puparial shape
varies specifically from subcylindrical to fusiform.
Some are noticeably attenuated posteriad and the
puparia of four species have almost symmetrical ends
in ventral view. In lateral view, the puparia are
cyphosomatic.
Conspicuous structures other than the creeping
welts are the head-lobe scar and the anal plate (both
ventral) and the dorsocephalothoracic cap (Figure
107). A better name for this latter structure is oper-
culum, for it is simply the dorsal thoracic section of
the puparium delimited by a continuous ecdysial
cleavage line. There is nothing cephalic about it.
Regarding the head-lobe scar, some authors such as
Brocher (1910) called it the mouth scar, but
TragSrdh (1903, p. 33), in discussing pupariation in
Ephydra riparia Fallen, stated: "Das Verschliessen
der vorderen Offnung erfolgt in der Weise dass die
Larve den Kopfabschnitt so tief hineinzieht, dass
eine tiefe trichterformige Einstiilpung der ventralen
Seite des Prothoracalsegmentes gebildet wird. Dieser
Trichter zieht sich im proximalen Teil zusammen
und erhartet stark (Fig. 6, Taf. 1) zu einem schwar-
zen, in das Puparium hineinragenden zapfenformigen
Gilde." This complete retraction of the head lobe
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
upon pupariation seems to occur also in Hydrellia;
thus, I have referred to the anterior end of the
puparium as prothoracic.
According to Wigglesworth (1956), in third-instar
larvae of higher Diptera the epidermis initiates
sclerotization of the soft endocuticle, making it
chemically identical to the exocuticle in the pupa-
rium, except in aquatic larvae where lime deposition
is substituted for this endocuticular sclerotization.
Because of their endophytic environment and per-
haps other factors, Hydrellia puparia do not exhibit
any apparent endocuticular lime deposition (Figures
126, 128, 131, 132). The translucency of the puparia
of some species, e.g., those in Figure 131, seems to
make the term "tanning" seem somewhat inappro-
priate. The puparial cuticle is sclerotized, but dark-
ening seems to vary ecologically and perhaps genet-
ically. Hering (1950) observed that the prevalent
translucency of Hydrellia puparia makes them some
of the most suitable of all Acalyptratae for inves-
tigating pupation.
Ecology
Ecological Role and Distribution
Adult Hydrellia are polyphagous, but since the
larvae are endophytophagous and since they consume
much more food than the adults the species can be
classified trophically as primary consumers. It is
probable that this classification will be found to
apply to all Notiphilinae.
No species of Hydrellia has been studied suffi-
ciently to integrate it completely in a food web, but
food chains involving some species are known. Larvae
of H. griseola parasitize hydrophytes of many species,
primarily grasses. Adults of this fly prey upon very
small insects and upon insects trapped in the surface
film. Among these are specimens of Hydrellia (in-
cluding Hydrellia griseola), Psyllidae, early-instar
Ephemeroptera and Odonata, Collembola, Braconi-
dae, several species of nematocerous Diptera, Musca
autumnalis, and Sarcophaga sp. Adults of some spe-
cies prey actively on Collembola. Yeasts and similar
fungi, Cyanophyta, Chrysophyta, nectars, and leaf
epidermis are other dietary items of adults. Sever-
al species of lycosid and ctenid spiders, species of
Ochthera (Ephydridae), and species of Lispe and
Hydromyza (Muscidae) prey upon adults of Hydrel-
lia. The chain continues through fishes, e.g., bluegills
and sunfishes, to ichthyophagous reptiles, birds, and
mammals. A side chain exists for the larvae. Several
hymenopterous species parasitize the larvae. From
these parasites, the energy flows through most of the
same species as it does from the adult flies. Similar
food chains exist for H. cruralis and H. pulla.
Host-plant preferences of the larvae likely influ-
ence the ecological distribution of Hydrellia species.
Predominantly, these preferences center on species
of Potamogetonaceae and Gramineae. Other prefer-
ences include species of Alismataceae, Cyperaceae,
Hydrocharitaceae, Lemnaceae, and Liliaceae. Mines
of Hydrellia in dicotyledonous plants of Caryophyl-
laceae, Labiatae, Chenopodiaceae, Scrophulariaceae,
and Compositae apparently serve only for puparia-
tion, and as such they are relatively short and simple.
Apparently, larvae that leave their old mines just
before pupariating have little host preference. I
found H. ischiaca puparia in the floating leaves of
Potamogeton natans growing near Zizania aquatica,
the preferred host plant of the locality, and H. bergi
puparia in leaves of Z. aquatica growing near a
preferred species, Potamogeton richardsonii. Nastur-
tium officinale (Cruciferae) is the one established
exception to the general absence of feeding mines
in dicotyledons. European authors reported certain
Hydrellia species parasitizing N. officinale, and I
have confirmed H. griseola as a fairly common miner
feeding in plants of this species in some localities in
the United States. The known host plants, including
those serving only for pupariation, are listed in
Table 1.
Hydrellia play an important role in many aquatic
ecosystems, especially eutrophic ones. Lange et al.
(1953) estimated that H. griseola destroyed from 10
to 20 percent of the California rice crop in 1959. At
Lake Itasca, Minnesota, I found that the infestation
of several stands of wild rice, Z. aquatica, by H.
ischiaca ranged from 33 to 89 percent in 1963. No
one has measured the effects of the mining of many
Hydrellia species in several Potamogeton species,
but they are obviously considerable judging from
observed leaf damage in many stands at several
localities. The same can be said of many other host
plants. Hydrellia larvae function, together with some
Aphididae, Delphacidae, donaciine Chrysomelidae,
Trichoptera, Lepidoptera (especially Nymphula
spp.), Chironomidae (Cricotopus, Polypedilum,
NUMBER 6 8 19
Glyptotendipes, and Tanytarsus spp.), Hydromyza
confluens, and numerous other phytophagous insects
in producing subtle, and sometimes conspicuous,
changes in the littoral macroflora. Temperature and
humidity requirements of adult and immature
Hydrellia as well as requirements of host plants
influence the effect and extent of mining. Wind and
wave action and temperature and humidity tolerance
tend to restrict oviposition to sheltered, eutrophic
habitats. In such habitats, low mineral availability,
low temperatures, wind, extensive algal and Lemna
mats, and excessive water depth make some species
of plants more susceptible to heavy infestation. The
newest plant growth is generally most susceptible to
infestation and is most severely damaged.
Berg (1949) found 26 insect species in addition to
those of Hydrellia living upon or in 17 species of
Potamogeton. Most of these insects may not have to
compete extensively with one another because of the
abundance of the plants and of narrowly defined
ecological roles. There is obviously interspecific com-
petition with aquatic caterpillars, snails, phytopha-
gous fishes, waterfowl, muskrats, beavers, deer, and
moose. According to Fassett (1960), bluegills eat the
leaves of several Potamogeton species and Vallisneria
americana; trout eat parts of Nasturtium officinale
and Zannichellia palustris; waterfowl and muskrats
use several Potamogeton species and Sagittaria
latifolia tubers as a staple; waterfowl, muskrats, deer,
and moose feed extensively on Zizania aquatica;
and ducks use Echinochloa species as a staple.
Parasitological Data
Burghele (1959a) stated: "It is a well established
fact that all aquatic Hymenoptera are parasites, the
female laying her eggs in the eggs or larvae of
aquatic insects." This cannot be entirely true, for I
observed a specimen of Chorebus aquaticus oviposit-
ing repeatedly for 5 minutes in a Z. aquatica leaf
harboring neither eggs nor larvae of Hydrellia. Also,
I watched a submerged specimen of Trichopria
columbiana oviposit in leaf tissue around an empty
puparium of H. pulla. One of my observations, in
which Chaenusa sp. parasitized three or four puparia
of H. ischiaca reared from eggs, indicates that some
eggs or larvae of parasites may enter the host larva
through the gut, for adult hymenopterans had no
access to these larvae. Grigarick (1959) believed
Pnigalio sp., Sympiesis sp., and Solenotus inter-
medius to be external larval feeders because they
pupated free of the host. He discovered abundant
eggs of S. intermedius and many larvae in the mines
of H. griseola. Some larvae fed externally on H.
griseola larvae.
Parasitic Hymenoptera undoubtedly exert consid-
erable control on population densities of Hydrellia,
especially in certain marginal habitats and when
population densities are very high. Grigarick found
parasitism of H. griseola by external parasites re-
peatedly higher in pools that were drying or very
low. In one sample the parasitism was 60 percent.
He found hymenopterous parasitism low on the first
host generation mining rice but rising rapidly on
succeeding generations to nearly 90 percent by July.
Opius hydrelliae and Chorebus aquaticus (Braconi-
dae) were most abundant. After July, parasitism
declined and remained low for the rest of the year.
This seemed to be correlated both with a great
decrease in host density and with less vulnerability
of Hydrellia larvae to parasitism when in certain
restricted survival habitats, e.g., fall rain pools. I
found 38 percent parasitism in 132 puparia of H.
ischiaca and 63 percent in 61 puparia of H. pulla
collected through one summer.
Hormone concentration and balance and perhaps
other physiological factors may stimulate develop-
ment of endoparasitic larvae, for in several observa-
tions they began to feed actively on host tissues only
after host pupariation. I have photographed an
opiine larva situated in the host larva (Figure 124).
After one host pupariated, I saw the leechlike larva
of Ademon niger devour the host tissues in a matter
of hours. When the parasite pupates, it usually
orients its head toward the prothoracic end of the
puparium (as in Figure 131). At emergence, it
usually leaves a dark meconium in its pupal exuviae
located in the puparium (as in Figure 128). I
observed several escape exits cut anteroventrally in
the puparium (as in Figure 132) and a few postero-
ventrally. I have observed only one case in which
the hymenopteron used the puparial operculum and
did not cut its own exit. According to Burghele
(1959a), the Dacnusinae cut the escape exit with
their mandibles, while the Chalcidoidea make the
exit by shaving off tiny pieces of the puparium.
Inflow of water would not necessarily kill the emerg-
ing adult hymenopteron for Ademon niger and
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Trichopria columbiana, at least, can remain sub-
merged for several hours. I noted four adults of the
latter species to live submerged for 24 hours. Two of
these had already emerged and then submerged
again.
The known hymenopterous parasites of Hydrellia
are listed in Table 2. In addition to these parasites,
species of Stigmatomyces (Laboulbeniales) parasitize
Hydrellia adults and larvae. This fungus apparently
killed several larvae of H. bilobifera during labora-
tory rearings.
Dispersal and Zoogeography
Most adults of Hydrellia locomote by a combination
of walking and short, hopping flight. Flight through
a distance of a few meters occurs in a zigzag pattern.
Of all the species, H. griseola most probably has the
longest flight range because it has the widest dis-
tribution and the largest wings. Perhaps its large
wings and flight behavior often subject it to upward
wind currents, for it is the most common species of
Hydrellia captured in the few aerial surveys per-
formed thus far in North America. Dispersal via
passive dissemination by high-altitude atmospheric
currents is probably efficient in most cases only over
a few hundred miles because of body water loss.
Strong wind often causes adults to congregate at the
downwind end of lakes and pools. Some dispersal in
most species probably occurs over relatively short
distances in or on plants caught in water currents.
Immature instars are particularly susceptible to this
passive dissemination. The phenomenon probably
accounts for the presence of larvae in plants at
depths of from 3.0 to 6.0 meters in some lakes.
Wind action and currents may carry floating puparia
for variable distances.
I have collated the following data for zoogeo-
graphic consideration: (1) the known host prefer-
ences center on plant species having their greatest
densities mostly between latitudes 25? and 65? north;
(2) H. griseola and probably many other species have
a temperature tolerance of 50?-90?F for adults and
larvae; (3) high temperatures lower the surface ten-
sion of water and thus probably adversely affect
locomotion of adults and migrating larvae; (4) most
of the known species are Holarctic, the greatest num-
ber being in the Nearctic Region; (5) H. griseola
has the most extensive distribution, being reported
from all zoogeographic regions except the Oriental
and Australian Regions; (6) H. griseola has the
greatest host-plant range within the genus.
Behavior
EMERGENCE.?The pharate adult must do more to
emerge than simply shed the pupal cuticle; it must
escape from the puparium and from the plant tissue
holding the puparium. Pharate adults start making
slight movements as early as twelve hours before
emergence. They use their ptilina to open the oper-
culum of the puparium. Initially, the operculum
remains attached posteriorly because the ecdysial
cleavage line extends only around the front and sides.
Once the fly gets its head through the opening it
pushes with its proboscis against the tip of the
puparium until its fore legs are free. After this, it
rapidly leaves the puparium. To emerge completely,
adults of most species have only to force their way
through thin, and often partially decomposed, plant
epidermis and then rise in an air bubble if the
puparium was submerged. However, adults of spe-
cies pupariating in stems and rhizomes sometimes
fail to escape and die. This can occur also in
H. griseola and other species mining grasses if the
leaves dry and become impliable. According to
Grigarick (1959), flies emerged several times during
the daylight and darkness at between 50? and 90?F.
After emergence, flies of at least several species walk
around slowly for several minutes, stopping period-
ically to clean their bodies and evert and then with-
draw the ptilinum. The wings remain folded until
about 15-30 minutes after emergence, when they
rapidly expand. Often, they cannot fly until 45 min-
utes after emergence.
FEEDING.?Adults often search over the surface
film for entrapped insects and sometimes mistakenly
pounce on inanimate floating objects and manipu-
late them with their fore tarsi as they do with small
insects. They often protect their catches by making
short rushes toward intruders. Adults have labellar
canalicular teeth, which apparently function carnas-
sially in lacerating conjunctival membranes of
trapped insects. According to Berg (1950), adult
H. cruralis chew small holes in floating leaves of
several Potamogeton species. They probably use their
labellar teeth for this chewing as well as for loosening
yeast, Chrysophyta, and other periphytic micro-
NUMBER 68 21
organisms. Adults of several species often congregate
within corollae of flowers of certain aquatic plants,
e.g., Nymphaea odorata, Nuphar advena, and
Ranunculus longirostris. Some species exhibit pecu-
liar behavior while feeding, e.g., H. biloxiae rhyth-
mically pushes its body up and down and H. ischiaca
sometimes touches its food-insects with its abdomen.
Hydrellia biloxiae and H. pulla protect their catches
of dead or dying arthropods by partially extending
their wings and making short rushes to drive away
intruders.
MATING.?Dahl (1959) distinguished six phases
in the mating process: (1) initiation, in which the
male approaches the female from the front or side;
(2) posturing, in which the male scissors his wings,
curves his abdomen, or assumes some other distinc-
tive posture, often while moving back and forth
before the female; (3) restimulation, in which the
male repeats his posturing to secure the female's full
attention; (4) mounting, in which the male climbs
on the female and grasps her partially extended
wings; (5) insemination, in which the male inserts
his phallus and ejaculates, often while titillating the
female's abdomen with his hind legs; (6) dismount-
ing, in which the male disengages and the female
pushes at him with her hind legs. Although I have
observed all of these phases in the mating of Hydrel-
lia, I found the terms epigamic and gamic more
flexible in describing mating behavior because of
variations in different species. One pair of H. nobilis
copulated without exhibiting any apparent epigamic
behavior. Another male H. nobilis repeatedly climbed
on this copulating pair and exerted his phallus each
time. Epigamic behavior in many species consists of
wing scissoring and walking to and fro before the
female, but males of H. definita rhythmically push
their bodies up and down, and H. bergi, H. surata,
and H. columbata touch their faces or antennae
before mounting. The mechanisms of species and sex
recognition remain obscure, but surfaces reflecting
ultraviolet rays may constitute at least one mechan-
ism. Highly reflective objects such as automobiles,
clean water surfaces, and polished aluminum pans
attract several insect species, especially Diptera and
Hymenoptera. One could hypothesize that the pre-
dominant silvery or yellow facial pruinosity in
Hydrellia (and other schizophorans) reflect consid-
erable ultraviolet radiation. Nickel and silver reflect
about 40 percent of radiation with wavelengths of
251 millimicra, and the percentage reflected should
be greater in the normal ultraviolet band of 292-400
millimicra. Density patterns of the pruinosity may be
important if this hypothesis is valid. Several authors,
e.g., Milne and Milne (1959), have emphasized the
importance of ultraviolet radiation in insect optics.
Mechanisms of sex and species recognition some-
times fail, as in cases where a male H. nobilis tried
to mount a male H. ischiaca after repeatedly scissor-
ing his wings, or a male H. nobilis tried to mount a
female H. ischiaca, or a female H. amnicola at-
tempted to mount another female of the same
species. Some cases of agonistic behavior are difficult
to distinguish from those involving failure in sex or
species recognition. In H. bergi, small males scissored
before large males in two instances, and the large
males grasped them by the thorax as if to mount.
However, most observed agonistic behavior was
interspecific, especially between Hydrellia species and
predators such as spiders, Lispe spp., and Ochthera
mantis. Adults of several Hydrellia species made
short rushes toward these predators, often with their
wings partially extended.
Copulation time varies specifically and with sev-
eral other conditions. A pair of H. ischiaca copulated
for 10 minutes and a pair of H. nobilis for 19 min-
utes. Hydrellia griseola have copulated from 1 to 50
minutes and at various times during daylight be-
tween 55? and 90?F. Copulation between the same
pair may occur repeatedly in one day and, in H.
griseola, for five consecutive days (according to
Grigarick, 1959). However, continuous mating is
unnecessary, for a female H. griseola has laid viable
eggs 93 days after being isolated. Grigarick (1959)
found the shortest time between emergence and
copulation for H. griseola of the same age was three
days, but Berg (1950) reported that H. cruralis
copulated 24 hours after they emerged. In the lab-
oratory, pairs of H. griseola have copulated as late
as 70 days after emergence (Grigarick, 1959). Pairs
may copulate in various types of microhabitats, but
most frequently on pleustonic vegetation. One pair
of H. cruralis copulated while skimming over the
surface film, and (according to Grigarick) pairs of
H. griseola copulated on emergent vegetation. In all
species observed, the female continues to walk and
occasionally to feed during copulation.
OVIPOSITION.?When forced by environmental
conditions, females oviposit on nearly any surface.
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
In the laboratory, they will oviposit on glass, and in
the field they will oviposit on dead stems, twigs of
shrubs, and several hydrophytes that are not their
hosts, e.g., Nelumbo lutea, Polygonum scabum,
Equisetum sp., and Sagittaria sp. Most Hydrellia
females can apparently distinguish preferred hosts
when they are available. Usually, they prefer to lay
their eggs in at least partially concealed places, such
as beneath folded leaf margins and on the adaxial
surfaces of sepals and stipules, but females of the
H. griseola group lay their eggs on the exposed leaf
blades of grasses close to the water surface. Most
species deposit eggs in irregular masses, but some,
notably H. spinicornis, scatter their eggs over the
leaf. It is not uncommon for females to lay eggs on
top of another's eggs. If females oviposit on linear
objects, e.g., leaf blades of grasses and various stems,
they align their eggs with the long axis of the object.
Hering (1951) reported that females of some
Hydrellia species submerge to oviposit. I have not
observed this nor have I seen any other report of it.
ECLOSION.?After about two to six days of incuba-
tion, the pharate larvae often begin to make slight
movements several hours before eclosion. They slit
the micropylar end of the chorion with their mouth-
hook and either crawl entirely out of the chorion or
only protrude their head lobe sufficiently to begin
excavating a mine. Sometimes the larvae remain in
the ruptured chorion for several hours. Mortality is
probably high among newly eclosed larvae judging
from oviposition sites quite removed from the host
plants and from the number of recently eclosed larvae
that fall into deeper water. Many larvae must eclose
while submerged because of water level fluctuations.
Those eclosing from eggs on the adaxial surface of
floating leaves usually avoid direct sunlight by crawl-
ing onto the abaxial surface. Larvae locomote on the
surface by anchoring their mouth-hook and con-
tracting the body segments and by pushing with
their spiracular peritremes and creeping welts. Par-
tial desiccation stimulates larvae, especially newly
eclosed ones, to twist and roll. This emergency re-
action often carries them into water, but once there
they may sink into an area devoid of plants.
MINING.?When the newly eclosed larvae find an
area they can penetrate because of either weakened
or thinner epidermis or some other condition, they
first break the epidermis by striking it perpendicu-
larly with their mouth-hook and then extend the
opening by rotating the head lobe and mouth-hook
and slicing the epidermis. The larvae ingest the
loosened tissues. The prothoracic end flattens down
to little more thickness than that of the feeding
apparatus to project the mouth-hook into the ex-
tremely thin mesophyll. By continuing this excava-
tion process, the first-instar larvae completely enter
their mines in 2 to 2.5 hours. Third-instar larvae
require only about half this time. The initial opening
does not have to be as broad as the larva because
of segmental contraction. Inside the mine, larvae
move forward by peristalsis, in which contraction
waves pass anteriad through the creeping welts,
and occasionally by pushing with the spiracular
peritremes and pulling with the mouth-hook. They
can move slowly backward by reversing the peri-
stalsis. Also, even late third-instar larvae can turn
about within the mine. During endophyllous loco-
motion, the spinules of the dorsum and creeping
welts press against the upper and lower epidermis
and sometimes penetrate it.
Mining larvae apparently depend on the host
plant for oxygen. According to Hering (1951), frass
and other debris seal the mine entrance and prevent
the entrance of water. Air then enters through the
plant tissue and accumulates in the mine around the
larva. This does not always happen, for I found
larvae in old mines partially filled with either water
or tissue fluid, and I observed larvae of H. griseola,
H. ischiaca, and H. bilobifera pierce several different
tissue areas in their mines with their spiracular
peritremes until they found an apparent oxygen
source. Larvae crawling on the underside of the
surface film or starting a mine just below the surface
film were observed periodically raising their spirac-
ular peritremes above the surface film. Observations
by Aldrich (1912) and Nemenz (1960) indicate a
strong possibility that some kind of plastron or
cuticular respiration operates in Hydrellia larvae.
Nemenz (I960, p. 222) showed in Ephydra that
"die Larven gelegentlich eine Luftblase aus den
Abdominalstigmen auspressen und wieder einziehen,
die offenbar als 'physiologische Kieme' wirkt."
Mines of first-instar larvae often are inconspicuous
because of the small larvae and also, perhaps, be-
cause of partial plant tissue repair. As the larva
grows, the mine is enlarged, but it may remain
inconspicuous if it is in the middle of a relatively
thick leaf, e.g., floating leaves of several Potamogeton
NUMBER 6 8 23
species. Feeding mines of first- and second-instar
larvae in narrow leaves such as those of grasses and
some narrow-leafed Potamogeton species are fairly
straight, but those of third-instar larvae are usually
tortuous and have blotch-like chambers. In broad
leaves, the feeding mines of even first-instar larvae
may be tortuous. Larvae eat the mesophyll and,
since various conditions will cause several larvae to
congregate in a single leaf, skeletonization often
results. If this occurs before the larvae are ready to
pupariate, they migrate to fresh leaves.
Some species of Hydrellia overwinter as larvae.
Larvae of H. bilobifera can survive in Potamogeton
nodosus encased in ice.
PUPARIATION.?Pupariation is a fairly new term for
the distinct process of the formation of a puparium.
It is not equivalent to pupation, since it is essentially
reformation of the third-instar cutide and since a
short fourth larval stadium occurs within the
puparium before pupation. This fourth instar is very
abortive, for a new feeding apparatus is not formed.
When larvae are ready to pupariate, they become
quiescent, with a greatly lowered heart rate. Puparia-
tion occurs within a period of eight hours in some
species. Larvae usually pupariate just under the
epidermis of thick leaves. This improves the emerging
adult's chances of escaping from the leaf. In thin
leaves, such as those of grasses, position of the pu-
parium seems unimportant. Third-instar larvae of
H. bergi and H. biloxiae make escape slits for the
adults in the stems and stolons that they mine. Third-
instar larvae of several species often migrate to a new
plant part for pupariation. This behavior probably
aids survival by providing a more stable anchorage
than the old, mined plant part. Also, this behavior
partly accounts for the broad range of host plants of
H. griseola. Apparently the larvae are nonspecific
for these pupariation plants. According to Berg
(1950), several species always pupariate with their
spiracular peritremes anchored in the midribs of
Potamogeton leaves. This fact seems to support the
hypothesis that puparia must take air from the plant,
but other data seem to refute this hypothesis. For
instance, larvae of H. ischiaca often pupariate in
skeletonized or wilted grass leaves, with their spi-
racular peritremes embedded only in the epider-
mis, and pupate successfully. Larvae of H. griseola
usually make a blotch mine and pupariate in its
center. Grigarick (1959) reported that H. griseola
larvae pupariated on glass, in sand, and in blotting
paper in the laboratory. Burghele (1959a) reported
finding many H. griseola puparia on the bottoms of
ice-covered pools. This may indicate overwintering
capacity of puparia. Schiitte (1921) reported the
existence of summer and winter puparia in Hydro-
myza livens (Muscidae). The winter puparium has
a wall eight times as thick as that of the summer one.
Such seasonal forms may exist in Hydrellia.
Environmental Tolerance
Hydrellia are fairly stenotopic and stenohygric. The
adult tolerance range for wind and temperature is
relatively small. Wind often precludes oviposition on
suitable host plants located in the limnetic zone of
ponds and lakes, and it often causes host plants to
break loose and pile up on the shore where the
mining larvae have no opportunity to escape. Wind
is probably one limitation to the height at which
oviposition can occur on emergent vegetation.
Grigarick (1959) presented the following tempera-
ture limits (in degrees Fahrenheit) to adult and
larval activities of H. griseola:
Adult
Heat death:
Heat paralysis:
Dropping:
Heat rest:
Normal locomotion:
Slow walking:
Heat death:
Heat paralysis:
Agitated movement:
Normal mining:
Quiescence:
110-114
107-110
104-107
98-104
52- 92
40- 50
Larva
112-114
104-111
95-103
50- 90
36- 50
In one of Grigarick's tests, 50 percent of a sample
of twelve wild adults lived 34 days at 29?F, and, in
another one, 20 percent of 80 eggs hatched after 120
hours of exposure to 29?F. The principal extrinsic
factors affecting the length of stadia of Hydrellia
are temperature, atmospheric saturation deficit, and
food (composition and quantity). The last factor
is also the principal extrinsic one affecting fecun-
dity.
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Host Plants and Parasitic Wasps
The known host plants and parasitic wasps of
Hydrellia are listed in tables starting on page 106.
Systematic Treatment
Classification
The type-species of Hydrellia Robineau-Desvoidy
(1830) is Hydrellia aurifacies Robineau-Desvoidy,
the first known valid designation of which was made
by Coquillett (1910b). Coquillett cited Notiphila
flaviceps Meigen (with H. aurifacies as a synonym)
as the type. According to Article 69a (iv) of the
International Code of Zoological Nomenclature
(1961), Coquillett effectively designated H. aurif-
acies as the type-species. According to Wirth (1965),
Westwood (1840) first designated H. aurifacies as
the type-species, but Westwood actually designated
H. flaviceps Meigen as the type-series. Since H.
flaviceps was not an originally included nominal
species and since Westwood did not synonymize it
with H. aurifacies, his designation is invalid.
Ever since 1896 (and perhaps before), Notiphila
griseola Fallen has generally been considered the
type-species of Hydrellia Robineau-Desvoidy (1830).
Macquart (1835) first reported the synonymy of
Hydrellia communis Robineau-Desvoidy (1830) and
Notiphila griseola Fallen (1813). Macquart also
transferred several other species from Notiphila to
Hydrellia. Kloet and Hincks (1945) rejected the
name Hydrellia as a homonym of Hydrelia Huebner
and replaced it with Hydropota Rondani. Article
56a of the International Code (1961) specifically
prohibits such rejection in this statement: "Even if
the difference between two genus-group names is
due to only one letter, these two names are not to be
considered homonyms."
The adults of Hydrellia can be distinguished from
those of other Ephydridae by the following char-
acters: ocular pubescence moderately dense; face
usually slightly convex (distinctly receding and
planate in one group) ; postocellar as long or longer
than ocellar; two or three dorsocentrals present, at
least one of which is as long or longer than longest
facial; median spermatheca cupuliform, heavily
sclerotized. Some additional characters present in
adults of most species are as follows: about three
palpal and five labellar setae; one very fine, ven-
trally directed, secondary facial setula inserted above
primary facial row; two fronto-orbitals (anteriorly
and posteriorly inclined) ; apicodorsal antennals not
projecting noticeably; three scutellars (basal, in-
termediate, and apical); one mesanepisternal,
mesokatepisternal, and basal coxal; ground color of
cuticle usually dark brown or black.
The larvae can be distinguished from those, of
other ephydrid genera by these characters: meta-
pneustic; spiracular peritreme spinous; mouth-hook
single.
Cresson (1944b) erected the tribe Hydrelliini
solely for Hydrellia and indicated its close relation-
ship to the tribes Hydrinini and Ilytheini. As char-
acters for Hydrelliini, he listed essentially the generic
characters. No author has disputed this classification.
However, some controversy has existed on the sub-
familial classification. Loew (1860) placed Hydrellia
with Atissa, Philygria (as Hydrina), Hyadina, and
Axysta in Hydrellina, which corresponded to a sub-
family. Becker (1926) grouped Hydrellia, Philygria,
and several parydrine genera in his subfamily "Hy-
drellinae." Cresson (1930, 1942, 1944b), Wirth and
Stone (1956), Dahl (1959), and Deonier (1964)
grouped Hydrellia with Notiphila, Dichaeta, llythea,
Philygria, Nostima, Oedenops, and several other
genera in the Notiphilinae.
I found 57 species of Hydrellia in the Nearctic
Region, of which 21 are new and one is a new
Nearctic record. I discovered and described im-
mature instars of twelve species and redescribed
those of seven species previously known.
The phylogeny of this genus remains obscure con-
cerning most details. The entire genus seems to be
in a very active speciation phase. The absence of
noticeable larval divergence within certain species
groups is very likely an indication of relatively recent
speciation. In the H. bilobifera species group, for
example, H. bilobifera and H. trichaeta can be dis-
tinguished by male and female terminalia, but the
larvae are very difficult to distinguish. Where the
species occur together, the larvae of each are often
found in the same host-plant species and even in the
same leaf of a plant. Berg (1950) and others have
reported this food (host) overlap in some other
Hydrellia species. The adults, however, are different
enough that they do not attempt to interbreed in a
laboratory colony nor, as so far observed, in natural
NUMBER 6 8 25
habitats. Most of the species groups of Nearctic
Hydrellia may well have in them similar situations
regarding speciation.
This state of the genus in the Nearctic Region has
prohibited any objective segregation of its consti-
tuents into morphologically discrete species groups.
A tentative, subjective grouping based upon overall
impressions of similarity in habitus is presented
below. This corresponds somewhat with Cresson's
impressions. A preliminary attempt to obtain a
grouping based upon computer analysis of ratios of
various measurements of male terminalia structures
proved inconclusive. In this attempt, 25 ratios were
established between the following characters: fused
surstyli length and breadth, lateral and median
lengths of fused surstyli, cercus length and breadth,
copulobus length and breadth, sterna 4 and 5
breadths, lateral and median lengths of sternum 5,
postgonite uncus length and thickness, cerci breadth
(spread), total phallus length, length and breadth
of phallus (including, in some cases, part of basi-
phallus) anterior to fused surstyli, and breadth of
distiphallus at midlength. The ratios represented
the mean of usually ten specimens (in a very few
species only one). These ratios for 53 of the species
were subjected to stepwise discriminant analysis and
then to clustering by the unweighted pair-group
method in a computer. Each species was categorized
as uncertain, present, or absent for each ratio char-
acter depending upon whether the actual ratio was
above or below the all-species mean of that ratio
character. Eleven species groups ranging from one
to ten species resulted from this program. All of
these groups had a similarity coefficient (SSm) of
0.62 or greater (where a coefficient of 1 is identity).
Only four of these correlated much with any of my
14 habitus groups. One group, bilobifera, was iden-
tical in both groupings. Another computer group
correlated fairly well with two habitus groups?
morrisoni and platygastra. A computer program
integrating all quantitative and qualitative charac-
ters gathered for the species is needed to obtain a
better generic analysis.
SPECIES GROUPS.?The following represent habitus
or physiognomic groups of Nearctic Hydrellia:
griseola group: (1) griseola, (2) valida, (3) flanicoxalis,
(4) spinicornis, (5) rixator, (6) ischiaca.
notiphiloides group: (7) notiphiloides, (8) deceptor, (9)
caliginosa, (10) pulla.
subnitens group: (11) subnitens, (12) suspecta, (13)
definita, (14) manitobae, (15) itascae, (16) amnicola,
(17) agitator,
morrisoni group: (18) morrisoni, (19) borealis, (20)
cessator, (21) serena, (22) lata, (23) penicilli.
cruralis group: (24) cruralis.
tibialis group: (25) tibialis, (26) americana, (27) biloxae.
formosa group: (28) formosa, (29) notata, (30) ains-
worthi.
crassipes group: (31) crassipes, (32) saltator, (33) proc-
teri, (34) advenae.
nobilis group: (35) nobilis, (36) idolator, (37) atro-
glauca, (38) cavator.
proclinata group: (39) proclinata, (40) melanderi, (41)
dec ens.
platygastra group: (42) platygastra, (43) wilburi.
bergi group: (44) bergi, (45) insulata, (46) per sonata.
prudens group: (47) prudens, (48) surata, (49) colum-
bata, (50) luctuosa, (51) floridana.
bilobifera group: (52) bilobifera, (53) gladiator, (54)
discursa, (55) ascita, (56) harti, (57) trichaeta.
REMARKS ON KEYS AND DESCRIPTIONS.?The high
degree of homogeneity in this genus has severely
hampered my construction of utilitarian keys to the
adults, larvae, and puparia. The extensive use of
various indices and the need to refer to male termi-
nalia limit the utility of the keys. I have not always
been able to group apparently closely related species
in the keys. Initial use of the keys will require refer-
ence to the section on definitions in the chapter on
morphology. Descriptions of male terminalia pertain
to the ventral view of the structures. Because of the
possibility that a specimen may represent a new
sibling species, one should consult the descriptions
after using the keys.
I studied the holotype or a syntype of each species
unless stated otherwise under the taxonomic remarks.
2 6 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Key to Adult Hydrellia
1. Ocellar present 2
Ocellar absent 50
2. Two subequal fronto-orbitals present 3
Three subequal fronto-orbitals present. (Male terminalia as in Figure 32.)
H. ainsworthi, new species
3. Anterior fronto-orbital anteriorly inclined, posterior one posteriorly inclined 4
Both fronto-orbitals anteriorly inclined 51
4. Face slightly convex or protruding in profile 5
Face planate and receding at constant angle in profile 53
5. Palpus moderate yellow or moderate orange-yellow 6
Palpus at least partly dark brown or black 35
6. Fore tarsus dark brown or gray in dorsal view 7
Fore tarsus moderate yellow or moderate orange-yellow 24
7. Some tibiae moderate brown or dark gray; mesokatepisternals variable 8
All tibiae moderate yellow; 1 large and 3 small mesokatepisternals. (Male terminalia as
in Figure 63.) H. cruralis Coquillett
8. Lower half of face not conically prominent; palpus moderate yellow; no sfa or 1-5
seriated sfa 9
Lower half of face conically prominent; palpus moderate orange-yellow; 4-7 scattered
sfa. (Male terminalia as in Figure 61.) H. pulla Cresson
9. Antennal segment 3 dark brown, or if partly yellow then hind basitarsus without or with
usually 2 black anterior basal setae (except H. cessator, with 4-7.) 10
Antennal segment 3 partly moderate yellow; hind basitarsus with 3?5 black, anterior
basal setae. (Male terminalia as in Figure 66.) H. penicilli Cresson
10. Fore coxa and tibia mostly dark gray or moderate brown; hind tibia without or with
usually 3-5 black, anterior apical setae 11
Fore coxa and tibia moderate yellow; hind tibia with 2 or 3 black, anterior apical setae.
(Male terminalia as in Figure 44.) H. ischiaca Loew
11. One large mesokatepisternal; basal coxals variable 12
One large and 3 or 4 small mesokatepisternals; 2 basal coxals. (Male terminalia as in
Figure 62.) H. jlavicoxalu Cresson
12. Thoracic pleuron and side of abdomen mostly light gray or light bluish gray, strongly
contrasting with mesonotum and abdominal dorsum (except in nearly uniformly light
gray H. valida) 13
Thoracic pleuron or side of abdomen mostly moderate brown or yellowish gray, not
strongly contrasting with mesonotum or abdominal dorsum 19
13. Female fore femur without distinct, black, anteroventral spines on distal half; costal
section I I : I 2.8 or less; mesofacial index 2.5 or less 14
Female fore femur with distinct, black, anteroventral spines on distal half; costal section
II : I 3.0 or more; mesofacial index 2.6 or more. (Male terminalia as in Figure 40.)
H. spinicornis Cresson
14. Large areas of body brown; usually 12 or fewer anterior interfractural costals 15
Body uniformly light gray (except mesonQtum and thoracic pleuron infrequently with
sparse light yellowish brown pruinosity in small areas) ; 12?15 anterior interfractural
costals. (Male terminalia as in Figure 22.) H. valida Loew
15. Antenna not velvety dark brown; frontal vitta and parafrontalia of different hues;
prementum usually glossy brownish black 16
Antenna velvety dark brown; frontal vitta and parafrontalia uniformly semiglossy dark
brown; prementum light gray. (Male terminalia as in Figure 42.)
H. deceptor, new species
16. Body length:wing length 0.9 or more; costal section I I : I 2.6 or less; vertex index usually
under 7.0 (except 7.5 in H. rixator) 17
Body length:wing length 0.8; costal section I I : I 2.6 or more; vertex index 7.0 or more.
(Male terminalia as in Figure 26.) H. griseola (Fallen)
17. Frontal vitta and parafrontalia of different hues; vertex index 5.5 or less; male length
usually over 1.8 mm; wing length usually over 2.0 mm 18
Frontal vitta and parafrontalia uniformly velvety dark brown; vertex index 5.5 or more;
NUMBER 6 8 27
male length usually 1.7 mm or less; wing length usually under 2.0 mm. (Male terminalia
as in Figure 29.) H. rixator, new species
18. Parafrontalia velvety black, contrasting with frontal vitta; 8-10 aristal rays; costal section
II I : IV 2.5 or less. (Male terminalia as in Figure 19.) H. caliginosa Cresson
Parafrontalia not velvety black; 6-8 aristal rays; costal section I I I : IV 2.5 or more.
(Male terminalia as in Figure 65.) H. notiphUoides Cresson
19. Male and female lengths under 2.3 and 2.7 mm respectively; fewer than 12 dorsal and
14 anterior interfractural costals; costal section I I : I usually over 2.0 20
Male and female lengths over 2.8 and 2.9 mm respectively; 12-14 dorsal and 14-16
anterior interfractural costals; costal section I I : I usually under 2.0. (Male terminalia
as in Figure 28.) H. itascae, new species
20. Face not silky light yellowish brown, without median crease or with one distinct in other
views; apicodorsal antennal prominent only in H. lata; most of fused surstyli concealed
in repose or if exposed, then not moderate yellow 21
Face silky light yellowish brown, with median crease distinct only in anteroventral view;
apicodorsal antennal prominent; most of fused surstyli always exposed and moderate
yellow. (Male terminalia as in Figure 17.) H. bergi Cresson
21. Prementum not glossy brownish black; antenna not velvety dark brown; costal section
V:IV usually 3.5 or less 22
Prementum glossy brownish black; antenna velvety dark brown; costal section V:IV 3.5
or more. (Male terminalia as in Figure 46.) H. insulata, new species
22. Prementum pale yellow; apicodorsal antennal not prominent; costal section I I : I usually
2.4 or less 23
Prementum light gray; apicodorsal antennal prominent; costal section I I : I 2.4 or more.
(Male unknown.) H. lata Cresson
23. Parafrontalia velvety black, contrasting with frontal vitta; 2 or 3 basal coxals; mesane-
pisternum bronzed in posterolateral view. (Male terminalia as in Figure 35.)
H. serena Cresson
Parafrontalia not velvety black; 1 basal coxal (infrequently 2 ) ; mesanepistemum not
bronzed in posterolateral view. (Male terminalia as in Figure 59.)
H. cessator, new species
24. Abdominal terga 2-4 not velvety purplish black medially; frontal vitta, parafrontalia,
occiput, and antennal segment 2 not uniformly velvety black 25
Abdominal terga 2?4 velvety purplish black medially; frontal vitta, parafrontalia, occiput,
and antennal segment 2 uniformly velvety black. (Male terminalia as in Figure 34.)
H. biloxiae, new species
25. Tibiae moderate brown with light gray pruinosity 26
Tibiae moderate yellow on one-third or more of their length 33
26. Antennal segment 3 at least partly moderate yellow 27
Antennal segment 3 dark brown or black 29
27. Parafrontalia not velvety black; wing length 2.4 mm or more; 6?9 aristal rays 28
Parafrontalia velvety black; wing length 2.1 mm or less; 4?6 aristal rays. (Male terminalia
as in Figure 54.) H. cavator, new species
28. Ocular index 13.0 or less; 8 or 9 aristal rays; costal section I I : I 2.4 or more. (Male
terminalia as in Figure 45.) H. subnitens Cresson
Ocular index 15.0 or more; 6-8 aristal rays; costal section I I : I 2.3. (Male terminalia not
figured.) H. suspecta Cresson
29. Tarsi moderate yellow; male hind femur without posteroventral flange and anteroventral
rows of close-set, short setae 31
Tarsi moderate orange-yellow; male hind femur with posteroventral flange and 2
anteroventral rows of close-set, short setae 30
30. Antenna dark brown; ocular index 8.0 or more; 1 basal coxal. (Male terminalia as in
Figure 33.) H- crassipes Cresson
Antenna velvety black; ocular index 7.0 or less; 2 basal coxals. (Male terminalia as in
Figure 56.) " . saltator, new species
31. Fore and mid coxae light gray anteriorly; 7 or more dorsal and 9 or more anterior
interfractural costals; ocular index usually more than 9.0 32
Fore and mid coxae moderate yellow anteriorly; 7 or fewer anterior interfractural costals;
ocular index usually under 9.0. (Male terminalia as in Figure 39.) H. advenae Cresson
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
32. Wing length 2.1 mm or less; epistomal index 1.4 or less; male hind femur very stout and
hind tibia expanded distally. (Male terminalia as in Figure 20.) H. procteri Cresson
Wing length 2.4 mm or more; epistomal index 1.4 or more; male hind femur not
stout and hind tibia not expanded distally. (Male terminalia as in Figure 45.)
H. subnitens Cresson
33. Face moderate or pale yellow; ocular index 13.0 or more; abdominal terga without
distinct bluish gray posterolateral wedges 34
Face light gray; ocular index 12.0 or less; abdominal terga with distinct light bluish
gray posterolateral wedges. (Male terminalia not figured.) H. atroglauca Coquillett
34. Wing length 2.3 mm or more; vertex index 6.0 or more; ocular index usually over
17.0. (Male terminalia as in Figure 36.) H. nobilis (Loew)
Wing length 2.1 mm or less; vertex index 5.5 or less; ocular index usually under 17.0.
(Male terminalia as in Figure 21.) H. idolater, new species
35. Mid and hind tarsi moderate orange-yellow; posterior part of male abdomen
compressed 36
Mid and hind tarsi dark brown; posterior part of male abdomen not compressed 37
36. Face velvety dark brown; antenna dark brown; 10-14 setae on basal end of costa.
(Male terminalia as in Figure 38.) ?? platygastra Cresson
Face white; antenna velvety black; 9-11 setae on basal end of costa. (Male terminalia
as in Figure 38.) H. wilburi Cresson
37. One basal coxal; 12 or fewer anterior interfractural costals; wing length 3.0 mm
or less 38
Two basal coxals; 11-14 anterior interfractural costals; wing length 3.0 mm or more.
(Male terminalia as in Figure 48.) H. manitobae, new species
38. Thoracic pleuron dark yellowish gray, brown, or light gray; wing length 2.7 mm or less;
9 or fewer aristal rays; male mid and hind tibiae not both expanded 39
Thoracic pleuron light yellowish gray; wing length 2.7 mm or more; 9-11 aristal rays;
male mid and hind tibiae expanded (Figure 8) . (Male terminalia as in Figure 16.)
H. morrisoni Cresson
39. At least one adc much longer than others; face not light yellowish brown; apicodorsal
antennal not prominent (except in H. personate); male hind tibia not expanded 40
All adc more or less uniordinate; face light yellowish brown; apicodorsal antennal
prominent; male hind tibia expanded as in Figure 8. (Male terminalia as in Figure 53.)
H. bore alts Cresson
40. Dorsal and anterior interfractural costals subequal; 2 or more pdc; male mid tibia
variable 41
Dorsal interfractural costals twice size of anterior ones; 1 pdc; male mid tibia not
expanded. (Male terminalia as in Figure 49.) H. personata, new species
41. Face not white; frontal vitta and parafrontalia not uniformly velvety black; 1 or more sfa;
male mid tibia variable 42
Face white; frontal vitta and parafrontalia (except light-brown ocellar triangle) uniformly
velvety black; no sfa; male mid tibia expanded. (Male terminalia as in Figure 27.)
H. arnerienna Cresson
42. Mesonotum and abdominal dorsum not glossy dark grayish green; face not both light gray
and slightly carinate (except in H. columbata) ; male mid tibia variable 43
Mesonotum and abdominal dorsum glossy dark grayish green; face light gray and slightly
carinate; male mid tibia expanded. (Male terminalia as in Figure 41.)
H. tibialis Cresson
43. Thoracic pleuron mostly light gray or yellowish gray; wing length 1.8 mm or more;
male mid tibia not expanded 44
Thoracic pleuron mostly moderate olive-brown (body mostly with dense moderate or
dark olive-brown pruinosity) ; wing length 1.8 mm or less (except 2.1 mm in H.
luctuosa) ; male mid tibia expanded 46
44. Ocular index 7.0 or less; mesofacial index 2.0 or less; costal section V:IV 3.5 or less 45
Ocular index 8.0 or more; mesofacial index 2.0 or more; costal section V:IV 3.6 or more.
(Male terminalia as in Figure 60.) H. dejinita Cresson
45. Antenna and parafrontalia velvety dark brown; 6-8 aristal rays; M1 + 2 index 1.5 or less.
(Male terminalia as in Figure 47.) H. amnicola, new species
NUMBER 6 8 29
Antenna and parafrontalia not velvety dark brown; 8 or 9 aristal rays; M 1 + 2 index 1.6
or more. (Male terminalia as in Figure 58.) B. agitator, new species
46. Face not entirely dark brown; usually 9 or fewer anterior interfractural costals 47
Face entirely dark brown; usually 9 or more anterior interfractural costals. (Male
terminalia as in Figure 64.) B. luctuosa Cresson
47. Ocular index 8.0 or less; face partly brown (except in H. columbata), protuberant
or carinate 48
Ocular index 9.0 or more; face entirely light gray or yellowish gray, slightly convex.
(Male terminalia as in Figure 31.) B. floridana, new species
48. Face centrally protuberant, light gray only on lower third; vertex index 4.5 or less;
mesofacial index usually 1.4 or more 49
Face slightly carinate on upper two-thirds, usually entirely light gray; vertex index 4.6
or more; mesofacial index 1.4 or less. (Male terminalia as in Figure 55.)
B. columbata, new species
49. Palpus, antenna, and parafrontalia velvety black; 4-7 sfa; lower half of mesokatepisternum
light gray. (Male terminalia as in Figure 25.) B. surata, new species
Palpus, antenna, and parafrontalia dark brown, not velvety; 1-3 sfa; thoracic pleuron
moderate olive-brown. (Male terminalia as in Figure 18.) B. prudens Curran
50. Thoracic pleuron nearly uniformly light gray; 3 fronto-orbitals. (Male terminalia as
in Figure 52.) B. notata, new species
Thoracic pleuron velvety brownish black and light gray; 2 fronto-orbitals. (Male
terminalia as in Figure 43.) B. formosa Loew
51. Thoracic pleuron with 1 continuous light-gray area 52
Thoracic pleuron with 2 distinct light-gray areas. (Male terminalia as in Figure 57.)
B. proclinata Cresson
52. Arista with 6-8 rays; 8-11 dorsal and 9-12 anterior interfractural costals. (Male
terminalia as in Figure 51.) B. melanderi, new species
Arista with 9-11 rays; 6-8 dorsal and 8-9 anterior interfractural costals. (Male
unknown.) B. decent Cresson
53. Male abdominal syntergum 9+10 rounded or only slightly bilobate posteriorly; female
cercus less than twice as long as wide 54
Male abdominal syntergum 9+10 prominently bilobate posteriorly; female cercus about
three times as long as wide (Figure 72). (Male terminalia as in Figure 50.)
B. bilobifera Cresson
54. Antennal segment 3 at least partly moderate yellow 55
Antennal segment 3 dark brown. (Female terminalia as in Figure 69; male as in Figure 15.)
B. harti Cresson
55. Ocular index over 6.0 56
Ocular index under 6.0 57
56. Male abdominal syntergum 9+10 rounded posteriorly; female cercus mucronate apically
(Figure 68). (Male terminalia as in Figure 37.) B. trichaeta Cresson
Male abdominal syntergum 9+10 truncate posteriorly; female cercus acute apically
(Figure 67). (Male terminalia as in Figure 23.) B. ascita Cresson
57. Wing length under 2.0 mm; female cercus ovoid apically, truncate basally (Figure 70).
(Male terminalia as in Figure 30.) B. discursa, new species
Wing length over 2.2 m m ; female cercus acute apically, rounded basally (Figure 7 1 ) .
(Male terminalia as in Figure 24.) B. gladiator, new species
Male Terminalia Key to Hydrellia
[The descriptions herein apply only to the ventral view and to the sclerotized portions of
structures unless otherwise stated. Only maximum dimensions are used unless otherwise stated.
The copulobus length is measured from a level perpendicular through the base of the copulobus
to the median point on the posterior margin of sternum 5.]
1. Phallus and cerci bilaterally symmetrical 2
Phallus and cerci bilaterally asymmetrical; copulobus about 25.0 times as long as median
length of sternum 5 (Figure 17) B. bergi Cresson
3 0 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
2. Copulobus length about 4.5 times (or more) as great as cercus breadth 47
Copulobus length about 3.2 times (or less) as great as cercus breadth 3
3. Basiphallus with black spinous process anterolaterally 4
Basiphallus without such process 5
4. Sternum 5 distinctly angular and prominent anterolaterally; posterolateral corner of
copulobus acutangular (Figure 26) H. griseola (Fallen)
Sternum 5 rounded anterolaterally; posterolateral corner of copulobus nearly 90* angular
(Figure 22) "? valida Loew
5. Sternum 5 with rounded prominence anterolaterally; posteromedial margin of sternum 5
and its copulobi forming squared recess; distiphallus not lamellate 6
Sternum 5 without rounded prominence anterolaterally, or if so then posteromedial margin
of sternum 5 and its copulobi not forming a squared recess; distiphallus variable 7
6. Fused surstyli narrowly and deeply cleft anteromedially, with anterolateral lobe, and
without posterolateral spicate projection; cercus about 1.5 times as long as broad
(Figure 62) H. flauicoxalis Cresson
Fused surstyli narrowly and shallowly notched anteromedially, without anterolateral lobe,
but with posterolateral spicate projection; cercus about 2.0 times as long as broad
(Figure 44) H. ischiaca Loew
7. Posteromedial part of sternum 5 with paired, long bispinate processes directed toward
anterior margin of fused surstyli; fused surstyli narrowly and deeply cleft anteromedially,
with anterolateral bladelike lobe and central gibba (Figure 40) H. spinicornis Cresson
Sternum 5 without such processes; fused surstyli variable, but without anterolateral
bladelike lobe and central gibba 8
8. Fused surstyli with 3 narrow, deep clefts anteriorly; median third of anterior margin of
sternum 5 broadly notched; copulobus with incurved posterior arm and shorter medial
arm, both with macrochaetae on ends (Figure 29) H. rixator, new species
Fused surstyli without or with only 1 cleft in anterior margin; sternum 5 not notched
anteromedially; copulobus without such arms 9
9. Postgonite uncus truncate apically and about 4.0 times as long as thick; cerci longer than
fused surstyli; copulobus sharply tapered to small tip nearly touching fused surstyli
(Figure 65) H. notiphiloides Cresson
Postgonite uncus acute or rounded apically, or if truncate then cerci shorter than fused
surstyli; copulobus not sharply tapered, or if so then not nearly touching fused
surstyli 10
10. Sternum 5 acutangular, hooklike anterolaterally; fused surstyli convex anteriorly with
narrow, shallow anteromedial notch; distiphallus longitudinally lamellate (Figure 42).
H. deceptor, new species
Sternum 5 not hooklike anterolaterally; fused surstyli not so formed anteriorly, or if so
then longer than broad; distiphallus variable 11
11. Fused surstyli about 2.0 times as broad as long and about 0.6?0.8 as long as cercus;
copulobus angularly notched medially and nearly touching fused surstyli (Figure 19).
H. caliginosa Cresson
Without such combination of characters 12
12. Pregonite 2.5-3.0 times as thick as postgonite; postgonite uncus buttonlike, about 1.5
times as long as thick; sternum 5 noticeably broader than fused surstyli (Figure 61).
H. pulla Cresson
Pregonite about 2.0 times (or less) as thick as postgonite; postgonite uncus not buttonlike;
sternum 5 variable 13
13. Cerci together as broad or broader than broadest part of fused surstyli and as long or
longer than median part of fused surstyli 14
Cerci not nearly as broad as broadest part of fused surstyli, or if so not as long as
median part of fused surstyli 22
14. Postgonite uncus projecting anteriad to or beyond distiphallus apex 15
Postgonite uncus not projecting much beyond midlength of distiphallus, or if so then
it is directed laterad 17
15. Sternum 5 noticeably broader than fused surstyli; postgonite uncus 12.0-15.0 times as
long as thick and hooked apically; copulobus about 4.0 times as long as broad
(Figure 55) H. columbata, new species
Sternum 5 not noticeably broader than fused surstyli; postgonite uncus about 4.0-6.0
NUMBER 6 8 31
times as long as thick, not distinctly hooked apically; copulobus about 2.0 times as
long as broad or broader than long 16
16. Copulobus about 2.0 times as long as broad; distiphallus broadest at midlength; postgonite
uncus nearly straight (Figure 25) H. surata, new species
Copulobus not prominent, broader than long; distiphallus broadest proximally; postgonite
uncus tip curved laterad (Figure 18) H. prudent Curran
17. Fused surstyli narrowly and deeply cleft medially along 0.9 of the length; uncus not
noticeably darker than rest of postgonite; copulobus about 4.0-5.0 times as long as
broad (Figure 28) H. itascae, new species
Fused surstyli cleft no deeper than about 0.7 of the length; uncus noticeably darker than
rest of postgonite; copulobus about 2.0-3.0 times as long as broad 18
18. Copulobus with sharp projection at midlength of medial margin; distiphallus longitudinally
lamellate (Figure 58) H. agitator, new species
Copulobus without such projection; distiphallus not longitudinally lamellate 19
19. Fused surstyli slightly concave anteriorly (nearly truncate); exposed part of basiphallus
near fused surstyli about 4.0 times as broad as distiphallus at midlength (Figure 59).
H. cessator, new species
Fused surstyli indented or cleft anteromedially; exposed part of basiphallus near fused
surstyli about 3.0 times (or less) as broad as distiphallus at midlength 20
20. Copulobus about 2.0 times as long as postgonite uncus; fused surstyli widely V-cleft
anteromedially (Figure 16) H. morrisoni Cresson
Copulobus about 5.0 times as long as postgonite uncus; fused surstyli narrowly cleft
anteromedially 21
21. Fused surstyli about 3.5 times as broad as copulobus and narrowly and deeply cleft
medially along about 0.7 of the length (Figure 66) H. penicilli Cresson
Fused surstyli about 6.0 times as broad as copulobus and narrowly cleft medially along
0.5 (or less) of the length (Figure 53) H. borealis Cresson
22. Cercus only about 0.6 as long as broad; distiphallus nearly uniform in breadth with
mucronate apex; fused surstyli broadly V-cleft anteriorly (Figure 60).
H. defbuta Cresson
Cercus more than 0.6 as long as broad, or if not then distiphallus broadly tapered to
apex that is not mucronate; fused surstyli variable 23
23. Fused surstyli about 1.4 times as long as broad, smoothly convex anterolaterally to narrow
shallow anteromedial indentation; distiphallus carinate and constricted at midlength
(Figure 48) H. mamtobae, new species
Fused surstyli usually more than or less than 1.4 times as long as broad, or if 1.4 then
concave or deeply cleft anteriorly; distiphallus not both carinate and constricted
(except in H. ainsworthi) 24
24. Anterolateral margin of sternum 5 a bare acutangular prominence; distiphallus longitudinally
lamellate; copulobus about 6.0 times as long as broad (Figure 45).
H. subnitens Cresson
Anterolateral margin of sternum 5 not a bare acutangular prominence, or if so then
copulobus only about 2.0 times as long as broad; distiphallus variable 25
25. Fused surstyli broadly and deeply V-cleft anteromedially; distiphallus not projecting much
beyond anterior margin of fused surstyli; postgonite above surstyli, not the copulobus
(Figure 47) B. amnicola, new species
Fused surstyli not broadly and deeply V-cleft anteromedially, or if so then distiphallus
projecting nearly to sternum 5; postgonite above copulobus (except in H. americana
and H. floridana) 26
26. Fused surstyli papilliform or triangular anteriorly (about 12.0-15.0 times as broad
posteriorly as anteriorly) 27
Fused surstyli not papilliform or triangular anteriorly 28
27. Fused surstyli 2.0-3.0 times as long as copulobus; pregonites and postgonites ending
near each other (Figure 63) H. cruraUs Coquillet
Fused surstyli about as long as copulobus; pregonite much larger and projecting more
anteriorly than postgonite (Figure 49) B. per sonata, new species
28. Sternum 5 with broad, flat, shallow recess anteriorly; copulobus concave posteromedially
and ending close to fused surstyli; fused surstyli about as long as broad, narrowly
notched anteromedially, and lobate anterolaterally (Figure 46) B. insulata, new species
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Sternum 5 convex or straight-edged anteriorly; copulobus without such posteromedial
concavity, or if so then not ending close to surstyli; fused surstyli not so formed 29
29. Fused surstyli slightly mucronate anteromedially, about 1.6 times as long as cercus, and
about 2.0 times as broad as sternum 4; copulobus papilliform posteriorly (Figure 41).
H. tibialis Cresson
Fused surstyli not mucronate anteromedially, usually more than or less than 1.6 times
cercus length, and less than 1.6 times as broad as sternum 4; copulobus variable 30
30. Fused surstyli, except for posterior end, evenly semicircular or ovoid and covering
basiphallus laterally 31
Fused surstyli cleft, indented, or some form other than semicircular or ovoid 32
31. Fused surstyli about 1.5 times as long as cercus; copulobus with 2 prominent macrochaetae
on tip; distiphallus apex not lamellate (Figure 64) H. luctuosa Cresson
Fused surstyli about 3.0 times as long as cercus; copulobus without macrochaetae;
distiphallus apex transversely lamellate (Figure 34) H. biloxiae, new species
32. Distiphallus distinctly expanded proximally and distally, entirely lamellate longitudinally
and bicarinate distally; fused surstyli 2.0 times (or more) as broad as long, with 3
broad anterior lobes (Figure 31) H. floridana, new species
Distiphallus not so expanded nor bicarinate, or if so then not lamellate; fused surstyli 2.0
times (or more) as broad as long in only one species (H. americana) 33
33. Postgonite absent; copulobus much broader than long; fused surstyli broadly and deeply
concave anteriorly and about 2.0 times as broad as long (Figure 27).
H. americana Cresson
Postgonite present; copulobus longer than broad; fused surstyli deeply cleft anteromedially,
or if broadly and deeply concave then not over 1.3 times as broad as long 34
34. Sternum 5 convex posteromedially, projecting as far or farther posteriad than papilliform
copulobus; fused surstyli broadly and deeply V-cleft anteromedially along 0.6 of the
length (Figure 52) H. notata, new species
Sternum 5 concave posteromedially; copulobus, if papilliform, much longer than median
length of sternum 5; fused surstyli not V-cleft along 0.6 of the length 35
35. Fused surstyli truncate anteriorly, about as long as broad and about 4.0?6.0 times as
long as cercus; copulobus notched medially (Figure 32) H. ainsworthi, new species
Fused surstyli concave, convex, cleft, or indented and variable in proportions; copulobus
variable 36
36. Fused surstyli broadly and shallowly concave anteriorly and slightly broader than long;
cercus about as long as broad; distiphallus constricted along middle portion and not
lamellate (Figure 43) H. jormosa Cresson
Fused surstyli deeply concave, V-cleft, notched or indented anteriorly, or if broadly and
shallowly concave then longer than broad or cercus only about 0.5 times as long as
broad; distiphallus lamellate if constricted along middle portion 37
37. Copulobus with 1 or 2 posterolateral winglike projections; basiphallus not covered
laterally by fused surstyli; distiphallus longitudinally lamellate 38
Copulobus without such projections; basiphallus covered laterally, or if not then distiphallus
not longitudinally lamellate throughout 39
38. Fused surstyli about 2.5 times as long as broad; copulobus with 1 posterolateral projection;
basiphallus broadest near anterior end of surstyli (Figure 36) H. nobilis (Loew)
Fused surstyli about 1.3 times as long as broad; copulobus with 2 posterolateral projections;
basiphallus broadest at about middle third of surstyli length (Figure 21).
H. idolator, new species
39. Fused surstyli convex anteriorly; copulobus with 2 setae on posterior tip about 0.5 times
as long as copulobus; distiphallus longitudinally lamellate distally (Figure 54).
H. cavator, new species
Fused surstyli concave, cleft, or notched anteriorly; copulobus without such long setae;
distiphallus variable 40
40. Fused surstyli shallowly concave anteriorly and about 1.4 times as long as broad; copulobus
verrucate on middle portion and acutangular posterolaterally (Figure 39).
H. advenae Cresson
Fused surstyli broader than long if shallowly concave anteriorly; copulobus not verrucate
on middle portion, tip variable 41
NUMBER 6 8 33
41. Distiphallus longitudinally lamellate from apex to anterior margin of fused surstyli 42
Distiphallus not longitudinally lamellate, or if so then only distally 44
42. Fused surstyli broadly and shallowly concave anteriorly and broader than long; median
length of sternum 5 about 1.3 times greater than copulobus breadth; distiphallus about
2.5 times longer than copulobus breadth (Figure 20) H. procteri Cresson
Fused surstyli U- or V-cleft anteriorly and longer than broad; median length of sternum
5 about 0.9 of copulobus breadth; distiphallus length about 5.0 times greater than
copulobus breadth 43
43. Postgonite uncus about 2.0 times as long as thick; sternum 5 nearly right-angled
anterolaterally; cercus about 1.4 times as long as broad; fused surstyli broadly U-cleft
or concave anteriorly (Figure 33) H. crassipes Cresson
Postgonite about 3.0 times as long as thick; sternum 5 broadly rounded anterolaterally;
cercus about as long as broad; fused surstyli V-cleft anteriorly (Figure 56).
H. saltator, new species
44. Fused surstyli about 1.2 times as long as broad and V- or somewhat U-cleft anteriorly 45
Fused surstyli about 1.2 times as broad as long and shallowly indented or notched
anteromedially 46
45. Sternum 5 acutangular and prominent anterolaterally; distiphallus acute apically, carinate
distally, but not longitudinally lamellate; cercus broader than long (Figure 35).
H. serena Cresson
Sternum 5 smoothly rounded anterolaterally; distiphallus rounded apically and longitudinally
lamellate on distal half; cercus longer than broad (Figure 38) H. platygastra group
46. Sternum 5 acutangular and prominent anterolaterally; copulobus and fused surstyli separated
by about 1.0 times copulobus breadth; postgonite uncus about 5.0 times (or more)
longer than thick (Figure 57) H. proclinata Cresson
Sternum 5 obtusangular anterolaterally; copulobus and fused surstyli separated by about
3.0 times copulobus breadth; postgonite uncus about 2.0 times longer than thick
(Figure 51) H. melanderi, new species
47. Posterior end of copulobus with fascicula of long macrochaetae 48
Posterior end of copulobus without such fascicula 51
48. Copulobus noticeably concave posteromedially just anterior to fascicula; sternum 5
with small acutangular prominence anterolaterally; fused surstyli shallowly concave
anteromedially (Figure 50) H. bilobifera Cresson
Copulobus not concave posteromedially; sternum 5 without small acutangular prominence
anterolaterally; fused surstyli concave or not 49
49. Sternum 5 with anterolateral right-angled projection from above; fused surstyli truncate
anteriorly; postgonite uncus about 3.0 times as long as thick (Figure 23).
H. ascita Cresson
Sternum 5 without such projection; fused surstyli shallowly or moderately concave
anteriorly; postgonite uncus about 5.0 times (or more) as long as thick 50
50. Copulobus with posteromedial digitiform lobe projecting posteriad beyond postgonite uncus
and without medial verruca; postgonite uncus bent (Figure 24).
H. gladiator, new species
Copulobus without such lobe, but with slight medial verruca; postgonite uncus nearly
straight (Figure 30) H- dhcursa, new species
51. Posterior margin of fused surstyli truncate; copulobus without noticeable convexity
posteromedially just anterior to postgonite uncus; basiphallus not covered laterally
by fused surstyli (Figure 15) # ? harti Cresson
Posterior margin of fused surstyli sinuous; copulobus with noticeable convexity
posteromedially just anterior to postgonite uncus; basiphallus covered laterally by
fused surstyli (Figure 37) ? ? trichaeta Cresson
Key to Third-Instar Larvae of some Hydrellia
1. Ventral frontoclypeal index 2.0 or less 2
Ventral frontoclypeal index over 2.0 3
2. Bifurcation index 2.5-2.7; cheliform spot touching clypeal arch margin in lateral view;
clypeal arch distinctly angular (Figure 101) H. biloxiae, new species
3 4 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Bifurcation index 4.8-5.4; cheliform spot not touching clypeal arch margin; clypeal arch
gradually sloping (Figure 89) ? ? bergi Cresson
3. Ventral frontoclypeal index under 4.0 4
Ventral frontoclypeal index 4.0 or more 14
4. Bifurcation index 4.0 or less 5
Bifurcation index over 4.0 10
5. Ventral frontoclypeal index 3.0 or more 6
Ventral frontoclypeal index under 3.0 9
6. Bifurcation index 3.4 or more; part of dorsal phragmatal ramus dark 7
Bifurcation index 3.0 or less; dorsal phragmatal ramus hyaline (Figure 90).
H. ainsworthi, new species
7. Mouth-hook, in lateral view, with distinct, thick base and thin beak; mouth-hook light
spot ovoid, less than 2.0 times as long as wide; cheliform spot arms connected 8
Mouth-hook, in lateral view, without distinct base and beak; mouth-hook light spot narrow,
triangular, and 3.0 times as long as wide; cheliform spot arms separate (Figure 103).
H. spinicornis Cresson
8. Dorsal phragmatal ramus all dark H. griseola (Fallen)
Dorsal phragmatal ramus partly hyaline (Figure 100) H. ischiaca Loew
9. Mouth-hook beak and base lengths subequal; mouth-hook light spot discal, encompassed
(Figure 97) " ? discursa, new species
Mouth-hook beak only 0.7-0.8 as long as base; mouth-hook light spot touching top margin
of base (Figure 88) H- trichaeta Cresson
10. Bifurcation index 5.5 or less; cheliform spot arms not removed from clypeal arch margin
by their length; clypeal arch not concave in lateral view 11
Bifurcation index 5.8 or more; cheliform spot arms removed from clypeal arch margin by
their length; clypeal arch slightly concave between 2 prominences in lateral view
(Figure 102) H. itascae, new species
11. Feeding apparatus partly hyaline 12
Feeding apparatus entirely dark H. morrhoni Cresson
12. Cheliform spot nearly parallel with dorsal phragmatal ramus; bifurcation index usually
under 4.5; ventral frontoclypeal index usually 3.0 or less 13
Cheliform spot distinctly oblique to dorsal phragmatal ramus (Figure 95); bifurcation
index usually over 4.5; ventral frontoclypeal index 3.0 or more....W. notiphiloides Cresson
13. Ventral frontoclypeal index 2.7 or less; mouth-hook beak distinctly longer than base;
ventral phragmatal ramus moderately dark (Figure 86) H. ascita Cresson
Ventral frontoclypeal index 2.8 or more; mouth-hook beak and base lengths subequal;
ventral phragmatal ramus mostly hyaline (Figure 91) H. bilobifera Cresson
14. Ventral frontoclypeal index over 4.5; bifurcation index 3.5 or more; mouth-hook light spot
small, elliptical; dorsal phragmatal ramus variable 15
Ventral frontoclypeal index 4.5 or less; bifurcation index 2.7 or less; mouth-hook light
spot large, ovoid; dorsal phragmatal ramus mostly hyaline (Figure 96).
H. cruralis Coquillett
15. Mouth-hook with light spot; clypeal-arch index 1.8 or more; mouth-hook base about
twice as thick as beak in lateral view 16
Mouth-hook without light spot; clypeal-arch index 1.7 or less; mouth-hook base about
1.5 times as thick as beak in lateral view H. caliginosa Cresson
16. Feeding apparatus hyaline except dark mouth-hook and cheliform spot (Figure 106);
mouth-hook beak distinctly longer than base H. pulla Cresson
Feeding apparatus mostly dark except partly hyaline ventral phragmatal ramus (Figure
92); mouth-hook beak only 0.9 as long as base H. luctuosa Cresson
Key to Puparia of some Hydrellia
1. Prothoracic end describing over half of a circle in ventral view; maximum breadth of
prothorax at midlength 2
Prothoracic end variable in shape, if describing more than half of a circle, then maximum
breadth of prothorax at or near its posterior margin 4
NUMBER 68 35
2. Head-lobe scar much longer than wide, nearly as long as prothorax 3
Head-lobe scar, nearly circular, its diameter about half of prothorax length (Figure 112).
H. ascita Cresson
3. Lengthrminimum breadth about 20.0; maximum breadth:minimum breadth about 4.2;
anal-plate index about 2.6 (Figure 115) H. trichaeta Cresson
Length:minimum breadth 12.0-15.0; maximum breadth:minimum breadth 2.4-3.2;
anal-plate 1.8-2.4 (Figure 109) H. bilobifera Cresson
4. Intersegmental constrictions inconspicuous; prothorax variable posterolaterally; bifurcate
supraspiracular spinules present or absent 5
Intersegmental constrictions extensive, making puparium distinctly scalloped laterally;
prothorax constricted posterolaterally; distinct bifurcate supraspiracular spinules present
(Figure 108) H. cruralis Coquillett
5. Maximum puparial breadth in abdomen 6
Maximum puparial breadth in metathorax 15
6. Maximum puparial breadth 5.0 times or less than maximum breadth of head-lobe scar 7
Maximum puparial breadth 5.5 times or more than maximum breadth of head-lobe
scar 13
7. Maximum breadth:minimum breadth 6.2 or less; length:minimum breadth 24.0 or less;
anal-plate variable 8
Maximum breadth:minimum breadth 7.0 or more; length:minimum breadth usually
24.0 or more; anal-plate index 2.8-3.2 (Figure 123) H. bergi Cresson
8. Maximum breadth of head-lobe scar subequal to minimum puparial breadth; length:
minimum breadth 10.0 or more; maximum breadth:minimum breadth variable 9
Maximum breadth of head-lobe scar distinctly less than minimum puparial breadth; length:
minimum breadth 10.0 or less; maximum breadth:minimum 2.6?3.6 (Figure 107).
H. luctuosa Cresson
9. Distinctly bifurcate supra- and subspiracular spinules absent 10
Distinctly bifurcate supra- and subspiracular spinules present H. mnsworthi, new species
10. Length:minimum breadth about 20.0 or less; spiracular peritremes terminal 11
Length:minimum breadth about 24.0; spiracular peritremes subterminal (Figure 121).
H. morrisoni Cresson
11. Anal-plate index 2.6 or more; anal plate ovoid or elliptical, definitely not nearly rec-
tangular 12
Anal-plate index 2.2 or less; anal plate subrectangular (Figure 118) H. tibialis Cresson
12. Length:minimum breadth 18.0 or more; anal plate reniform, anterior margin convex;
maximum puparial breadth 1.7 times or less than maximum prothoracic breadth (Figure
117) H. spinicornis Cresson
Length:minimum breadth 18.0 or less; anal plate subelliptical, anterior margin straight or
slightly concave; maximum puparial breadth 1.8 times or more than maximum pro-
thoracic breadth (Figure 122) H. griseola (Fallen) and H. ischiaca Loew
13. Maximum puparial breadth 6.0 times or less than maximum breadth of head-lobe scar;
maximum puparial breadth 2.3 times or less than maximum prothoracic breadth 14
Maximum puparial breadth 7.5 times or more than maximum breadth of head-lobe scar;
maximum puparial breadth 2.5 times maximum prothoracic breadth (Figure 116).
H. notiphiloieUs Cresson
14. Anal-plate index 3.8 or more; maximum breadth:minimum breadth 5.5 or less;
length:minimum breadth 15.0 or less (Figure 110) H. itascae, new species
Anal-plate index 3.0 or less; maximum breadth:minimum breadth 6.0 or more;
length:minimum breadth 18.0 or more (Figure 111) H. caliginosa Cresson
15. Prothorax constricted posterolaterally; anal-plate index 3.0 or less 16
Prothorax not constricted posterolaterally (Figure 113); anal-plate index 3.5 or more.
H. discursa, new species
16. Prothorax convex or somewhat triangular anteriorly, angular laterally; length:minimum
breadth 16.0 or less; abdomen nearly uniform in breadth posteriad to segment 7
(Figure 114) ?? ??**? Cresson
Prothorax semicircular anteriorly, rounded laterally; length:minimum breadth 16.0 or
more; abdomen tapering posteriad from segment 4 (Figure 119).
H. biloxiae, new species
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Hydrellia advenae Cresson
FIGURE 39
Hydrellia advenae Cresson, 1934, p. 236; 1936, p. 257;
1944, pp. 165, 174.?Procter, 1938, p. 351.?Wirth, 1965,
p. 743 [catalog listing].
DIAGNOSIS.?Palpus moderate yellow; 5-8 aristal
rays; vertex index 3.6-4.5; ocular index 6.0-9.3;
thoracic pleuron indistinctly splotched light gray and
moderate olive-brown; fore and mid coxae moderate
yellow anteriorly. Male length 1.29-1.58 mm; female
1.62-1.99 mm. Male terminalia as in Figure 39.
HEAD.?Face yellowish gray; 4?5 pfa; epistomal
index 1.2-1.7; mesofacial index 1.8-2.2; vertex
index 3.6-4.5; A-index 1.6-1.9; ocular index 6.0-
9.3. Palpus moderate yellow; 5-8 aristal rays;
antenna dark brown; frontal vitta dark gray; para-
frontale strong yellowish brown; 10-14 postoculars.
THORAX.?Ppn and mesonotum moderate brown;
3-4 adc and 2 pdc; pleuron indistinctly splotched
light gray and moderate olive-brown; fore and mid
coxae moderate yellow anteriorly, light gray laterally;
rest of legs light-gray pruinose except moderate-
yellow tarsi and mid and hind tibial apices. Wing
length 1.63-1.90 mm; veins light yellowish brown
or moderate brown; 5-7 setae on basal end of costa;
5-7 dorsal and 6-8 anterior interfractural costals;
costal-section ratios: I I : I 2.0-2.3; III:IV 2.6-3.3;
V: IV 2.9-3.5; Mi ?
 2 index 1.5-1.8.
ABDOMEN.?Terga moderate brown medially, light
gray laterally and ventrally. Male terminalia: median
third of posterior margin of sternum 5 deeply con-
cave; anterolateral margin of sternum 5 smoothly
rounded; posterolateral angle of copulobus about
35? from lateral corner; copulobus irregularly setose
and slightly verrucate at midlength laterally. Postgon-
ite bent anteriad and postgonite uncus slightly laterad;
distiphallus constricted near basiphallusr apex acute.
Anterior margin of fused surstyli concave and narrow
(only about half the midbreadth of structure) ;
B-index about 5.2; C-index 1.7-2.0.
TYPE.?Holotype male, ANSP 6513.
TYPE-LOCALITY.?Bar Harbor, Mount Desert
Island, Maine (VII-15-1933, W. Procter).
SPECIMENS EXAMINED.?9 (4o*d\ 5 $ $ ) from 2
localities:
Maine: Bar Harbor, Mount Desert Island (VII-15-1933,
W. Procter), 2?$<J, 19; Mount Desert Island (VII-12
and 25, 1935, W. Procter), 1$, 49 9.
Hydrellia agitator, new species
FIGURE 58
DIAGNOSIS.?Palpus dark brown; 8-9 aristal rays;
ocular index 6.0-7.0; Mi * 2 index 1.6-1.8; meso-
notum strong yellowish brown; thoracic pleuron light
gray except upper half of mesanepistemum semi-
glossy moderate olive-brown; abdominal dorsum
glossy dark grayish green in posterior view. Male
length 1.70-1.79 mm; female 1.79-2.21 mm. Male
terminalia as in Figure 58.
HEAD.?Face yellowish gray or light gray; 3-5
pfa; epistomal index 1.3-1.5; mesofacial index 1.8-
2.0; vertex index 6.3-8.7; A-index 1.8-2.4; ocular
index 6.0-7.0. Palpus dark brown; 8-9 aristal rays;
antenna dark brown; frons usually uniformly light
brown; 14-16 postoculars.
THORAX.?Ppn and mesonotum moderate olive-
brown; 3-4 adc and 2 pdc; pleuron light gray except
upper half of mesanepistemum semiglossy moderate
olive-brown; legs sparsely moderate olive-brown
pruinose except light-gray or yellowish-gray coxae
and trochanters. Wing length 1.87-2.30 mm; veins
light yellowish brown; 6?7 setae on basal end of
costa; 6-8 dorsal and 7-8 anterior interfractural
costals; costal-section ratios: I I : I 2.3-3.4; III:IV
2.8-3.2; V:IV 3.2-3.5; Mx + 2 index 1.6-1.8.
ABDOMEN.?Terga semiglossy and with moderate
olive-brown pruinosity over dark brown (glossy dark
grayish green in posterior view). Male terminalia:
median third of posterior margin of sternum 5 deeply
concave and congruent with distiphallus; anterolat-
eral margin of sternum 5 rounded; copulobus regu-
larly setose, acutangular posterolaterally and with
acuminate, peninsulate, posteromedial projection.
Postgonite bent anteromediad; postgonite uncus
short, bent laterad; distal half of distiphallus narrow,
spa tula te, and ventrally lamellate (distinctly delim-
ited from basiphallus and protruding far beyond
fused surstyli). Anterior margin of fused surstyli
deeply and broadly concave; B-index about 1.2;
C-index 2.0-2.2.
TYPE.?Holotype male, USNM 70525.
TYPE-LOCALITY.?Port Saint Joe Beach, Gulf
County, Florida (111-17-1954, G. Steyskal).
PARATYPES.?3 9 $ from 3 localities:
Florida: Green Cove Springs (111-24-1952, J. Vocke-
roth), 1$ (ISC).
Georgia: Billy's Island (VI-1912, collector unknown),
19 (ANSP).
NUMBER 68 37
Mississippi: Lake Shady, Lamar County (VII-11-1962,
D. L. Deonier), 19 (ISC).
REMARKS.?Adult. Habitat recorded: on mats of
Hydrochloa caroliniensis.
Hydrellia ainsworthi, new species
FIGURES 32, 83, 90, 99, 120
DIAGNOSIS.?Palpus moderate yellow; 6-8 aristal
rays; vertex index 5.2-6.8; body length:wing length
1.0; 3 fronto-orbitals, all anteriorly inclined; ante-
costa of male tergum 3 strongly curved posteriad in
lateral view. Male length 1.33-1.53 mm; female
1.45-1.79 mm. Male terminalia as in Figure 32.
HEAD.?Face light yellowish brown; 4-5 pfa; 3-7
sfa; epistomal index 1.6-1.8; mesofacial index 2.5-
3.5; vertex index 5.2-6.8; A-index 1.5-2.0; ocular
index 7.0-7.5. Palpus moderate yellow; 6-8 aristal
rays; antenna and most of frons moderate brown;
fronto-orbital area strong yellowish brown; 3 fronto-
orbitals, all anteriorly inclined; 11-15 postoculars.
THORAX.?Ppn and mesonotum dark gray except
light-gray postalar wall; 2?4 adc and 2 pdc; pleuron
light gray; legs light-gray pruinose except dark-
brown tarsi. Wing length 1.36-1.82 mm; veins dark
brown; 5-7 setae on basal end of costa; 5-7 dorsal
and 6-9 anterior interfractural costals; costal-
section ratios: II:I 2.0-2.5; III:IV 2.6-3.0; V:IV
3.0-3.5; Mi
 + 2 index 1.2-1.8.
ABDOMEN.?Terga semiglossy and bronzed medi-
ally, light gray laterally and ventrally; antecosta of
male tergum 3 strongly curved posteriad in lateral
view; ventral lobes of male terga 1-4 overlapped to
form hillock. Male terminalia: median third of
posterior margin of sternum 5 concave; anterolateral
margin of sternum 5 rounded (115?-120? angular) ;
copulobus slightly convex posteriorly, notched at
midlength on medial side, and regularly setose.
Postgonite bent slightly mediad and postgonite uncus
slightly laterad from general anterior direction;
distiphallus slightly constricted at midlength, ven-
trally carinate, and mucronate apically. Anterior
margin of fused surstyli truncate; B-index about 6.0;
C-index 1.0-1.2.
SECOND-INSTAR LARVA.?Length 1.75-2.50 mm;
maximum breadth 0.35-0.50 mm. Frontoclypeal
length 0.29 mm (Figure 90). Thorax with several
rows of dorsal spinules. Other characters similar to
those in third-instar larva.
THIRD-INSTAR LARVA.?Length 2.50-3.50 mm;
maximum breadth 0.50-0.65 mm. Frontoclypeal
length 0.40-0.45 mm; dorsal phragmatal ramus
hyaline (Figure 99). Ventral frontoclypeal index
3.4-3.6; phragmatal index 0.9-1.0; bifurcation
index 2.8-3.0; clypeal-arch index 1.4-1.6. Clypeal
arch sloping at 25?-30? and slightly concave.
Mouth-hook beak and base lengths subequal; maxi-
mum mouth-hook base thickness about 1.2 times
that of beak. First two thoracic segments moderately
spinulose; prothorax faintly rugulose ventrally;
abdominal segment 8 with only about 5 annuli of
spinules, many black and distinctly bifurcate; other
dorsal spinules mostly intersegmental. Body trans-
lucent, yellowish gray.
PUPARIUM.?Length 2.25-3.25 mm; maximum
breadth 0.65-0.75 mm; subcylindrical (Figure 120).
Puparial length: minimum breadth 14.0-15.5; maxi-
mum breadth: minimum breadth 4.0-4.5; anal-plate
index 2.3-2.6. Prothoracic end semicircular in ven-
tral view; head-lobe scar subcircular or ovoid; maxi-
mum puparial breadth in abdomen and 5.0 times or
less than maximum breadth of head-lobe scar;
breadth of head-lobe scar and minimum puparial
breadth subequal; anal plate subovoid, with anterior
margin straight or slightly convex. Empty puparium
translucent, light yellowish brown.
TYPE.?Holotype male, USNM 70526.
TYPE-LOCALITY.?Lake Shady, 7 miles west of
Hattiesburg, Lamar County, Mississippi (VII-11-
1962, D.L. Deonier).
PARATYPES.?433 (261 d d\ 172 ? ? ) from 9 local-
ities :
Florida: Inglis (VII-7-1948, E. Todd), 13 (UKC);
Lake Worth's Bay Shore (VIII-8-1951, W. Wirth), 1$
(USNM); Lacoochee (VI11-18-1930, P. Oman), 1?
(UKC); Orange Park (111-25-1952, J. Vockeroth), 19
(CNC); Punta Gorda (IV-11-1952, J. Vockeroth), 19
(CNC); Silver Springs, near Ocala, Marion County (IV-
5-1953, W. Mason), 19 (CNC).
Mississippi: Dickinson's Pond, sec. 3, T.4 north, R. 14
west, Lamar County (VII-11-1962, D. L. Deonier), 48,5 3 ,
459 9, (VII-24-1962, D. L. Deonier), 163 3 , 209 9
(USNM and ISC); Lake Shady, Lamar County (VI-29-
1962, D. L. Deonier), 413 3 , 199 9, (VI-29-1962, J. H.
Ainsworth), 4 3 3 , 39 9 , (VII-11-1962, D. L. Deonier),
151 3 3 , 78 9 9 (USNM, ISC, and DLD); Vancleave Road,
Jackson County (VI-15-1962, D. L. Deonier), 1 3 , 19
(USNM and DLD).
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
IMMATURE SPECIMENS EXAMINED.?23 from 2
localities:
Mississippi: Dickinson's Pond, sec. 3, T.4 north, R.I4
west, Lamar County (collected and reared by D. L. Deo-
nier); Lake Shady, Lamar County (collected and reared by
D. L. Deonier).
REMARKS.?This species is similar in some respects
to H. notata, but it differs from it and other species
by the orientation of the male ventral tergal lobes.
Adult. In June and July, I collected adults from
leaves of Hydrochloa caroliniensis and Nelumbo
lute a and one specimen with several H. biloxiae in a
Grigarick floating trap in Jackson County, Mississippi.
Larva and puparium. Larvae mined in the stems,
leaf sheaths, and leaf blades of the host plants. Three
larvae pupariated in leaf sheaths in the laboratory.
One adult emerged six days later. I found no escape
slits in any of the many mines containing puparia.
Host plants. Of several species examined, I found
water grass, Hydrochloa caroliniensis, to be the sole
host plant. My survey included water grass and its
associates, Polygonum hyperpiperoides, Xyris caro-
liniana, Hypericum punctatum, Potamogeton diver-
sifolius, and Juncus repens. The mats of water grass
seemed to have a low percent of infestation. I found
no infestation in several samples collected in Sep-
tember, during a drought.
Hydrellia americana Cresson
FIGURE 27
Hydrellia americana Cresson, 1931, p. 106; 1944b, pp. 164,
172.?Wirth, 1965, p. 743 [catalog listing].
DIAGNOSIS.?Palpus black; 5-6 aristal rays; face
white; body length:wing length 0.8-1.0; frons
(except light-brown ocellar triangle and fronto-
orbital sockets) velvety black; thoracic pleuron dark
brown except light-brown mesanepisternum and
mesepimeron. Male length 1.29-1.68 mm; female
1.45-1.80 mm. Male terminalia as in Figure 27.
HEAD.?Face white; 4-5 pfa; no sfa; epistomal
index 1.0?1.3; mesofacial index 1.5-1.8; vertex
index 4.4-5.6; A-index 1.5-2.7; ocular index 5.5-
6.5. Palpus black; 5-6 aristal rays; antenna very
dark brown (segment 2 velvety); frons (except light-
brown ocellar triangle and fronto-orbital sockets)
velvety black; postocular area dark brown except
lower one-fourth moderate olive-brown; 12-14
postoculars.
THORAX.?Ppn and mesonotum dark gray; 3-4
adc and 2 pdc; pleuron dark brown except light-
brown mesanepisternum and mesepimeron; legs dark
brown except yellowish-gray coxae; male mid tibia
expanded. Wing length 1.53-1.80 mm; veins light
yellowish brown; 6-7 setae on basal end of costa;
6-8 dorsal and 6-10 anterior interfractural costals;
costal-section ratios: I I : I 2.0-2.5; I I I : IV 3.0-3.7;
V:IV 3.0-3.7; Mx + 2 index 1.5-2.0.
ABDOMEN.?Terga semiglossy dark gray. Male
terminalia: median third of posterior margin of
sternum 5 broadly notched; anterolateral corner of
sternum 5 right-angled; copulobus broader than long
and irregularly setose except 4-5 macrochaetae seri-
ated on posterior margin. Postgonite inapparent;
distiphallus uniformly tapering to about two-thirds
of length, the remaining distal third of uniform
breadth to blunt apex. Anterior margin of fused
surstyli broadly notched; B-index about 0.8.
TYPE.?Holotype female, USNM 43454.
TYPE-LOCALITY.?Chesapeake Beach, Maryland
(VIII-2-no year, J. M. Aldrich).
SPECIMENS EXAMINED.?13 (3<Sd, 10$ $ ) from
11 localities:
Maine: Machias (VII -17 and 19-no year, C. Johnson),
2 9 9 .
Maryland: Chesapeake Beach (VIII -18 -1922 , J. Mal-
loch), \$, (VII -3-1924 , J. Malloch), 1 9 ; Kent Narrows
(VII-1954, M. Wheeler), 1 9 .
Massachusetts: Woods Hole (VII -21 -1954 , M. Wheeler),
1 9 .
Michigan: Baraga County (111-27-1952, R. Dreisbach),
1 9 .
Mississippi: Bay St. Louis (VI-17-1917 , collector un-
known), 1 9 ; Bellefontaine Point, Jackson County ( V I I -
10-1962, D. L. Deonier), 1 9 ; Dickinson's Pond, Lamar
County (VII -24-1962 , D . L. Deonier), 1 9 ; Gulf Coast
Research Laboratory, Ocean Springs ( V I - 5 - 1 9 6 2 , D. L.
Deonier), 1 <$ ; Vancleave Road, Jackson County, 30?25.1'
N, 88?46' W (VI-23-1962 , D. L. Deonier), 1 3 .
New York: Cold Spring Harbor, Long Island (VH-date
and collector unknown), 1 9 ; Wilmington Notch, Adiron-
dacks (VII -3 -no year, J. Aldrich), 1 9 .
REMARKS.?This species is very similar to H.
tibialis, but its male terminalia are distinctive. This
species also reportedly occurs in California.
Adult. Habitat recorded: leaves of Sporobolus
indica on sea beach.
NUMBER 6 8 39
Hydrellia amnicola, new species
FIGURE 47
DIAGNOSIS.?Palpus dark brown; 6-8 aristal rays;
body length:wing length 0.9-1.0; antenna velvety
dark brown; thoracic pleuron light gray except upper
one-sixth of mesanepimeron light brown; Mi+2
index 1.3-1.5. Male length 1.87-2.04 mm; female
2.04-2.38 mm. Male terminalia as in Figure 47.
HEAD.?Face light gray or yellowish gray; 4-5
pfa; epistomal index 1.1-1.5; mesofacial index 1.5-
1.8; vertex index 5.5-7.0; A-index 1.6-2.0; ocular
index 5.5-6.2. Palpus dark brown; 6-8 aristal rays;
antenna and most of parafrontale velvety dark
brown; frontal vitta semiglossy dark brown; fronto-
orbital area moderate olive brown; 13?17 post-
oculars.
THORAX.?Ppn and mesonotum dark brown; 3-4
adc and 2 pdc; pleuron light gray except upper one-
sixth of mesanepimeron light brown; legs dark brown
with light-gray pruinosity sparse except on coxae and
trochanters. Wing length 2.13-2.38 mm; veins dark
brown; 7-8 setae on basal end of costa; 6-10 dorsal
and 8-11 anterior interfractural costals; costal-
section ratios II:I 2.0-2.3; III:IV 2.5-3.0; V:IV
2.7-3.3; Mi ? 2 index 1.3-1.5.
ABDOMEN.?Terga dark brown dorsally, light gray
laterally and ventrally. Male terminalia: median
third of posterior margin of sternum 5 slightly con-
cave; anterolateral margin of sternum 5 smoothly
rounded; copulobus inapparent (actually widely
curving and acuminate). Postgonite covered by
surstyli and nearly straight except preapical part bent
sharply laterad; postgonite uncus straight and di-
rected laterad; distiphallus acuminate; basiphallus
completely covered by surstyli. Anterior margin of
fused surstyli cleft medially to about midlength of
structure; B-index about 2.2; C-index 1.8-2.2.
TYPE.?Holotype male, USNM 70527.
TYPE-LOCALITY.?Sucker Creek, 100 yards north
of Highway 31, Clearwater County, Minnesota
(VII-23-1963, D. L. Deonier).
PARATYPES.?28 (80* cf, 20 9 $. ), with same collec-
tion data as for holotype (USNM, ISC, and DLD).
REMARKS.?There are some similarities between
the male terminalia of this species and those of the
H. prudens species group, but the habitus is very
dissimilar.
Adult. Habitat recorded: leaves of Glyceria grandis.
At the type-locality, I observed three pairs attempting
to copulate. Also, I saw one female try to mount
another female, and a male advance aggressively upon
a female H. griseola. I tried unsuccessfully to force
two captive females to oviposit. Both of them died
after three days of captivity.
Hydrellia ascita Cresson
FIGURES 23,67,86, 112
Hydrellia ascita Cresson, 1942, p. 78; 1944b [in part], pp.
170, 175.?Berg, 1949 [in part], p. 284; 1950 [in part],
pp. 375, 376, 378, 384, 385 (pi. 2, fig. 2), 388, 389 (pi. 3,
fig. 2), 391-392, 393, 396 [biology, morphology, and tax-
onomy of immatures].?Deonier, 1964, pp. 115, 125, fig.
4; 1965, p. 500 [ecology].?Wirth, 1965, p. 743 [catalog
listing].
Diagnosis.?Palpus moderate yellow; 5-8 aristal
rays; face planate; vertex index 5.4-6.8; ocular index
6.2-8.0; male abdomen truncate posteriorly; female
cercus triangular with truncate base; antennal seg-
ment 3 at least partly moderate yellow; apicodorsal
antennal prominent. Male length 1.79-2.10 mm;
female 1.60-2.28 mm. Male terminalia as in Figure
23; female as in Figure 67.
HEAD.?Differing mainly from H. bilobifera in the
following: epistomal index 1.4-1.8; mesofacial index
1.8-2.4; ocular index 6.2-8.0.
THORAX.?Differing mainly from H. bilobifera in
the following: 3-4 adc and 2 pdc; wing length
1.87-2.04 mm.
ABDOMEN.?Terga semiglossy dark gray medially,
light gray laterally and ventrally. Male terminalia:
median third of posterior margin of sternum 5 deeply,
rectangularly recessed; anterolateral margin of ster-
num 5 with dorsal process projecting laterad (form-
ing 90?-100? angle with copulobus); posterior mar-
gin of copulobus only slightly convex (partly or
wholly obscured by loose fascicula of 5-7 strong
macrochaetae) ; medial margin of copulobus dis-
tinctly concave on posterior third and with seriated
setae on medial third, verrucate laterally with bare
space just posteriad from verruca. Postgonite bent
mediad; acuminate postgonite uncus curved anteriad;
distiphallus constricted proximally and preapically,
with notched ventrally lamellate apex, and bicarinate
ventrally on basal two-thirds; most of basiphallus
exposed ventrally. Anterior margin of fused surstyli
truncate (much narrower than posteriorly) ; B-index
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
about 1.8; C-index 0.8-1.0. Syntergum 9 + 1 0 trun-
cate posteriorly. Female terminalia: tergum 8 shorter
than sternum 8; cercus irregularly setose dorsally and
laterally, triangular with truncate base, and less than
1.5 times as long as wide (lateral view). Segment 7
with group of about 6 seriated setae laterally on
posterior margin; median spermatheca similar to that
of H. discursa (Figure 70).
FIRST-INSTAR LARVA.?Length 0.80-1.20 mm;
maximum breadth 0.16-0.18 mm. Frontoclypeal
length 0.16-0.18 mm; frontoclypeus moderate brown
except black cheliform spot. As with most of the
other species investigated, the creeping welts of the
first instar are larger and more distinct than in later
instars. This description in part after Berg (1950).
SECOND-IN STAR LARVA.?Length 1.20-2.40 mm;
maximum breadth 0.20-0.50 mm. Frontoclypeal
length 0.23-0.29 mm. Other characters similar to
those in the first-instar larva.
THIRD-INSTAR LARVA.?Length 2.40-5.00 mm;
maximum breadth 0.60-0.80 mm. Frontoclypeal
length 0.35-0.49 mm; ventral phragmatal ramus
moderately dark (Figure 86). Ventral frontoclypeal
index 2.5-2.7; phragmatal index 1.0-1.2; bifurca-
tion index 4.2-4.4; clypeal-arch index 1.5-1.8.
Clypeal arch sloping at 10?-15?, with slight inden-
tation at level of cheliform spot. Mouth-hook beak
distinctly longer than base; maximum mouth-hook
base thickness about 2.0 times that of beak. Prothorax
and mesothorax sparsely spinulose; abdominal seg-
ment 8 without ventral, transverse row of setulae,
but with 3-4 annuli of spinules. Body translucent,
with greenish-yellow tinge.
PUPARIUM.?Length 2.80-4.25 mm; maximum
breadth 0.80-1.15 mm; subcylindrical (Figure 112).
Puparial length: minimum breadth 14.0-21.0; maxi-
mum breadth: minimum breadth 3.5-4.0; anal-plate
index 1.8-2.2. Prothoracic end describing more than
half of a circle in ventral view; head-lobe scar sub-
circular, with maximum breadth about half of
prothorax length; anal plate subovoid, with distinctly
convex anterior margin. Empty puparium trans-
lucent, light yellowish brown. Early pupa with
greenish-yellow tinge.
TYPE.?Holotype male, ANSP 6621.
TYPE-LOCALITY.?Nigger Creek, Cheboygan Coun-
ty, Michigan (VIII-21-1941, C. O. Berg; reared
from Potamogeton tenuifolius).
ADULT SPECIMENS EXAMINED.?32 ( H c f c T ,
21 $ $ ) from 12 localities:
Illinois: Havana (IX-17-1895, C. Hart) , 19 .
Iowa: Ledges State Park, Boone County, 41?59.5' N,
93?53.5' W (IX-12-1959, J. Laffoon, 1 $ , (VII-28-1954,
J. Laffoon), 1 9 ; Springbrook State Park, Guthrie County
(VI-23-1961, D. L. Deonier), 2 9 9-
Kansas: Leavenworth County Lake (VIII-27-1962, D. L.
Deonier), 39 9-
Michigan: Cheboygan County (VI-12-1940, C. Berg),
It, (VI-26-1946, C. Berg), 1$ , 29 9 , (VII-5-1946,
C. Berg), 1 9 , (VIII-7-1947, C. Berg), 1$ ; Bessey Creek,
Cheboygan County (VIII-14-1941, C. Berg), 1 9 ; Douglas
Lake, Cheboygan County (VI1-3-1941, C. Berg), 1 9 ;
Nichol's Bog, Cheboygan County (VII-5-1940, C. Berg),
1 $, (VII-5-1941, C. Berg), 1 9 ; Nigger Creek, Cheboygan
County (VI-20-1940, C. Berg), 2$$, (VI-21-1941, C.
Berg), 6 9 9.
Minnesota: Chisago County (no date, O. Oestlund), 1 9.
Ontario: Midland (VIII-18-1955, J. Chillcott), 3 t t ,
19.
Tennessee: Samburg, Reelfoot Lake (VIII-11-1962, D.
L. Deonier), 2$ $.
IMMATURE SPECIMENS EXAMINED.?20 from 2
localities:
Michigan: Nigger Creek, Cheboygan County (collected
and reared by C. Berg); Sodon Lake, Oakland County (col-
lected and reared by C. Berg).
REMARKS.?As with many other species, especially
those of the H. bilobifera group, there is some pos-
sibility of misdetermination, since I was not allowed
to prepare the holotype terminalia for study.
Adult. Habitats recorded: leaves of Potamogeton
nodosus, P. angustifolius, P. gramineus, and Nuphar
advena. I found too few adults of H. ascita to make
any observations on their behavior. According to
Berg (1950), H. ascita showed no emergence maxima
in Cheboygan County and Oakland County,
Michigan.
Larva and puparium. Berg found that the larvae
preferred flaccid, submerged leaves, but that many
mined in floating leaves and in linear submerged
leaves of P. foliosus. He discovered that third-instar
larvae commonly pupariate in the leaf axil and insert
their spiracular peritremes in the main stem. Berg
observed no puparia situated entirely in stems. Many
times the leaves disintegrated, leaving the puparia
projecting out from the nodes.
No stadial measurements have been made for this
species.
Host plants. Berg reared this fly from Potamogeton
NUMBER 6 8 41
alpinus, P. amplifolius, P. ephihydrus, P. foliosus,
P. illinoensis, P. oakesianus, P. richardsonii, and
P. zosteriformis.
Parasites. Berg reared two braconids?Chorebidella
bergi and Dacnusa sp.?and the diapriid Trichopria
columbiana from the puparia of H. ascita.
Hydrellia atroglauca Coquillett
Hydrellia atroglauca Coquillett, 1910a, p. 131.?Cresson,
1931, pp. 107, 108; 1944b, pp. 169, 172.?Wirth, 1956,
p. 16; 1965, p. 744 [catalog listing].
DIAGNOSIS.?Palpus moderate yellow; 9-11 aristal
rays; face light gray; vertex index 5.8-6.4; ocular
index 12.0 or less; antennal segment 3 partly or
wholly moderate yellow; thoracic pleuron light gray;
tibiae (one-third or more of length) moderate yellow;
abdominal dorsum semiglossy moderate brown medi-
ally, and with distinct light blue-gray posterolateral
wedges. Male length 2.13 mm; female 2.55-2.64
mm.
HEAD.?Face light gray; 6-8 pfa; 1-2 sfa; epis-
tomal index 1.5-1.8; mesofacial index 2.2-2.7;
vertex index 5.8-6.4; A-index 1.6-2.0; ocular index
7.0-12.0. Palpus moderate yellow; 9-11 aristal rays;
frons uniformly strong yellowish brown; antenna
moderate brown except segment 3 partly or wholly
moderate yellow; 16-18 postoculars.
THORAX.?Ppn and mesonotum light brown; 3-4
adc and 2 pdc; pleuron light gray; legs light-gray
pruinose except moderate-yellow (one-third or more
of length) tibiae and tarsi. Wing length 2.72-2.79
mm; veins light yellowish brown; 6-7 setae on
basal end of costa; 8-10 dorsal and 10-13 anterior
interfractural costals; costal-section ratios: I I : I 2.2-
2.5; I I I : IV 2.7-3.1; V:IV 3.0-3.8; Mi ?
 2 index
1.4-1.6.
ABDOMEN.?Terga semiglossy moderate brown me-
dially and with conspicuous light blue-gray postero-
lateral wedges (in some females entire posterior
margin of tergum 5 light blue-gray).
TYPES.?Syntype male and two females (the male
and one female, USNM 13101).
TYPE-LOCALITY.?Biscayne Bay Florida (no date,
Mrs. A. T. Slosson).
SPECIMENS EXAMINED.?1 cf, 1 9 syntype, and 1 9
from Royal Palm Park, Florida (1-29-1933, A. L.
Melander).
REMARKS.?Hydrellia idolator is very similar to
this species, but the former has a pale-yellow premen-
tum, an ocular index of 13.0-17.5, and 6-8 dorsal
and 6-10 anterior interfractural costals.
Hydrellia bergi Cresson
FIGURES 17,74,89, 123
Hydrellia bergi Cresson, 1941, p. 37; 1944b, pp. 170, 175.?
Berg, 1950, pp. 375, 376, 378, 382, 383 (pi. 1, figs. 1-3)
384, 385 (pi. 2, fig. 3), 387-388, 389 (pi. 3, fig. 3), 390-
391, 393, 396 [biology, morphology, and taxonomy of im-
matures].?Judd, 1964, p. 411 [ecology].?Wirth, 1965, p.
744 [catalog listing].
DIAGNOSIS.?Palpus moderate yellow; 7-8 aristal
rays; vertex index 5.8-6.4; apicodorsal antennal
prominent; median facial crease distinct only in
anteroventral view; thoracic pleuron mostly light
gray. Male length 2.04-2.13 mm; female 2.04-2.21
mm. Male terminalia as in Figure 17.
HEAD.?Face silky light yellowish brown with
median crease distinct only in anteroventral view;
4?5 pfa; epistomal index 1.3-1.7; mesofacial index
1.9-2.3; vertex index 5.8-6.4; A-index 1.5-1.8;
ocular index 6.0-7.0. Palpus moderate yellow; 7-8
aristal rays; antenna and most of parafrontale very
dark brown or black; fronto-orbital area and frontal
vitta light brown; 13-17 postoculars.
THORAX.?Ppn semiglossy dark gray; mesonotum
dark brown; 3-4 adc and 2 pdc; pleuron light gray
except moderate olive-brown on posterior half of
mesanepimeron; legs sparsely light-gray pruinose
except dark-brown tarsi. Wing length 1.87-2.21 mm;
veins usually dark brown; 6?8 setae on basal end of
costa; 5-7 dorsal and 7-9 anterior interfractural
costals; costal-section ratios: I I : I 2.0-2.3; I I I : IV
3.0-3.4; V . IV 3.4-3.7; Mx + 2 index 1.2-1.6.
ABDOMEN.?Terga semiglossy dark gray. Male
terminalia: median third of posterior margin of
sternum 5 notched at obtuse angles; anterolateral
margin of sternum 5 angular (90?-100?); copulobus
acutangular and slightly incurved posteriorly, and
regularly setose. Postgonite covered by copulobus
and postgonite uncus bent 90? anteriad; disti-
phallus long, asymmetrical, with apex curved to
right. Anterior margin of fused surstyli slightly con-
vex; surstyli yellow and mostly exposed; B-index 3.2;
C-index 0.2-0.4.
EGG.?Length 0.59-0.68 mm; maximum breadth
0.14-0.17 mm. Chorion (Figure 74) yellowish gray,
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
corrugate, with light-brown (frequently dark) ridges
flanked by regular, wavy, perpendicular striae. Gen-
eral color aspect variable within one population,
with some being uniformly yellowish gray and others
with ridges light brown or dark gray. Micropylar
protuberance infundibulate, concealed in dorsal view
by a hoodlike chorionic projection.
FIRST-INSTAR LARVA.?Length 0.80-1.50 mm;
maximum breadth 0.16-0.22 mm. Feeding apparatus
similar in shape and coloration to that of third-
instar larva; frontoclypeal length 0.16-0.18 mm.
General color aspects of body similar to those of
H. ischiaca.
SECOND-INSTAR LARVA.?Length 1.50-3.00 mm;
maximum breadth 0.20-0.50 mm. Frontoclypeal
length 0.25-0.27 mm. Other characters similar to
those of first-instar larva.
THIRD-INSTAR LARVA.?Length 2.00-5.20 mm;
maximum breadth 0.25-0.75 mm. Frontoclypeal
length 0.40-0.52 mm (Figure 89). Ventral fronto-
clypeal index 1.6-1.8; phragmatal index 0.9-1.0;
bifurcation index 4.8-5.4; clypeal-arch index 1.6-
1.8. Clypeal arch sloping slightly convexly at about
20?. Mouth-hook beak and base equal in length;
maximum mouth-hook base thickness about 1.5 times
that of beak. Body translucent, greenish yellow.
PUPARIUM.?Length 3.50-4.50 mm; maximum
breadth 0.80-1.15 mm; subcylindrical (Figure 123).
Puparial length: minimum breadth 23.0-27.0; maxi-
mum breadth: minimum breadth 7.0-9.0; anal-plate
index 2.8-3.2. Prothoracic end slightly convex in
ventral view; head-lobe scar subcircular and 0.2 or
less of maximum puparial breadth; prothorax dis-
tinctly rugulose in ventral view; abdominal segment
8 without ventral, transverse row of setulae and with
spinules sparse, irregular; anal plate subovoid, with
anterior margin straight or slightly convex. Empty
puparium translucent, light brown.
TYPE.?Holotype male, USNM 7451.
TYPE-LOCALITY.?Nigger Creek, Cheboygan
County, Michigan (VI-27-1940, C. O. Berg).
ADULT SPECIMENS EXAMINED.?108 (46c?1 cf,
6 2 ? ? ) from 20 localities:
Michigan: Bessey Creek, Cheboygan County (VI-10-
1940, C. Berg), *$$, 2 9 9 , (VIII -14-1941 , C. Berg),
\$, 2 9 9 ; Cheboygan County (VII-12-1946, C.
Berg), 1 9 , (VII-18-1946 , C. Berg), \$, ( V I I I - 1 8 -
1946 , C. B e r g ) , 1 9 , ( V I I - 2 1 - 1 9 4 7 , C. Berg) ,
1 9 , (VIII -7 -1947 , C. Berg), 1 9 ; Douglas Lake,
Cheboygan County (VI1-5-1940, C. Berg), 1 3 , ( V I -
16-1941, C. Berg), 1$, (VII -9 -1941 , C. Berg), 2$$,
7 9 9 ; Gaylord (VII-17-1938 , C. Sabrosky), 1 9 ; Ocqueoc
Lake, Presque Isle County (VII -13-1941 , C. Berg), 1 9 ,
(VIII -1 -1941 , C. Berg), 2 $ $ ; Third Sister Lake, Wash-
tenaw County (VI-28-1942, C. Berg), 1 9 -
Minnesota: Eaglenest ( IX-1-1958 , W. Balduf), 1 9 ;
Iron Corner Lake, Itasca State Park (VI -18 -1963 , D. L.
Deonier), 1 9 ; Mississippi River at Sucker Creek, Clear-
water County (VII -9-1963 , D. L. Deonier), \$, 2 9 9 ;
Mississippi River, 150 yards north of Highway 31, Clear-
water County (VII -9-1963 , D. L. Deonier), 1 4 , 3 9 9 ;
Squaw Lake, Itasca State Park (VIII -14-1963 , D. L. Deo-
nier), 5$ $, 3 9 9 ; Rapid River Logging Camp, Hubbard
County (VII-2-1963 , D. L. Deonier), 1 9 ; 1.3 miles south
of main entrance, Itasca State Park ( V I - 2 6 - 1 9 6 3 , D. L.
Deonier) 9 3 4 , 7 9 9 .
New York: Dryden Lake, Tompkins County ( V I I I - 8 -
1961, D. L. Deonier), \1 $ &, 199 9 ; Franklinton ( V I -
20-1923, M.Leonard), 2 9 9 .
Ontario: London ( IX-12-1958 , W. Judd) , 1 9 ; Mar-
mora ( IX-9-1952 , J. McAlpine), 1 9 ; Ottawa ( V I I - 2 -
1947, G. Shewell), \$.
Quebec: Lac Bernard (VIII -7 -1938 , G. Shewell), 1 9 ;
Mt. Albert (VII-24-1954, J. Martin), 2 9 9 .
I M M A T U R E S P E C I M E N S E X A M I N E D . ? 3 2 f rom 6
localities:
Michigan: Douglas Lake (collected and reared by C.
Berg) ; Cheboygan County (collected and reared by C.
Berg); Presque Isle County (collected and reared by
C. Berg); Washtenaw County (collected and reared by
C. Berg).
Minnesota: Rapid River Logging Camp, Hubbard County
(collected and reared by D. L. Deonier); 1.3 miles south of
main entrance, Itasca State Park (collected and reared by
D. L. Deonier).
REMARKS.?The one external character of most
utility for rapid determination is the median facial
crease distinct only in anteroventral view.
Adult. Habitats recorded: leaves of Potamogeton
nodosus, P. natans, P. amplifolius, and Nuphar
advena.
No data is available on the adult stadium. On food
habits, I observed a female rapidly probing a dead,
fungus-covered chironomid. In other behavioral
observations, one male became aware of a second
male about 6 cm away and advanced upon it, then,
when both were only a few millimeters apart, the
second male jumped and grasped the thorax of the
protagonist as if to mount. A male advanced actively
upon two other specimens, and, in another obser-
vation, a smaller male scissored its wings after facing
a larger male, and then the larger male flew across
and rapidly grasped the thorax of the smaller one.
NUMBER 6 8 43
On another occasion, two females "faced off' and
began to slowly approach each other. This agonistic
behavior of H. bergi extended to its enemies. In
several observations made at Squaw Lake, Itasca
State Park, H. bergi rapidly approached and re-
treated from water spiders repeatedly.
In the only example of epigamic behavior observed,
a male and female stood with faces touching for
several seconds before the wind disturbed the meet-
ing. I captured two females after they oviposited on
sepals of Potamogeton natans. Berg (1950) found
an egg mass on a stipule of this species.
Egg. The incubation period for 24 of 36 eggs laid
by two females was four days.
Larva. All of the larvae that eclosed from the eggs
died except one, which was a late third instar when
it died 14 days later. According to Berg (1950), the
larvae mine nearly exclusively in stems and petioles
where they can be readily located only when the
epidermis over the longitudinal mine collapses.
Puparium. The puparia often bulge partially
through the epidermis of stems and petioles. Berg
(1950, p. 390) stated that prior to pupariation "the
larva provides for emergence of the adult by cutting
a U-shaped incision in the epidermis of the stem."
The incision is always made in the epidermis cover-
ing the future puparial operculum.
Host plants. Berg (1950) found larvae and
puparia in Potamogeton natans, P. richardsonii, and
P. zosteriformis. I reared an adult from a puparium
in Zizania aquatica. This host plant grew near a
stand of P. richardsonii. Judd (1964) recorded one
specimen of H. bergi from an emergence trap float-
ing near P. amplifolius.
Parasites. Berg reared three species of Braconidae?
Ademon niger, Chorebidea sp., and Dacnusa sp.?
and one of Diapriidae?Trichopria columbiana?
from puparia of H. bergi.
Hydrellia bilobifera Cresson
FIGURES 2, 12, 14, 50, 72, 73, 75, 85, 91, 109
Hydrellia bilobifera Cresson, 1936, p. 262; 1942, p. 78;
1944b [at least in part], pp. 170-175.?Deonier, 1964, p.
115; 1965, p. 500 [ecology].?Wirth, 1965, p. 744 [catalog
listing].
DIAGNOSIS.?Palpus moderate yellow; 5-8 aristal
rays; face planate; vertex index 5.4-6.8; male
abdomen prominently bilobate posteriorly; female
cere us about 3.0 times as long as wide; antennal
segment 3 at least partly moderate yellow; apicodor-
sal antennal prominent. Male length 1.63-2.04 mm;
female 1.70-2.31 mm. Male terminalia as in Figure
50; female as in Figure 72.
HEAD.?Face planate, light yellowish brown, yel-
lowish gray, or light gray; 5-6 pfa; epistomal index
1.7-2.2; mesofacial index 2.4-2.7; vertex index 5.4-
6.8; A-index 1.8-2.8; ocular index 5.3-6.2. Palpus
moderate yellow; 6-8 aristal rays; apicodorsal an-
tennal prominent; antennal segments 1 and 2 dark
brown, segment 3 at least partly moderate yellow;
fronto-orbital area moderate olive-brown; frontal
vitta semiglossy dark gray; most of parafrontale dark
brown; 14-17 postoculars.
THORAX.?Ppn light gray; mesonotum semiglossy
dark brown; 3 adc and 2 pdc; pleuron light gray
except light brown around metathoracic spiracle;
legs mostly moderate yellow except light-gray
pruinose femora. Wing length 1.79-1.87 mm; veins
light brown; 6-7 setae on basal end of costa; 5-8
dorsal and 9-11 anterior interfractural costals;
costal-section ratios: II:I 2.0-3.2; III:IV 2.5-3.2;
V: IV 3.0-3.5; Mi *
 2 index 1.2-1.7.
ABDOMEN.?Terga semiglossy dark gray medially,
light gray laterally and ventrally. Male terminalia:
median third of posterior margin of sternum 5 deeply
recessed and congruent with distiphallus; antero-
lateral margin of sternum 5 rounded to a posteriorly
directed notch; posterior third of copulobus dis-
tinctly concave in relation to sagittal plane (arcuate
and digitiform); copulobus verrucate laterally and
with a compact terminal fascicula of 5-7 strong
macrochaetae projecting anteriad to one-third of
copulobus length. Postgonite completely concealed;
distiphallus expanded proximally and bicarinate
ventrally on proximal two-thirds; most of basiphallus
exposed ventrally. Anterior margin of fused surstyli
concave medially; B-index about 1.6. Syntergum
9-f-10 prominently bilobate posteriorly. Female ter-
minalia: tergum 8 longer than sternum 8; cercus
setose mostly dorsally, digitiform and 3.0 times as
long as wide (lateral view). Segment 7 with group
of 4 seriated setae laterally on posterior margin;
median spermatheca cupuliform and nearly as long
as cercus.
EGG.?Length 0.47-0.64 mm; maximum breadth
0.14-0.18 mm. Chorion (Figure 75) white or light
yellowish gray, with inconspicuous, frequently
anastomosing longitudinal ridges; wide spaces be-
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
tween ridges smooth (in contrast to Figure 78).
Micropylar protuberance infundibulate, visible in
dorsal view; end opposite micropylar protuberance
with distinctly lacunose papilla.
FIRST-INSTAR LARVA.?Length 0.35-0.75 mm; max-
imum breadth 0.10-0.15 mm. Clypeal arch dis-
tinctly more angular at level of cheliform spot in
lateral view than in third-instar larva; frontoclypeal
length not measured. Very similar to H. discursa
(Figure 80), especially in having supraspiracular
protuberance with central spinous seta; dorsal setae
and spinules apparently absent. Newly eclosed larva
translucent, light yellowish gray; late first-instar larva
translucent, with greenish-yellow tinge.
SECOND-INSTAR LARVA.?Length 1.00-3.75 mm;
maximum breadth 0.20-0.50 mm. Frontoclypeal
length not measured. Dorsal setae absent; dorsal
spinules restricted to intersegmental grooves on
abdominal segments 1-7; abdominal segment 8 with
3-4 annuli of spinules and 1 conspicuous dorso-
lateral seta (perhaps supraspiracular spinous seta of
first-instar larva). Body translucent, with light-green
tinge.
THIRD-INSTAR LARVA.?Length 2.17-5.85 mm; max-
imum breadth 0.30-0.85 mm. Frontoclypeal length
0.45?0.52 mm; ventral phragmatal ramus mostly
hyaline (Figure 91). Ventral frontoclypeal index
2.8-3.2; phragmatal index 1.0-1.2; bifurcation
index 4.2-4.5; clypeal-arch index 1.7?2.0. Cly-
peal arch gradually sloping at about 35? in rela-
tion to lower frontoclypeal margin. Mouth-hook beak
and base lengths subequal; maximum mouth-hook
base thickness about 2.2 times that of beak. Prothorax
and mesothorax sparsely spinulose ventrally and
laterally; abdominal segment 8 without ventral,
transverse row of setulae, but with 3-5 annuli of
spinules. Body translucent, with light-green tinge.
PUPARIUM.?Length 3.10-4.25 mm; maximum
breadth 0.75-1.00 mm; fusiform (Figure 109).
Puparial length:minimum breadth 12.0-15.0; maxi-
mum breadth: minimum breadth 2.4-3.2; anal-plate
index 1.8-2.4. Prothoracic end semicircular in ven-
tral view; head-lobe scar obovoid, nearly as long as
prothorax; prothorax moderately rugulose ventrally
and laterally; abdominal segment 8 ornamented as
in third-instar larva; anal plate subrectangular, with
anterior margin straight or slightly convex. Empty
puparium translucent, light yellowish brown; early
pupa light green.
TYPE.?Holotype male, ANSP 6531.
TYPE-LOCALITY.?Atherton, Missouri (VI-no year,
C. F. Adams).
ADULT SPECIMENS EXAMINED.?349 (210cfcf,
139? ? ) from 40 localities:
California: Buena Park (V-19-1944, A. Melander), 1 3 ;
Davis (VIII-13-1954, A. Grigarick), 1 3 , (X-21-1954, A.
Grigarick), 33 3 , 19 , (VII-26-1955, W. Lange), 1 3 ,
(IX-10-1957, A. Grigarick), 2 9 9 , (IX-17-1957, A. Gri-
garick), 2 3 3 ; Maxwell, Colusa County (VIII-12-1954, A.
Grigarick), 11 3 $, 7$ 5 , (VIII-3-1955, W. Lange), 1$ ;
Putah Canyon, Yolo County (X-20-1954, A. Grigarick),
3 3 3 ; Sacramento (IX-3-1957, A. Grigarick), 33 3 .
District of Columbia: Washington (VIII-13-1923, W.
McAtee), 1 3 .
Iowa: Banner Mine Area, Warren County, 41?26.4' N,
93?33.8' W (VIII-7-1960, D. L. Deonier), 1 3 ; Ledges
State Park, Boone County (VII-28-1954, J. Laffoon), 1 9 ;
Ledges State Park, Boone County, south pond at 41?58.8'
N, 93?53.5' W (VII-10-1960, D. L. Deonier), 19 ; Lewis
and Clark State Park, Monona County 42?2' N, 96? 10' W
(VI-6-1960, D. L. Deonier), 1 9 ; Little Wall Lake, Hamil-
ton County (VIII-27-1960, D. L. Deonier), 2 3 3 , 1 9 ;
Mclntosh Woods State Park, Clear Lake (IX-10-1961,
D. L. Deonier), 2$ 3 ; Pilot Knob State Park, Hancock
County (VI-15-1960, D. L. Deonier), 1 $ ; Siewer's Springs
State Park near Decorah (IX-9-1961, D. L. Deonier)
83 3 ; Spring Lake, Greene County (IX-19-1961, D. L.
Deonier), 4 3 3 , 79 9 ; Springbrook State Park, Guthrie
County (IX-2-1955, J. Laffoon), 1 3 , (VII-19-1960,
D. L. Deonier), 2$ 3 .
Kansas: Kansas State College Natural History Area near
Pittsburg (VI-4-1963, D. L. Deonier), 28 3, 19 ; Kansas
University Natural History Reservation near Lawrence,
Douglas County (VI-7-1963, D. L. Deonier), 35 3 3 ,
219 9 , (VI-10-1963, D. L. Deonier), 1 3 , 29 9 ; Leaven-
worth County Lake (VIII-27-1962, D. L. Deonier), 213 3 ,
17 9 9 ; Marais des Cygnes Wildlife Refuge, Linn County
(IX-5-1961, D. L. Deonier), 1 3 ; 1 mile northeast of
Lawrence, Douglas County, in sandpits (VI-2-1963, D. L.
Deonier), 1 3-
Michigan: Bessey Creek, Cheboygan County (VIII-14-
1941, C. Berg), 19 ; Cheboygan County (VII-26-1946, C.
Berg) 1 9 , (XI-26-1946, C. Berg), 23 3 , (VIII-11-1947,
C. Berg), 1 3 .
Minnesota: Biological Station, Itasca State Park (VI-
26-1963), D. L. Deonier), 1 3 ; Bohall Lake, Itasca State
Park (VI-25-1963, D. L. Deonier), 2 3 3 , 19 (VIII-17-
1963, D. L. Deonier), 23 3 ; Chambers Creek, Itasca State
Park (VII-18-1963, D. L. Deonier), 1 3 ; Headwaters of
La Salle Creek, Itasca State Park (VIII-6-1963, D. L.
Deonier), 5 3 3 ; Mississippi River near north entrance,
Itasca State Park (VII-9-1963, D. L. Deonier), 7 3 3 ,
89 9 ; Two Island Lake, Itasca State Park (VI-28- 1963,
D. L. Deonier), 1 3 , (VIII-8-1963, D. L. Deonier), 1 3 ;
6.5 miles east of Waubun (VIII-18-1963, D. L. Deonier),
5 9 9 .
NUMBER 6 8 45
Mississippi: Vancleave Road, Jackson County, 30? 28' N.
88?43' W (VI-15-1962, D. L. Deonier), 19 .
North Carolina: Highlands Biological Station, Highlands,
Macon County (VI-15-26-1968, D. L. Deonier), 20 5 5 ,
319 9.
Ontario: Marmora (VIII-25-1952, E. H. Smith), 15 ,
(IX-9-1957, J. McAlpine), 1 $.
South Carolina: Jamestown (VII-2-1953, M. Wheeler),
1*.
Tennessee: Brewer's Bar Ditch, Reelfoot Lake, 36?27.1,
N, 89?21.2' W (VIII-10-1962, D. L. Deonier), 45 5 ,
2 9 2 ; Dale Hollow Reservoir, 6 miles south of Byrdstown,
Pickett County (VIII-19-1962, D. L. Deonier), 4 5 5 ; Mil-
ler's Camp, Reelfoot Lake, 36?24.3' N, 89?20' W (VIII-
12-1962, D. L. Deonier), 15 5 5 , 152 9 ; Samburg, Reel-
foot Lake, 36?22.9' N, 89?21.3' W (VIII-10-16-1962, D. L.
Deonier), 30 5 5, 112 2-
Texas: Galveston (IV-11-1952, M. Wheeler), 15 , (IX-
13-1953, M. Wheeler), 1 5 ; Kerrville (111-17-1955, W.
Wirth), 15 .
IMMATURE SPECIMENS EXAMINED.?296 from 7
localities:
Iowa: Siewer's Springs State Park near Decorah (col-
lected and reared by D. L. Deonier); Spring Lake, Greene
County (collected and reared by D. L. Deonier).
Kansas: Kansas University Natural History Reservation
near Lawrence, Douglas County (collected and reared by
D. L. Deonier); 1 mile northeast of Lawrence, Douglas
County (collected and reared by D. L. Deonier).
North Carolina: Highlands Biological Station, Highlands,
Macon County (collected and reared by D. L. Deonier).
Tennessee: Brewer's Bar Ditch and Samburg, Reelfoot
Lake (collected and reared by D. L. Deonier).
REMARKS.?Determination should be confirmed by
examination of the terminalia. The H. bilobifera
group includes this species, H. ascita, H. gladiator,
H. harti, H. discursa, and H. trichaeta. These sibling
species constitute the most well-defined group in
Hydrellia. There is some probability that I misdeter-
mined some females of this group because I was
unable to dissect the genitalia of every specimen
examined.
Adult. Habitats recorded: floating parts of
Potamogeton nodosus, P. gramineus, P. epihydrus,
Zizania aquatica, Oryza sativa, and Nymphaea
tuberosa; mats of Lemna minor; limnic wrack. I
collected one male by the lighted-receptacle method.
At the Kansas University Natural History Reser-
vation, I observed one female catching an adult
chironomid, another female grasping an adult
chironomid, and yet another one probing a specimen
of Podura aquatica that it held with its fore tarsi.
The gut contents of ten adults from this locality con-
sisted of greenish-yellow, granular material, the
identity of which could not be determined. At this
locality also, several pairs of adults approached each
other until their antennae touched. These meetings
usually culminated in one adult jumping rapidly
away or attempting to mount the other from the side.
At Itasca State Park, Minnesota, I watched a male
approach a female H. trichaeta; the latter struck
aggressively at the male. One adult female emerged
from a host plant collected at Highlands, North
Carolina, mated, oviposited, and lived a total of 18
days while being fed 5 percent sucrose solution.
Egg and larva. I found eggs on the floating leaves
of Potamogeton nodosus at Spring Lake, Greene
County, Iowa, and on those of P. epihydrus at
Ravenel Lake, Highlands, North Carolina. I observed
deposition of 10 eggs at the latter locality. The
incubation period ranged from 4 to 5 days. Eggs
from the Iowa locality incubated 2 days after collec-
tion. The first larval stadium ranged from 5-10
days; the second and third, from 5 to 14 and 4 to 9
days, respectively, at laboratory room temperatures
for 20 specimens.
Puparium. The puparial phase ranged from 6 to
15 days in the laboratory for 37 specimens. In one
case, one larva pupariated in the open water of a
culture dish. The adult did not emerge.
Host plants. I found larvae and puparia only in
Potamogeton nodosus, P. epihydrus, and P. gramineus
in the field, but field observations indicate that
they infest any Potamogeton with floating leaves.
Grigarick reared Hydrellia bilobifera from Zanni-
chellia palustris in Davis, California, during his re-
search on Hydrellia griseola. It is uncertain whether
this represents a larval host or an incidental puparial
host. My survey of potential host plants included
also Potamogeton pectinatus, P. natans, P. spirillus,
Mimulus glabratus, Alisma sp., Heteranthera dubia,
Ceratophyllum demersum, Sagittaria australis, Jus-
siaea repens, Nasturtium officinale, Eleocharis palus-
tris, Polygonum amphibium, Elodea canadensis,
Zannichellia palustris, and Echinodorus cordifolius.
Parasites. Fungi of the genus Stigmatomyces
(Laboulbeniales) killed several specimens in two lots
of larvae being reared in the laboratory. It is com-
monly stated that these insect parasites apparently
do not injure the host in any way. Every larva
infested with this parasite in my laboratory died. The
thalli first appeared as small dark brown spots on the
46 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
cuticle. The hyphae appeared to penetrate just
through the cuticle.
I reared Chorebidella bergi, Aphanta sp. (prob-
ably new), and Opius sp. (all Hymenoptera:
Braconidae) from puparia of this species. The time
from pupariation to emergence of the hymenopterans
ranged from 12 to 23 days. Five hymenopterans
emerged from 42 puparia reared or collected.
Hydrellia biloxiae, new species
FIGURES 3 4 , 8 1 , 101, 119
DIAGNOSIS.?Palpus moderate yellow; 6-10 aristal
rays; vertex index 4.4-5.0; abdominal terga 2^4
velvety purplish black medially; thoracic pleuron
moderate olive-brown with bronze reflections; frons
(except light brown around fronto-orbital sockets)
velvety black. Male length 1.22-1.45 mm; female
1.36-1.56 mm. Male terminalia as in Figure 34.
HEAD.?Face light yellowish brown; 4-5 pfa;
epistomal index 1.1-1.5; mesofacial index 2.0-2.4;
vertex index 4.4-5.0; A-index 2.0-2.4; ocular index
12.0-14.0. Palpus moderate yellow; 6-10 aristal
rays; antennal segment 1 pale yellow medially, 2
velvety black, and 3 moderate brown with moderate
orange-yellow splotches; frons (except light brown
around fronto-orbital sockets) and occiput velvety
black; 13-16 postoculars.
THORAX.?Ppn, mesoscutum posterior to transverse
sulcus, and scutellum semiglossy moderate brown;
mesoscutum anterior to transverse sulcus semiglossy
dark brown; notopleuron and postalar wall mod-
erate olive-brown; 3-4 adc and 2-3 pdc; pleuron
moderate olive-brown with bronze reflections; coxae
light gray laterally, moderate yellow anteriorly and
ventrally; trochanters, tarsi, and distal fifth of tibiae
moderate yellow; remaining areas of legs light-gray
pruinose. Wing length 1.39-1.70 mm; veins light
brown; 6-7 setae on basal end of costa; 5-7 dorsal
and 7-10 anterior interfractural costals; costal-
section ratios: 11:1 1.4-2.2; I I I : IV 2.6-3.5; V:IV
3.2-3.6; Mi *
 2 index 1.6-2.2.
ABDOMEN.?Terga 2-4 velvety purplish black
medially, the sides, ventral lobes, and tergum 5 dark
brown. Male terminalia: median third of posterior
margin of sternum 5 concave and congruent with
distiphallus; anterolateral margin of sternum 5 about
110? angular; copulobus acute posteriorly and irreg-
ularly setose. Postgonite bent anteriad and post-
gonite uncus slightly laterad; distiphallus tapering
to ovoid, ventrally transversely lammellate apex.
Anterior margin of fused surstyli ovoid; B-index
about 3.5; C-index 2.0-2.5.
FIRST-INSTAR LARVA.?Length 0.52 mm. Feeding
apparatus lighter and less heavily sclerotized, but of
same general shape as that of third-instar larva.
Body translucent, yellowish gray.
SECOND-INSTAR LARVA.?Length 2.75-3.10 mm;
maximum breadth 0.30-0.40 mm. Frontoclypeus not
measured. Abdominal segment 8 with bidentate and
tridentate spinules dorsally.
THIRD-INSTAR LARVA.?Length 3.40-4.00 mm;
maximum breadth 0.40-0.50 mm. Frontoclypeal
length 0.35-0.40 mm; cheliform spot touching
clypeal arch margin (Figure 101). Ventral fronto-
clypeal index 1.5-2.0; phragmatal index 0.75-0.85;
bifurcation index 2.5-2.7; clypeal-arch index 1.5-
1.8. Clypeal arch sloping upward very slightly from
subrectangular prominence dorsal to labial-gland
duct opening. Mouth-hook beak and base lengths
subequal; maximum mouth-hook base thickness
about 1.2 times that of beak. Body opaque, yellowish
gray.
PUPARIUM.?Length 2.40-3.25 mm; maximum
breadth 0.60-0.80 mm; subfusiform (Figure 116).
Puparial length: minimum breadth 16.0-19.0; maxi-
mum breadth: minimum breadth 3.6-4.2; anal-plate
index 1.8-3.2. Prothoracic end semicircular in ven-
tral view; maximum puparial breadth in metathorax;
prothorax and mesothorax very sparsely spinulose;
prothorax distinctly transversely rugulose in ventral
view; head-lobe scar subcircular to transversely
elliptical; dorsal spinules restricted to thorax and
abdominal segment 8; caudal segment without ven-
tral, transverse row of setulae, but with 3-4 annuli
of spinules; anal plate subovoid, with anterior mar-
gin slightly convex. Empty puparium translucent,
light yellowish brown.
TYPE.?Holotype male, USNM 70528.
TYPE-LOCALITY.?Gravel pit near U. S. Highway
90, sec. 20, T.7 south, R.5 west, Jackson County,
Mississippi (VI-17-1962, D. L. Deonier).
PARATYPES.?84 (290*0*, 55? $ ) from 4 locali-
ties:
Mississippi: Gravel pit near U. S. Highway 90, sec. 20,
T. 7 south, R. 5 west, Jackson County (VI-17-1962,
D. L. Deonier), 6$S, 4 9 $ (USNM and DLD); Lake
Shady, 7 miles west of Hattiesburg, Lamar County (VII-
11-1962, D. L. Deonier), 1$ (ISC); Lake Shelby State
NUMBER 6 8 47
Park, Forrest County (VII-24-1962, D. L. Deonier), 2$ $,
49 9 (USNM); Vancleave Road, Jackson County, 30?28'
N, 88?43' W (VI-15-1962, D. L. Deonier), 18$ $, 47 9 9
(USNM, ISC, and DLD).
IMMATURE SPECIMENS EXAMINED.?23 specimens
from 2 localities:
Mississippi: Gravel pit near U. S. Highway 90, sec. 20,
T. 7 south, R. 5 west, Jackson County (collected and reared
by D. L. Deonier); Vancleave Road, Jackson County, 30?28'
N, 88C43' W (collected and reared by D. L. Deonier).
REMARKS.?Adult. Habitats recorded: leaves of
Juncus repens, Hydrochloa caroliniensis, and Nym-
phaea odorata. In one locality near Vancleave Road,
Jackson County, Mississippi, from 23 to 28 June and
from 7 to 10 July I collected 54 and 68 adults,
respectively, in three Grigarick model floating traps
(25 cm diameter).
At the above-mentioned locality, an adult probed
a partially dessicated chironomid larva on a leaf of
Nymphaea odorata for about 5 minutes. When it
left the larva and touched its proboscis to the water,
another adult approached the larva. The first adult
drove off the intruder by scissoring its wings and
rushing rapidly toward it. While feeding, the speci-
men pushed its body forward and upward rhyth-
mically. Shortly afterward, in another case, I observed
the same behavior and sequence of events, except for
a more vigorous defense against the intruder. Here,
too, I saw a ctenid spider grasping an adult and a
smaller undetermined spider leap at an adult that
landed near its resting site.
I did not observe oviposition, but some females
among several adults on leaves of Juncus repens and
Eleocharis wolfii protracted their ovipositors and
touched them to various places on the leaves.
Larva and puparium. Mines in stolons and the
presence of agromyzid leaf miners hampered the dis-
covery of the larvae. For four specimens, the time
from about the middle of the first larval stadium to
adult emergence ranged from 19 to 21 days. One
larva passed the third larval stadium in 12 days.
One third-instar larva mined through a 4-cm length
of stolon into a leaf sheath. I did not measure the
puparial phase. Adults may encounter occasional
difficulty in escaping from mines, for I found two
adults that had emerged from the puparia but re-
mained trapped in the mines. One of these adults
was trapped about 5 mm from the puparium in a
stolon, without an escape slit, growing about 0.75 m
below the surface. In some cases, the escape slit in
the stolon mine was over the operculum, and in
others it was anterior to it. Evidently, the larvae
made escape slits before pupariation.
Host plants. I found larvae and puparia only in
Juncus repens. At the Vancleave Road locality, I
estimated the infestation to be light to moderate.
In addition to /. repens, I examined Orontium
aquaticum, Zizaniopsis miliacea, Polygonum seta-
ceum, Rhynchospora cymosa, Eleocharis tuberculosa,
E. wolfii, E. mammillata, and Cyperus strigosus.
Hydrellia borealis Cresson
FIGURE 53
Hydrellia borealis Cresson, 1944b, pp. 164, 171, 172.?
Wirth, 1965, p. 744 [catalog listing].
DIAGNOSIS.?Palpus dark brown (infrequently
partly moderate orange-yellow); 7-8 aristal rays;
vertex index 4.5-9.5; body length:wing length 0.8-
0.9; apicodorsal antennal prominent; adc more or
less uniordinate; thoracic pleuron light gray; male
hind tibia expanded. Male length 1.96-2.21 mm;
female 1.79-2.13 mm. Male terminalia as in
Figure 53.
HEAD.?Face silky light yellowish brown; 4-6 pfa;
1-3 sfa; epistomal index 1.0-1.4; mesofacial index
1.7-2.3; vertex index 4.5-9.5; A-index 1.5-2.0;
ocular index 6.8-8.0. Palpus dark brown (infre-
quently partly moderate orange-yellow); 7-8 aristal
rays; apicodorsal antennal prominent; antenna and
frontal vitta brown; most of parafrontale olive-
brown; lower third of postocular area light gray,
upper two-thirds light brown; 13?16 postoculars.
THORAX.?Ppn and mesonotum dark gray; 3-4
adc (none macrochaetous) and 2 pdc; pleuron light
gray; legs light-gray pruinose except moderate-yellow
trochanters; male hind tibia slightly expanded (as
in Figure 8). Wing length 2.13-2.47 mm; veins light
brown; 5-7 setae on basal end of costa; 6-8 dorsal
and 6-9 anterior interfractural costals; costal-
section ratios: II:I 2.3-2.9; III:IV 2.8-3.3; V:IV
3.5-4.0; Mi
 + 2 index 1.2-1.8.
ABDOMEN.?Terga dark gray. Male terminalia:
median third of posterior margin of sternum 5 deeply
concave; anterolateral margin of sternum 5 rounded
through 110?-115? angle; copulobus slightly convex
posteriorly and regularly setose, with about 7-8
48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
seriated setae on medial margin. Postgonite bent
strongly anteromediad and postgonite uncus slightly
laterad; distiphallus slightly expanded (oblong) on
distal two-thirds. Anterior margin of fused surstyli
narrowly cleft medially to about one-third of length
of structure; B-index about 1.0; C-index 0.5-0.7.
TYPE.?Holotype male, type no. 6653, which may
be at the Academy of Natural Sciences of Philadel-
phia in an unlabelled condition, but is probably
either lost or in the Melander Collection.
TYPE-LOCALITY.?Mer Bleue, Ottawa, Canada
(VII-2-1938, A. L. Melander).
SPECIMENS EXAMINED.?11 (6c? d\ 5 $ $ ) from 9
localities:
Alaska: King Salmon, Naknek River (VIII-1-1952, W.
Mason), 1 $.
British Columbia: Atlin, 2,200 feet (VII-6-1955, B.
Gibbard), 1 $ ; Brilliant (VIII-3-1947, H. Foxlee), 1$.
Michigan: Dickinson County (VII-1-1955, R. Dreis-
bach), 1$ .
Nebraska: Crete (VII-3-1960, W. Rapp), 1$.
Ontario: Ottawa (VI-1-1951, J. McAlpine), 1 $, (VII-
14-1952, G. Shewell), 1$, (VII-2-1958, J. Vockeroth),
\$.
Quebec: Beech Grove (VII-18-1951, J. McAlpine), 1$ .
Washington: 23 miles west of Republic (VII-14-1960,
collector unknown), 1 $ .
Wyoming: Lander (VIII-16-1950, M. Wheeler), 1$ .
REMARKS.?I did not see the holotype; thus, I
based my determinations entirely upon Cresson's
description and key and upon specimens that had
been compared with the holotype.
Hydrellia caliginosa Cresson
FIGURES 19,98, 111
Hydrellia caliginosa Cresson, 1936, pp. 257-258; 1942, p.
78; 1944b, pp. 170, 175.?Berg, 1950, pp. 375, 376, 378,
384-385, 392-393, 396 [biology].?Wirth, 1965, p. 744
[catalog listing].
DIAGNOSIS.?Palpus moderate yellow; 8-10 aristal
rays; vertex index 4.5-5.5; thoracic pleuron light
gray; costal section I I I : IV 2.3-2.5; fronto-orbital
area and frontal vitta strong yellowish brown; most
of parafrontale velvety black. Male length 2.11-2.60
mm; female 1.97-2.47 mm. Male terminalia as in
Figure 19.
HEAD.?Face light yellowish brown or light gray;
4-6 pfa; epistomal index 1.2-1.6; mesofacial
index 1.6-2.2; vertex index 4.5-5.5; A-index 0.8-
2.0; ocular index 4.2-7.0. Palpus moderate yellow;
8-10 aristal rays; prementum glossy brownish black;
antenna very dark brown; fronto-orbital area and
frontal vitta strong yellowish brown; most of para-
frontale velvety black; 14-17 postoculars.
THORAX.?Ppn and notopleuron strong yellowish
brown; remainder of mesonotum usually moderate
brown; 3-4 adc and 2-4 pdc; pleuron light gray;
1-2 basal coxals (1 macrochaetous) ; legs light-gray
pruinose except dark-brown tarsi. Wing length 2.30-
2.75 mm; veins moderate brown; 8-9 setae on basal
end of costa; 8-10 dorsal and 9-12 anterior inter-
fractural costals; costal-section ratios: I I : I 2.2-2.6;
I I I : IV 2.3-2.5; V:IV 2.8-3.3; Mx ? 2 index 1.0-1.3.
ABDOMEN.?Terga moderate brown medially and
anterolaterally, light gray elsewhere. Male terminalia:
median third of posterior margin of sternum 5 deeply
concave; anterolateral corner of sternum 5 right-
angled (90?-100?); posterolateral angle of
copulobus roundly 45? from the lateral corner; the
medial margin broadly notched; copulobus irregu-
larly setose. Postgonite bent mediad and then sharply
anteriad; with peninsulate projection along lateral
margin of copulobus; postgonite uncus distinctly
clawlike; distiphallus tapering to mucronate apex.
Anterior margin of fused surstyli truncate; B-index
about 0.6; C-index 0.1-0.2.
THIRD-INSTAR LARVA.?Body of larva not exam-
ined. Frontoclypeal length 0.46-0.59 mm (Figure
98). Ventral frontoclypeal index 5.0-5.5; phrag-
matal index 1.0-1.2; bifurcation index 3.6-3.8;
clypeal-arch index 1.6-1.7. Clypeal arch sloping at
about 20? in relation to lower frontoclypeal margin.
Mouth-hook beak longer than base; maximum
mouth-hook thickness about 1.5 times that of beak;
mouth-hook without light spot.
PUPARIUM.?Length 3.30-4.15 mm; maximum
breadth 0.95-1.40 mm; subfusiform, suddenly taper-
ing at segment 7 (Figure 111). Puparial length:
minimum breadth 18.0-20.0; maximum breadth:
minimum breadth 6.0-8.0; anal-plate index 2.5-
3.0. Prothoracic end semicircular in ventral view;
head-lobe scar circular; maximum puparial breadth
5.5-6.0 times diameter of head-lobe scar and 2.3
times or less than maximum prothoracic breadth;
prothorax with fewer rugulae ventrally than H.
itascae; lateral spinules conspicuous, but no large
spinules in annuli of abdominal segment 8 as in
H. itascae (Figure 107) ; anal plate subovoid, with
NUMBER 68 49
anterior margin slightly convex. Empty puparium
moderate brown.
TYPE.?Holotype male, ANSP 6528.
TYPE-LOCALITY.?New Mill Pond, Mount Desert
Island, Maine (VII-25-1935, W. Procter).
ADULT SPECIMENS EXAMINED.?-66 (40cT d1,
2 6 ? ? ) from 11 localities:
Alaska: Matanuska Valley (VI-27-1950, C. Berg), 26 6,
(VII-6-1950, C. Berg), 36 6, (VII-20-1950, C. Berg),
86 6, 59 9, (VIII-5-1950, C. Berg), 13 .
Idaho: Priest Lake, Tule Bay (VIII-19-1919, A. L. Me-
lander), 1 6 ?
Maine: Mount Desert Island (VII-12-1935, W. Procter),
IS, (VII-25-1935, W. Procter), 2 6 6, 39 9.
Michigan: Cheboygan County (VII-21-1947, C. Berg),
43 6, 79 9, (VIII-7-1947, C. Berg), 43 5 , 5 9 9.
Minnesota: Eaglenest (IX-1-1958, W. Balduf), 1 3 ;
Rapid River Logging Camp, Hubbard County (VII-2-
1963, D. L. Deonier), 1 6-
Montana: St. Marys (VII-22-1935, A. L. Melander),
16-
Quebec: Mt. Albert (VII-24-1954, J. Martin), 11 6 6,
39 9 ; Rupert House (XI-17-1949, E. LeRoux), 19.
Wyoming: Kemmerer (VIII?14?1950, collector un-
known), 16, 1 9 ; Yellowstone National Park, 4,750 feet
(VII-20?no year, collector unknown), 19-
IMMATURE SPECIMENS EXAMINED.?10 from 2
localities:
Alaska: Matanuska Valley (collected and reared by C.
Berg).
Minnesota: Rapid River Logging Camp, Hubbard County
(collected and reared by D. L. Deonier).
REMARKS.?Using the key of Cresson (1944b,
p. 175), one encounters difficulty in the critical
couplet mainly in the ocellar: postocellar length ratio.
I found this character unreliable.
Adult. Habitat recorded: leaves of Nuphar advena.
Host plants. Berg (1950) reared one adult from a
puparium in a leaf of Potamogeton praelongus. In
personal communication with Berg, he indicated he
had reared several adults from an unspecified spe-
cies of Potamogeton in Matanuska Valley, Alaska.
I reared one adult from a puparium in a leaf of
P. richardsonii collected at Rapid River Logging
Camp, Hubbard County, Minnesota.
Hydrellia cavator, new species
FIGURE 54
DIAGNOSIS.?Palpus moderate yellow; 4-6 aristal
rays; parafrontale velvety black; wing length 1.53-
2.13 mm; antennal segment 3 moderate yellow;
mesoscutum and abdominal dorsum dark brown
medially; thoracic pleuron and side of abdomen light
gray. Male length 1.63-1.70 mm; female 1.45-1.84
mm. Male terminalia as in Figure 54.
HEAD.?Face light gray; 5-6 pfa; epistomal
index 1.5-1.8; mesofacial index 2.1-2.5; vertex in-
dex 4.5-4.8; A-index 1.6-2.2; ocular index 6.5-12.0.
Palpus moderate yellow; 4-6 aristal rays; antennal
segments 1 and 2, fronto-orbital area, and frontal
vitta moderate brown; antennal segment 3 moderate
yellow; most of parafrontale velvety black; 13-16
postoculars.
THORAX.?Ppn and notopleuron moderate olive-
brown; remainder of mesonotum dark brown; 3-4
adc and 2 pdc; pleuron light gray; legs light-gray
pruinose except moderate-yellow tarsi. Wing length
1.53-2.13 mm; veins light yellowish brown; 6-7
setae on basal end of costa; 5-7 dorsal and 7-9
anterior interfractural costals; costal-section ratios:
I I : I 2.0-2.5; I I I : IV 2.6-3.5; V:IV 3.4-4.0; Mi ?
 2
index 1.2-1.7.
ABDOMEN.?Terga dark brown medially, light
gray laterally and ventrally. Male terminalia: median
third of posterior margin of sternum 5 concave;
anterolateral margin of sternum 5 truncate between
angles of about 120? and 90?-100?; copulobus acut-
angular posteriorly and irregularly setose. Postgonite
bent anterolaterad and postgonite uncus sharply
laterad; distiphallus uniform in breadth to truncate
and ventrally lamellate apex. Anterior margin of
fused surstyli smoothly convex; B-index about 3.0;
C-index 2.0-2.5.
TYPE.?Holotype female, AMNH (no. not used).
A male specimen originally intended to be the holo-
type was damaged in shipment. It was collected at
Biscayne Bay, Florida, by Mrs. A. T. Slosson.
TYPE-LOCALITY.?Lake Worth, Florida (collection
of Mrs. A. T. Slosson, Ace. 26226).
PARATYPES.?4 (lef, 3 ? $ ) from 4 localities:
Florida: Biscayne Bay (no date, Mrs. A. T. Slosson), 1 6
(AMNH); Gainesville (IV-23-1952, J. Vockeroth), 19
(CNC); Lake Worth (no date, Mrs. A. T. Slosson), 19
(AMNH); Lake Worth, bay shore (VIII-8-1951, W.
Wirth), 19 (USNM).
REMARKS.?I found some specimens of this species
in material determined as H. atroglauca. Hydrellia
atroglauca is much larger and has 8?10 aristal rays.
50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Hydrellia cessator, new species
FIGURE 59
DIAGNOSIS.?Palpus moderate yellow; 7-9 aristal
rays; vertex index 4.0-4.8; mesofacial index 1.5-
2.0; ocular index 6.0-7.0; thoracic pleuron mod-
erate olive-brown except mesokatepisternum and
mesomeron light gray. Male length 1.79-2.19 mm;
female 2.38-2.55 mm. Male terminalia as in Fig-
ure 59.
HEAD.?Face light gray or light yellowish brown;
3-6 pfa; 0-5 sfa; epistomal index 1.0-1.4; meso-
facial index 1.5-2.0; vertex index 4.0-4.8; A-index
1.5-2.0; ocular index 6.0-7.0. Palpus moderate
yellow; 7-9 aristal rays; prementum pale yellow;
antenna dark gray; fronto-orbital area and frontal
vitta moderate olive-brown; most of parafrontale
dark brown; 13?16 postoculars.
THORAX.?Ppn light gray; mesonotum moderate
brown; 4-5 adc and 2-3 pdc; pleuron moderate
olive-brown except mesokatepisternum and meso-
meron light gray; 1-2 basal coxals (1 macro-
chaetous); legs sparsely light-gray pruinose except
densely gray pruinose coxae and dark-brown tarsi.
Wing length 2.04-2.64 mm; veins moderate brown;
6-8 setae on basal end of costa; 6-8 dorsal and 8-11
anterior interfractural costals; costal-section ratios:
11:1 2.2-2.4; I I I : IV 2.4-3.3; V:IV 3.0-3.6; Mi ?
 2
index 1.2-1.4.
ABDOMEN.?Terga moderate brown medially and
light gray laterally. Male terminalia: median third
of posterior margin of sternum 5 deeply concave;
anterolateral margin of sternum 5 with 2 angles
(100? and 140?) ; copulobus acutangular posteriorly
and irregularly setose. Postgonite bent laterad and
postgonite uncus anteriad; distal half of distiphallus
tapering to acute and ventrally bicarinate apex.
Anterior margin of fused surstyli only slightly con-
cave; B-index about 0.6; C-index 1.0-1.2.
TYPE.?Holotype male, USNM 70529.
TYPE-LOCALITY.?Mississippi River, sec. 34, T.145
north, R.36 west, Clearwater County, Minneosta
(VII-8-1963, D. L. Deonier).
PARATYPES.?23 (7cT d", 16? $ ) from 6 localities:
Manitoba: Two miles north of Forrest (VII-19-1958, J.
Chillicott), 1$ (CNC).
Minnesota: Headwaters of La Salle Creek, Itasca State
Park (reared adults between V-15 and VI-7-1967, D. L.
Deonier), 4<J $, 79 9 (DLD); Mississippi River, sec. 34,
T.145 north, R.36 west, Clearwater County (VII-8-1963,
D. L. Deonier), 69 9 (ISC and USNM) ; Mississippi River
near north entrance, Itasca State Park (reared adults V I -
4-1967, D. L. Deonier), \$, 19 (DLD); Rapid River
Logging Camp, Hubbard County (VII-2-1963, D. L. Deo-
nier), 2$ (DLD); Sucker Lake at dam (reared adult V-
27-1967, D. L. Deonier), 19 (DLD).
REMARKS.?Adult. Habitats recorded: floating
leaves of Glyceria grandis, Zizania aquatica, and in
sedge meadow. The adults appeared to move about
very little on the leaves and for this reason they were
difficult to detect.
Host plants. Immatures were found in the sub-
merged leaves of Glyceria grandis during the late
winter and early spring of 1967 in Itasca State Park
and vicinity. Of the 12 specimens found in this
grass, two were second-instar larvae, six were third-
instar, and four were puparia. The larvae (probably
first- or second-instar) evidently overwinter in the
green, submerged leaves of this species along with
the larvae of H. ischiaca, H. morrisoni, and H.
nobilis . The second-instar larvae were collected in
the host plants on 30 April, 15 days after ice had
thawed in the lakes and while mean daily tempera-
tures were still in the 35-45?F range. Circumstantial
evidence indicates that G. grandis plants are also
normal summer hosts of the larvae. I also surveyed
Potamogeton epihydrus, P. praelongus, P. zosteri-
formis, Zizania aquatica, and Glyceria striata as
species of potential host plants.
Hydrellia columbata, new species
FIGURE 55
DIAGNOSIS.?Palpus dark brown; 5-7 aristal rays;
vertex index 4.6-6.0; mesofacial index 1.0-1.4; face
light gray, carinate only on upper two-thirds;
thoracic pleuron moderate olive-brown except lower
half of mesokatepisternum light gray; male mid tibia
expanded. Male length 1.53-1.99 mm; female 1.70-
2.09 mm. Male terminalia as in Figure 55.
HEAD.?Face light gray, carinate only on upper
two-thirds; 4-5 pfa; 5-7 sfa; epistomal index 0.9-
1.4; mesofacial index 1.0-1.4; vertex index 4.6-6.0;
A-index 1.8-2.2; ocular index 4.3-5.0. Palpus dark
brown; 5-7 aristal rays; antenna and most of frons
velvety dark brown; fronto-orbital area moderate
olive-brown; 13-15 postoculars.
THORAX.?Ppn and mesonotum moderate olive-
brown; 3-4 adc and 2-4 pdc; pleuron moderate
NUMBER 68 51
olive-brown except lower half of mesokatepistemum
light gray; legs moderate olive-brown pruinose ex-
cept dark-brown tarsi; male mid tibia expanded.
Wing length 1.62-1.87 mm; veins dark brown; 6-7
setae on basal end of costa; 5-7 dorsal and 7-10
anterior interfractural costals; costal-section ratios:
II:I 2.0-2.4; III:IV 2.5-3.0; V:IV 2.8-3.4; Mi
 + 2
index 1.3-1.8.
ABDOMEN.?Terga moderate olive-brown pruinose
over dark gray. Male terminalia: median third of
posterior margin of sternum 5 broadly and shallowly
concave; anterolateral corner of sternum 5 roundly
right-angled (90?-100?) and projecting distinctly
more laterad than surstyli; copulobus unusually long
with end bent posteromediad, with terminal cluster
of setae, and with 5-7 setae anteriorly. Postgonite
long and bent anteromediad; postgonite uncus dis-
tinctly long and hooklike; distiphallus narrow, of
uniform breadth to acute apex. Anterior margin of
fused surstyli deeply cleft medially to midlength of
structure; B-index about 1.0; C-index 0.5-0.8.
TYPE.?Holotype male, USNM 70530.
TYPE-LOCALITY.?Squaw Lake, Itasca State Park,
Clearwater County, Minnesota (VIII-14-1963, D.
L. Deonier).
PARATYPES.?43 (180*0", 25? $ ) from 4 locali-
ties:
Maine: Mount Desert Island (VIII-12-1935, W. Proc-
t e r ) ^ ? 9 (ANSP).
Minnesota: Basswood Lake, Lake County (VIII?16?
1950, R. Namba), 1$ (UMC); Squaw Lake, Itasca State
Park (VIII-14-1963, D. L. Deonier), 6$$, 29 9 (ISC
and USNM); Two Island Lake, Itasca State Park (VI-28-
1963, D. L. Deonier), 6$ <$, 16 9 9, (VIII-8-1963, D. L.
Deonier), 5$ $, 59 9 (USNM, ISC, and DLD).
REMARKS.?Hydrellia columbata, H. surata, and
H. prudens are sibling species and, as in several other
such cases in the genus, examination of the male
terminalia may be necessary for positive diagnosis.
Adult. Habitats recorded: leaves of Potamogeton
natans, P. gramineus, Nuphar advena, Sparganium
chlorocarpum, and Eleocharis palustris.
I collected many adults of H. columbata by the
lighted-receptacle method on Two Island Lake,
Itasca State Park, Minnesota. There, on a small
floating island, the adults covered most of nearly
every spike-rush stem examined. During the daytime
I saw adults nearly covering leaves of S. chlorocarpum
and P. natans at several localities in this vicinity.
Many of these adults stood with faces or antennae
touching. This seemed to be epigamic behavior.
Host plants. In my unsuccessful search for im-
matures of this species I examined Potamogeton
amplifolius, P. natans, P. gramineus, P. robbinsii,
Zannichellia palustris, Eleocharis palustris, Spar-
ganium chlorocarpum, and Glyceria sp.
Hydrellia crassipes Cresson
FIGURES 7, 33
Hydrellia crassipes Cresson, 1931, p. 107; 1944b, pp. 169,
174.?Wirth, 1965, p. 744 [catalog listing].
DIAGNOSIS.?Palpus moderate yellow; 7-10 aristal
rays; antenna dark brown; ocular index 8.0-10.0;
1 basal coxal; male hind femur grooved ventrally
and hind tibia flanged. Male length 2.13-2.30 mm;
female 2.30-2.55 mm. Male terminalia as in Fig-
ure 33.
HEAD.?Face yellowish gray or occasionally light
gray; 4-8 pfa; epistomal index 1.2-1.7; mesofacial
index 2.0-2.5; A-index 1.7-2.2; ocular index 8.0-
10.0. Palpus moderate yellow; 7-10 aristal rays;
antenna, fronto-orbital area, and frontal vitta dark
brown; most of parafrontale velvety black; 15-19
postoculars.
THORAX.?Ppn and mesonotum dark gray; 3-4
adc and 2-3 pdc; pleuron light gray; legs light-gray
pruinose except moderate orange-yellow tarsi; male
hind femur flanged posteroventrally and with two
anteroventral rows of close-set short setae; male mid
tibia flanged anteroventrally (Figure 7). Wing length
2.21-2.55 mm; veins usually dark brown; 5-9 setae
on basal end of costa; 8-9 dorsal and 10-13 anterior
interfractural costals; costal-section ratios: I I : I 2.2-
2.6; I I I : IV 2.8-3.2; V: IV 3.5-4.0; M, ? 2 index
1.2-1.8.
ABDOMEN.?Terga semiglossy dark brown medi-
ally, light gray laterally and ventrally. Male ter-
minalia : median third of posterior margin of sternum
5 deeply concave; anterolateral margin of sternum 5
rounded through angle of 100?; copulobus acut-
angular (about 20?) posteriorly, notched medially at
midlength, and irregularly setose. Postgonite bent
anterolaterad; postgonite uncus short, blunt, and
straight; distiphallus finely lamellate ventrally and
tapering to mucronate apex. Anterior margin of
fused surstyli broadly emarginate with about 8-9
setae on lateral corners; B-index about 2.5; C-index
1.8-2.0.
52 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TYPE.?Holotype male, in Ohio State University
collection (type no. not used).
TYPE-LOCALITY.?Sandusky, Cedar Point, Ohio
(VI11-4-1902, collector unknown).
SPECIMENS EXAMINED.?153 (66c? cf, 8 7 9 ? )
from 24 localities:
Illinois : Havanna (IX-19-1895, C. Hart), 19 .
Iowa: Goose Lake, Hamilton County (VII-2-1961, D. L.
Deonier), 7 $ 5 , 12$ 9 ; Little Wall Lake, Hamilton County
(IX-22-1962, D. L. Deonier), 1,$, 1$ ; Springbrook State
Park, Guthrie County (VI-23-1961, D. L. Deonier), 1 $ ;
Spring Lake, Greene County (IX-19-1961, D. L. Deonier),
115 5 , 7 9 $ .
Maine: Mount Desert Island (VII-12-1935, W. Procter,
1 $ .
Michigan: Hamburg, Livingston County (VI-8-1930, G.
Steyskal), 1 5 , (VIII-13-1933, G. Steyskal), 19 .
Minnesota: Basswood Lake, Lake County (VIII-16-1950,
R. Namba), 1 9 ; Chambers' Creek, Itasca State Park (VII-
18-1963, D. L. Deonier), 4 5 3 , 2 9 9 ; Douglas Lodge Bay,
Itasca State Park (VI-20-1963, D. L. Deonier), 1 9 ; Mis-
sissippi River near north entrance Itasca State Park (VII?
9-1963, D. L. Deonier), 1 5 ; Squaw Lake, Itasca State Park
(VIII-14-1963, D. L. Deonier), 29 9 ; Two Island Lake,
Itasca State Park (VI-28-1963, D. L. Deonier), 3 5 5 ,
79 9 ; 1.3 miles south of main entrance Itasca State Park
(VI-26-1963, D. L. Deonier), 2 5 5 , 69 9 ; 6.5 miles east
of Waubun (VIII-18-1963, D. L. Deonier), 1 5 , 19-
New Jersey: Manahawkin (VIII?1?1912, collector un-
known), 1 5 ?
North Carolina: Highlands Biological Station, Highlands,
Macon County (VI-15-VII-10-1968, D. L. Deonier),
105 5 , 69 9 ; Trussell's Pond, Horse Cove near Highlands,
Macon County (IX-2-1968, D. L. Deonier), 65 5 , 8 9 9 .
Ohio: Cedar Point, Sandusky River (VIII-4-1902, col-
lector unknown), 85 5 , 59 9 ; (VII-14-1903, collector
unknown), 2 5 5 , 2 9 9 .
Ontario: Bark Lake (VII-22-1939, D. Bullock), 1 9 ;
Marmora (VII-24 and VIII-7-1952, C. Boyle), 2 9 9 ,
(VIII-7-1952, J. McAlpine), 35 5 , 59 9 , (IX-9-1952,
J. McAlpine), 1 5 , 69 9 , (VIII-25 and IX-8-1952, E.
Smith), 39 9 .
Oregon: Oregon City (IX-no date and year, collector un-
known), 1 9 .
Pennsylvania: Pittsburgh (VIII-22-1931, H. Kahl), 19 .
Quebec: Perkins' Mills (VIII-25-1949, E. Shewell),
45 5,39 9.
REMARKS.?This species would be very distinct,
because of the male hind femoral modification, if
the cryptic sibling species H. saltator did not exist.
I did not see the holotype, but I did study several
paratypes of this species.
Adult. Habitats recorded: leaves of Potamogeton
nodosus, P. natans, P. epihydrus, Nuphar advena,
Nymphaea tuberosa, and N. odorata; sedge meadow,
riffle rocks, and limnic wrack.
I observed copulation many times in the field. In
one instance, at a pond near Itasca State Park,
Minnesota, the pair was surrounded by an aggrega-
tion of eight to ten specimens circling and following
one another in an agitated manner and showing
repeated mounting attempts. A large specimen of
Notiphila sp. kept walking through this aggregation
with its wings raised; this may have contributed to
the highly active, repetitious movement of the aggre-
gation. Observed under a microscope, a male in
copulation can be seen to have the posterolateral
edges of tergum 5 or 6 of the female firmly clamped
between his hind femora and tibiae with his hind
tarsi pushing against the female's sterna. I have
illustrated in Figure 7 the enlarged, grooved male
hind femur and flanged hind tibia. The short, stout,
seriated setae in the male hind femoral groove may
function for both sexes during copulation. In the
male, they could receive stimuli that aid in proper
mount positioning, and in the female the setae could
stimulate various copulatory movements of her
abdomen. I could detect no differences in the male
hind femoral apparatus between H. crassipes and
H. saltator.
This species seems to mate more readily in cap-
tivity than several others of the genus. If this is true
it offers opportunities of establishing a laboratory
colony for the purpose of studying the copulatory act
microscopically. Perhaps because the male tightly
clamps the female abdomen, quick freezing of
specimens while they are in copulation could be
accomplished.
Host plants. During the summer of 1968 I reared
three adults from Juncus debilis at Highlands
Biological Station, North Carolina. I found eggs
deposited on the tips of these rushes, which were
growing in very shallow water. Several white
Hydrellia larvae, determined as H. crassipes, were
found in these rush plants and in Scirpus smithii
plants, however most of these could not be reared to
the adult stage. My host-plant determinations for this
fly have not been confirmed by a plant taxonomist.
The adults spend most of their time feeding and
mating on floating vegetation, such as water lilies
and pond weeds, and they very seldom go to the
host rushes and sedges except to oviposit. This infor-
mation I gathered during about 20 hours of field ob-
NUMBER 6 8 53
servation. Various population studies have now been
started on this species, one of which hopefully will
provide data on the behavior of these flies at locali-
ties lacking floating vegetation.
Hydrellia cruralis Coquillett
FIGURES 63,96, 108, 128, 132
Hydrellia cruralis Coquillett, 1910a, p. 131.?Cresson, 1924,
p. 162; 1931, pp. 104, 106; 1934, p. 235; 1944b, pp. 169,
172.?Berg, 1950, pp. 375, 376, 377, 378-382, 383 (pi. 1,
figs. 5-8), 384, 385 (pi. 2, fig. 4) , 386, 387, 388, 389
(pi. 3, fig. 4) , 390, 396 [biology, morphology, and taxon-
omy of immatures].?Wirth, 1956, p. 16.?Judd, 1964,
p. 411 [ecology].?Deonicr, 1964, p. 116; 1965, pp. 500,
505 [ecology].?Wirth, 1965, p. 744 [catalog listing].
DIAGNOSIS.?Palpus moderate yellow; 6-9 aristal
rays; body length:wing length 1.0-1.2; antennal
segment 3 moderate yellow; thoracic pleuron light
gray; 4 mesokatepisternals; tibiae moderate yellow.
Male length 1.96-2.55 mm; female 2.21-3.23 mm.
Male terminalia as in Figure 63.
HEAD.?Face light yellowish brown or light gray;
4-7 pfa; no sfa; epistomal index 1.4-1.9; meso-
facial index 1.5-2.3; vertex index 3.3-4.7; A-index
1.6-1.8; ocular index 4.8-8.6. Palpus moderate
yellow; 6-9 aristal rays; antenna dark brown except
moderate-yellow segment 3; fronto-orbital area and
frontal vitta strong yellowish brown; most of para-
frontale velvety dark brown; 12-16 postoculars.
THORAX.?Ppn light gray; mesonotum strong yel-
lowish brown; 3-4 adc and 3-4 pdc; pleuron light
gray; 4 mesokatepisternals (1 macrochaetous) ; 2
basal coxals (1 macrochaetous); legs light-gray
pruinose except moderate-yellow tibiae. Wing length
1.94-2.64 mm; veins light yellowish brown; 8-11
setae on basal end of costa; 7-9 dorsal and 9-14
anterior interfractural costals; costal-section ratios:
I I : I 1.8-2.2; I I I : IV 2.8-3.2; V: IV 3.0-3.5: M, ?
 2
index 1.1-1.5.
ABDOMEN.?Terga sparsely strong yellowish-brown
pruinose over dark brown medially and light gray
laterally and ventrally. Male terminalia: median
third of posterior margin of sternum 5 nearly square-
notched and congruent with distiphallus; anterolat-
eral margin of sternum 5 roundly right-angled;
copulobus irregularly setose, acutangular posteriorly
(apex rounded). Postgonite bent anteriad and
postgonite uncus about 90? laterad; distiphallus very
short, tapering to acute apex. Anterior margin of
fused surstyli varying from acutangular (posterior
breadth of surstyli about 15.0 times anterior breadth)
to truncate; B-index about 2.5; C-index 0.8-1.2.
EGG.?Length 0.50-0.60 mm; maximum breadth
0.12-0.17 mm. Chorion (not illustrated) white or
light yellowish brown, corrugate, with about twice
as many ridges as in H. ischiaca (Figure 76); these
infrequently anastomosing and somewhat more un-
dulate than in H. ischiaca. Micropylar protuberance
infundibulate and visible in dorsal view. This de-
scription in part after Berg (1950).
FIRST-INSTAR LARVA.?Length 0.80-1.50 mm;
maximum breadth 0.15-0.22 mm. Frontoclypeal
length 0.15-0.19 mm. Creeping welts large and con-
spicuous. Newly eclosed larva translucent light
yellowish gray; late first-instar larva translucent,
light green. This description in part after Berg (1950).
SECOND-IN STAR LARVA.?Length 1.50-4.15 mm;
maximum breadth 0.25-0.60 mm. Frontoclypeal
length 0.25-0.40 mm. Creeping welts not so con-
spicuous. Body translucent, light green.
THIRD-INSTAR LARVA.?Length 3.00-6.50 mm;
maximum breadth 0.60-1.20 mm. Frontoclypeal
length 0.40-0.62 mm; dorsal phragmatal ramus
mostly hyaline (Figure 96). Ventral frontoclypeal
index 4.0-4.5; phragmatal index 1.3-1.4; bifurca-
tion index 2.4-2.7; clypeal-arch index 1.8-2.3.
Clypeal arch smoothly sloping at about 20? in rela-
tion to lower frontoclypeal margin. Mouth-hook beak
distinctly longer than base; maximum mouth-hook
base thickness about 1.5 times that of beak; mouth-
hook light spot large and ovoid. Prothorax and meso-
thorax only moderately spinulose; creeping welts of
8-13 transverse spinular rows; abdominal segment
8 with ventral, transverse row of 6 setulae and about
3?4 annuli of spinules; these spinules heavy and
bidentate or tridentate dorsally. Body translucent,
light green.
PUPARIUM.?Length 3.50-4.75 mm; maximum
breadth 1.00-1.60 mm; distinctly scalloped laterally
and subcylindrical (Figure 108). Puparial length:
minimum breadth 12.0-13.0; maximum breadth:
minimum breadth 3.7-4.2; anal-plate index 2.8-3.2.
Prothoracic end truncate or slightly convex in ven-
tral view (constricted posterolaterally) ; prothorax
only slightly rugulose ventrally; head-lobe scar
obovoid to subtriangular; anal plate subrectangular
to crescentric, with anterior margin slightly concave.
54 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Empty puparium usually dark brown. Early pupa
light green.
TYPE.?Holotype female, USNM 13102.
TYPE-LOCALITY.?Riverton, New Jersey (IX-1909,
H. S. Harbeck).
ADULT SPECIMENS EXAMINED.?560 (2O7cfcf
353? ?. ) from 48 localities:
Alabama: Mobile (VI-14-1953, M. Wheeler), 1?.
Alaska: Valdez (VIII-15-1948, E. Marks), 2 9 9-
Connecticut: Candlewood Lake (VIII-30-1941, A.
Melander), 1$.
Florida: Fruit-fly survey (no dates?1951, collector un-
known), 59 9 .
Idaho: Priest Lake, Tule Island (VIII-19-no year, A.
Melander), 1$.
Illinois: Pistakee Bay (VII-7-1933, A. Melander), 2 9 9-
Iowa: Spring Lake, Greene County (IX-19-1961, D. L.
Deonier), 4 3 3 , 2 9 9 .
Kansas: Leavenworth County Lake (VIII-27-1962, D. L.
Deonier), 7 3 3 , 8 9 9 .
Maryland: Chesapeake Beach (IX-8-1920, J. Aldrich),
23 3 , 39 9 , (IX-18-1922, J. Malloch), 1 9 ; Plummer's
Island, Potomac River (IX-1-1962, K. Krombein), 10 3 3 ,
7 9 9 .
Massachusetts: Fall River (VII-3-1930, A. Melander),
19 .
Michigan: Black River, Cheboygan County (VI-20-
1941, C. Berg), 8 3 3 , 8 9 9 ; Carp River, Cheboygan
County (VII-1-1941, C. Berg), 59 9 ; Cheboygan Pool,
Cheboygan County (VIII-6-1941, C. Berg), 1 3 , 1 9 ;
Cheboygan County (VI-12-1940, C. Berg), 1 3 , (VIII-
14-1940, C. Berg), 1 3 , (VIII-6-1941, C. Berg), 1 3 , (VI-
26-1946, C. Berg), 3 9 9 , (VII-5-1946, C. Berg), 2 3 3 ,
1 9 , (VII-16-1946, C. Berg), 1 3 , (VIII-15-1946, C.
Berg), 19 , (VII-21-1947, C. Berg), 23 3 ; Chippewa
County (VIII-26-1941, R. Dreisbach), 19 ; Douglas Lake,
Cheboygan County (VI-25-1940, C. Berg), 1 3 , 19 , (VII-
3-17-1941, C. Berg), 39 9 , (VI-26-1946, C. Berg), 1 9 ,
Grosse Isle, Wayne County (IX-12-1948, G. Steyskal),
1 9 ; Hamburg, Livingston County (VIII-13-1933, G. Stey-
skal), 1 9 ; Huron River, Washtenaw County (all reared by
C. Berg: 111-26 and V-7-1941, 111-24 and IV-15-1942),
7 3 3 , 69 9 ; Indian River, Cheboygan County (VI-29-
1940, C. Berg), 1 9 , (VI-6-1941, C. Berg), 1 3 , (VI-29-
1941, C. Berg), 33 3 , 109 9 ; Midland County ( IX-5-
1937, G. Steyskal), 1 9 ; Monroe (VII-2-1939, G. Stey-
skal), 1 9 ; Ocqueoc Lake, Presque Isle County (VII-13-
1941, C. Berg), 2 3 3 , 2 9 9 , (VII-15-1941, C. Berg), 1 3 ;
Third Sister Lake, Washtenaw County (all reared by C.
Berg: 111-15-30-1940, V- l 1-20-1940, VI-29-1940, VII -
2-1940, 1-3-1941, II-1-3-1941, VI-20-1941, H-2-1942,
IV-3-1942, and V-3-1942), 213 3 , 239 9 ; Washtenaw
County (all reared by C. Berg: VI-24-25-1940, 1-31-1941,
II?5?11-1941 and III-7-21-1941), 63 3 , 10 9 9 ; Whit-
more Lake, Washtenaw County (all reared by C. Berg:
II-1-1941, III-3-18-1941, and 1-23-1942), 5 3 3 , 89 9.
Minnesota: Biological Station, Itasca State Park (VI-26-
1963, D. L. Deonier), 4 9 9 ; Detroit Lakes (VIII-9-1935,
A. Melander), 29 9 ; Eaglenest (IX-11-1958, W. Balduf),
1 9 ; Headwaters of LaSalle Creek, Itasca State Park (VIII-
6-1963, D. L. Deonier), 1 9 ; Long Lake, Clearwater County
(VII-15-1963, W. Schmid), 29 9 , (VIII-8-1963, W.
Schmid), 23 3 ; Rapid River Logging Camp, Hubbard
County (VII-2-1963, D. L. Deonier), 1 9 ; Squaw Lake,
Itasca State Park, (VIII-14-1963, D. L. Deonier), 4 3 3 ,
49 9.
New Jersey: Trenton (VII-21-1909, collector unknown),
2 9 9 .
Ohio: Cedar Point, Sandusky River (VIII-5-1902, col-
lector unknown), 1 3 ?
Ontario: Arnprior (VIII-19-22-1930, C. Curran), 1 3 ,
29 9 ; Black Rapids, Ottawa (VIII-6-1959, J. Vockeroth),
39 9 ; London (all trapped by W. Judd: VIII-27-1956,
V-29-1958, VI-2-1958, VI-5-1958, VI-15-1958, VI I -
8-30-1958, VIII-14-31-1958, IX-2-27-1958, and X-9-
1958), 1013 3 , 1869 9 ; Marmora (VII-21-1952, J. Mc-
Alpine), 23 3 , 19 , (VIII-4-1952, J. McAlpine), 19 ,
(VIII-7-1952, J. McAlpine), 1 9 , (VIII-11-1952, J. Mc-
Alpine), 19 , (IX-9-1952, J. McAlpine), 19 .
Pennsylvania: West Fairview (Vll-no date-1960, M.
Wheeler), 13-
Quebec: Lac Bernard (VIII-7-1938, G. Shewell), 2 9 9-
Texas: Comal River (111-24-1942, A. Melander), 2 3 3 ,
89 9 ; Devil's River (VIII-29-1940, A. Melander), 1 3 ,
(IV-27-1942, A. Melander), 1 9 ; Garner State Park,
Uvalde County (IV-3-1955, W. Wirth), 1 3 ; Pedernales
River, Gillespies County (111-19-1955, W. Wirth), 49 9 .
Virginia: Alexandria (VIII-5-1922, J. Aldrich), 1 3 .
West Virginia: Marlington (VIII-15-1930, G. Auxier),
1 3 , 49 9 .
IMMATURE SPECIMENS EXAMINED: 60 from 6 local-
ities :
Michigan: Cheboygan County (collected and reared by
C. Berg); Third Sister Lake, Washtenaw County (collected
and reared by C. Berg).
Minnesota: Biological Station, Itasca State Park (col-
lected and reared by D. L. Deonier) ; Long Lake, Clear-
water County (collected by W. Schmid) ; Rapid River
Logging Camp, Hubbard County (collected and reared by
D. L. Deonier); Squaw Lake, Itasca State Park (collected
and reared by D. L. Deonier).
REMARKS.?Adult. Habitats recorded: leaves of
Potamogeton amplifolius, P. natans, P. richardsonii,
P. nodosus, Polygonum scabum, Nuphar advena, and
fern.
According to Berg (1950), adult H. cruralis feed
on Potamogeton leaves by chewing small holes in
them. However, they are polyphagous, for they attack
insects, especially congeners, trapped in the surface
film. From these and from dead and decaying
arthropods, they suck tissue fluids and possibly fungal
fragments.
NUMBER 6 8 55
Berg surmised that the unperfected motor co-
ordination observed in newly emerged adults of
this species indicated a high risk of drowning during
emergence in rough water.
I observed what may have been epigamic behavior
at Squaw Lake, Itasca State Park, when two speci-
mens approached each other while holding their
wings about 45? from their abdomina. They sud-
denly flew away before I could capture them. At this
same locality, I briefly observed a pair, in copulation,
skim rapidly over the water surface for about 2
meters between two floating leaves. Berg reared flies
which mated within 24 hours after emergence and
oviposited within 24 hours after mating. He found
that females deposited eggs parallel in a single-
layered mass in partially concealed sites, especially
in broken midribs of leaves and in old, open leaf
mines. Oviposition observed was always near the
water surface. In natural microhabitats, the females
oviposited in folded leaves and stipules, and on stems
near the water surface.
Egg and larva. In Berg's work, the incubation
period ranged from 2 to 7 days. The newly hatched
larva immediately tried to excavate a mine in any
nearby leaf. This leaf could be floating or submerged.
The first larval stadium ranged from 2 to 3 days, the
second from 5 to 8 days, and the third from 10 to 18
days. I found the third larval stadium ranged from
5 to 7 days for eight larvae. Berg found that the
larvae migrated from dying or skeletonized leaves to
fresh ones. In several observations on infestations of
Potamogeton richardsonii, he found a lower leaf
with a mine containing first-instar exuviae, the next
higher leaf with second-instar exuviae in a mine,
and two successively higher leaves containing, re-
spectively, an empty mine and a mature third-instar
larva or a puparium.
Dr. W. Schmid collected a second-instar larva and
several puparia in Potamogeton amplifolius growing
at a depth of 5.8 meters in Long Lake, Clearwater
County, Minnesota. Occurrence of larvae at this
depth indicates that larvae are sometimes affected
by water current and wind either directly during
their random migrations between leaves or indirectly
by dislocation of host plants. Judd (1964) caught
297 adults in floating emergence traps over sites with
depths ranging from 0.6 to 9.0 meters. Potamogeton
amplifolius grew at all of these sites in Saunders
Pond, near London, Ontario.
Berg found larvae of the three instars overwinter-
ing in a quiescent state in the winter buds of several
species of Potamogeton under the ice.
Puparium. The puparial phase ranged from 8 to
14 days in Berg's laboratory. For six puparia, I
found the phase range to be 7-9 days. The third-
instar larva usually pupariated with its spiracular
peritremes inserted in a leaf midrib. The life cycle
from egg to egg completed several times in Berg's
laboratory ranged from 32 to 53 days, with a mean
of 41 days.
Host plants. Berg collected larvae and puparia
from Potamogeton alpinus, P. amplifolius, P.
epihydrus, P. foliosus, P. gramineus, P. illinoensis,
P. natans, P. nodosus, P. praelongus, P. richardsonii,
and P. zosteriformis. Potamogeton amplifolius, P.
richardsonii, and P. praelongus had the highest
percentages of infestation.
Parasites. Berg reared the braconids Ademon niger,
Chorebidea sp., and Chorebidella bergi and the
diapriid Trichopria columbiana from puparia of H.
cruralis. I have reared Ademon niger from specimens
collected in Virginia.
Hydrellia deceits Cresson
Hydrellia deceits Cresson, 1931, p. 107; 1944b, pp. 170,
171.?Wirth, 1965, p. 744 [catalog listing].
DIAGNOSIS.?Palpus dark brown; 9-11 aristal
rays; all fronto-orbitals anteriorly inclined; 6-8 dor-
sal and 8-9 anterior interfractural costals; face light
yellowish brown or yellowish gray medially, with
light gray vitta along secondary facial row; thoracic
pleuron with one continuous light-gray area. Female
length 2.04-2.21 mm.
HEAD.?Face light yellowish brown or yellowish
gray medially with lunule and vitta along secondary
facial row light gray; 4-6 medially inclined pfa;
5-8 medially inclined sfa; epistomal index 1.0-1.2;
mesofacial index 2.0-2.2; vertex index 5.0-7.0;
A-index 1.8-2.2; ocular index 5.5-8.0. Palpus dark
brown; 9-11 aristal rays; antenna dark brown;
frontal vitta semiglossy dark brown; parafrontale
(except strong yellowish-brown fronto-orbital area)
opaque very dark brown; fronto-orbitals anteriorly
inclined: 13-16 postoculars.
THORAX.?Ppn semiglossy dark brown and meso-
notum glossy dark grayish green; 3-4 adc and 2 pdc;
pleuron light gray except semiglossy light brown on
56 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
upper fifth of mesanepisternum, around wing base,
and on laterotergite; legs (except light-gray coxae
and dark-brown tarsi) dark brown with sparse light-
gray pruinosity. Wing length 2.30-2.48 mm; veins
dark brown; 6 setae on basal end of costa; 6-8
dorsal and 8-9 anterior interfractural costals; costal-
section ratios: I I : I 2.2-2.6; I I I : IV 2.8; V:IV 3.2-
3.6; M i + 2 index 1.3-1.5.
ABDOMEN.?Most of terga glossy dark grayish
green.
TYPE.?Holotype female, USNM 21842.
TYPE-LOCALITY.?Near Plummer's Island, Potomac
River, Maryland (VIII-12-1914, R. C. Shannon).
SPECIMENS EXAMINED.?2? $ from 2 localities:
Maryland: Plummer's Island, Potomac River (VIII-13-
1913, W. McAtee), 1 9 ; near Plummer's Island, Potomac
River (VII-14-1914, R. Shannon), 19.
REMARKS.?Although the male is still unknown,
I consider this a distinct species on the basis of
geographic distribution and the speciation trend in
the H. proclinata group. At the present stage, one
must rely on the geographic distribution and the
small color differences to distinguish H. decens from
H. proclinata and H. melanderi.
Hydrellia deceptor, new species
FIGURES 42, 94
DIAGNOSIS.?Palpus moderate yellow; prementum
light gray; 7-9 aristal rays; vertex index 6.5-7.5;
ocular index 7.5-9.5; lower half of postocular area
light gray, upper half light brown; thoracic pleuron
light gray. Male length 1.79 mm; female 2.04-2.13
mm. Male terminalia as in Figure 42.
HEAD.?Face light gray or yellowish gray; 4-6
pfa; epistomal index 1.1-1.4; mesofacial index 2.0-
2.4; vertex index 6.5-7.5; A-index 2.0-2.3; ocular
index 7.5-9.5. Palpus medium yellow; 7-9 aristal
rays; prementum light gray; antenna velvety dark
brown; frons semiglossy dark brown except moderate
olive-brown fronto-orbital area; 13-17 postoculars.
THORAX.?Ppn and mesonotum moderate brown;
3?4 adc and 2 pdc; pleuron light gray; legs densely
light-gray pruinose except dark-brown tarsi. Wing
length 1.99-2.55 mm; veins dark brown; 6-8 setae
on basal end of costa; 6-9 dorsal and 8-11 anterior
interfractural costals; costal-section ratios: I I : I 2.2-
2.5; I I I : IV 2.5-3.2; V:IV 3.0-4.0; Mi ?
 2 index
1.3-1.6.
ABDOMEN.?Terga moderate brown except light
gray laterally and ventrally. Male terminalia: median
third of posterior margin of sternum 5 deeply con-
cave; anterolateral margin of sternum 5 acutangular
in form of blunt hook; copulobus acuminate pos-
teriorly and irregularly setose. Postgonite and post-
gonite uncus bent slightly laterad; distiphallus ven-
trally lamellate and tapering to acutangular, retuse
apex. Anterior margin of fused surstyli narrowly
indented medially from broad convexity; B-index
about 2.2; C-index 0.6-1.0.
TYPE.?Holotype male, USNM 70531.
TYPE-LOCALITY.?Sacramento, California ( IX-
20-1957, A. A. Grigarick).
PARATYPES.?3 $ 9 from Sacramento, California
(IX-21-1957, A. Grigarick), in UCD and ISC.
REMARKS.?This species is so similar to H.
notiphiloides that one should study the male ter-
minalia to confirm determinations.
Host plants. Grigarick reared a few specimens
from Sagittaria sp. collected near Sacramento, Cali-
fornia, 20 and 21 September, 1957 during his re-
search on H. griseola. I found no indication as to
whether this Sagittaria species was an incidental
puparial host or a more constant host for larvae and
puparia. Grigarick very probably collected the host
plants in or near a rice field.
Hydrellia definita Cresson
FIGURE 60
Hydrellia definita Cresson, 1944b, pp. 165-166.?Deonier,
1964, p. 116; 1965, pp. 500, 505 [ecology].?Wirth, 1965,
p. 744 [catalog listing].
DIAGNOSIS.?Palpus mostly moderate yellow, but
partly dark brown; 6-8 aristal rays; mesofacial index
2.0-2.3; body length:wing length 0.9-1.0; thoracic
pleuron yellowish gray except light brown on lower
half of propleuron, anterior part of mesokatepister-
num, and laterotergite; costal section V: IV 3.6-4.2.
Male length 1.73-2.13 mm; female 1.87-2.55 mm.
Male terminalia as in Figure 60.
HEAD.?Face yellowish gray medially, light gray
laterally; 4-6 pfa; 1-4 sfa; epistomal index 1.1-1.7;
mesofacial index 2.0-2.3; vertex index 4.8-5.6;
A-index 1.5-2.7; ocular index 8.0-13.0. Palpus
mostly moderate yellow but partly, or infrequently
NUMBER 6 8 57
wholly, dark brown; 6-8 aristal rays; antenna,
fronto-orbital area, and frontal vitta dark brown;
most of parafrontale velvety dark brown; 12-16
postoculars.
THORAX.?Ppn and mesonotum moderate brown;
4-6 adc and 2-3 pdc; pleuron yellowish gray except
light brown on lower half of propleuron, anterior
part of mesokatepisternum, and laterotergite; legs
light-gray pruinose except dark brown tarsi. Wing
length 1.96-2.72 mm; veins light yellowish brown;
6-10 setae on basal end of costa; 7-10 dorsal and
8-12 anterior interfractural costals; costal-section
ratios: 11:1 2.5-3.0; I I I : IV 2.8-3.5; V:IV 3.6-
4.2; Mi ?
 2 index 1.3-1.8.
ABDOMEN.?Terga semiglossy dark gray. Male
terminalia: median third of posterior margin of
sternum 5 concave; anterolateral margin of sternum
5 roundly obtusangular; copulobus somewhat sinuate
posteriorly and irregularly setose. Postgonite bent
anteriad and postgonite uncus straight; distiphallus
of uniform breadth to mucronate and ventrally
lamellate apex. Anterior margin of fused surstyli
broadly emarginate; B-index about 2.8; C-index
1.8-2.0.
TYPE.?Holotype male, ANSP 890.
TYPE-LOCALITY.?Spanish Fork, Utah (no dates,
D. E. Hardy).
SPECIMENS EXAMINED?64 (19c? d, 45 $ $ ) from
24 localities:
Alaska: King Salmon, Naknek River (VII-31-1952, W.
Mason), 19.
California: Jenks Lake (VII-7-1950, A. Melander), IS.
Illinois: Centerville, Sangamon River (VIII-16-1914,
collector unknown), 19.
Iowa: Ames (VIII-4-1960, D. L. Deonier), 19 ; Banner
Mine Area, Warren County (VIII-7-1960, D. L. Deonier),
1<J, 19; Lake Odessa, Louisa County (VIII-9-1960, D.
L. Deonier, 1 $, 7 9 9 ; Little Wall Lake, Hamilton County
(IX-22-1962, D. L. Deonier), 19; Springbrook State
Park, west /2 sec. 33, T. 8 north, R. 31 west, Guthrie
County (VII-19-1960, D. L. Deonier), 1$.
Kansas: Kansas University Natural History Reservation
near Lawrence, Douglas County (VI-7-1963, D. L.
Deonier), 19; Leavenworth County Lake (VIII-27-1962,
D. L. Deonier), 1 9 ; Sappa Lake, Decatur County (VIII-
31-1961, D. L. Deonier), 19.
Michigan: Emmett County (VII-14-1954, C. Berg),
2$ $, 39 9 ; Vineyard Lake (VIII-1-1954, G. Steyskal),
19.
Minnesota: Basewood Lake, sec. 9 T. 64 north, R. 10
west, Lake County (VIII-16-1950, R. Namba), 1 9 ; Itasca
State Park (VI-20-1938, H. Milliron), 19; west side
across from Biological Station, Lake Itasca (VII-28-1963,
D. L. Deonier), 39 9; 2.5 miles west of Waubun (VIII-
18-1963, D. L. Deonier), 43 5 ,49 9.
Nebraska: Lincoln, Lancaster County (VIII-16-1960,
W. Rapp), 19.
New York: Dryden Lake, Tompkins County (VIII-8-
1961, D. L. Deonier), 7$$, 139 9.
Ontario: Pembroke (VII-3-1938, A. Melander), 19.
Saskatchewan: Rock Glen (VIII-2-1955, C. Miller),
It.
South Dakota: Hot Springs (VII-13-1924, collector
unknown), 19-
Wyoming: Slide Lake (VIII-14-1951, collector un-
known), 1 9 ; 12 miles northwest of Lusk (VH-no date-
1895, collector unknown), 1 $ .
REMARKS.?Hydrellia definita, H. manitobae, H.
suspecta, H. subnitens, and H. amnicola seem to
constitute a distinct species group.
Adult. Habitats recorded: leaves of Potamogeton
nodosus and Polygonum scabrum; tundra, limnic
wrack, sedge meadow, and at light.
I observed males pushing their head and thorax
up and down before females at Dryden Lake, Tomp-
kins County, New York.
Hydrellia discursa, new species
FIGURES 30, 70, 77, 80, 97, 113, 125, 127
Hydrellia ascita; Cresson, 1942 [in part], p. 78.
DIAGNOSIS.?Palpus moderate yellow; 5-8 aristal
rays; face planate; vertex index 4.0-5.4; ocular index
4.4-5.6; male abdomen slightly convex posteriorly;
female cercus ovoid apically and truncate basally;
antennal segment 3 at least partly moderate yellow;
apicodorsal antennal prominent. Male length 1.79-
1.96 mm; female 1.70-2.30 mm. Male terminalia as
in Figure 30; female as in Figure 70.
HEAD.?Differing mainly from H. bilobifera in the
following: mesofacial index 1.8-2.4; vertex index
4.0-5.4; ocular index 4.4-5.6.
THORAX.?Mainly as in H. bilobifera.
ABDOMEN.?Terga semiglossy dark gray medially,
light gray laterally and ventrally. Male terminalia:
median third of posterior margin of sternum 5 deeply
recessed and nearly congruent with distiphallus
apex; anterolateral margin of sternum 5 with dorsal
process projecting anterolaterad and covered by
sternum 4; posterolateral angle of copulobus about
75? from lateral corner (partly or wholly obscured
by loose fascicula of 4-5 strong macrochaetae) ;
posteromedial margin of copulobus with notch
58 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
paralleled and mostly obscured by 7-10 seriated
stout setae (almost a pecten); copulobus with dis-
tinct bare space posteriad from indistinct central
verruca. Postgonite bent laterad then mediad; pos-
gonite uncus spicate, directed anteromediad; dis-
tiphallus slightly constricted in middle third and
indistinctly carinate and lamellate ventrally to
mucronate apex; most of basiphallus exposed ven-
trally. Anterior margin of fused surstyli deeply and
broadly concave; B-index about 2.0; C-index 0.5-
0.8. Syntergum 9-f-10 slightly convex posteriorly.
Female terminalia: tergum 8 slightly shorter than
sternum 8; cercus irregularly setose laterally, ovoid
apically, truncate basally and less than 1.5 times as
long as wide (lateral view). Segment 7 with setae all
along posterolateral margin; median spermatheca
cupuliform and usually not as long as cercus.
EGG.?Length 0.47-0.56 mm; maximum breadth
0.14-0.16 mm. Chorion (Figure 77) white (initially)
with inconspicuous, occasionally anastomosing longi-
tudinal ridges; wide spaces between ridges smooth
as in those of H. bilobifera. Micropylar protuberance
infundibulate, visible in dorsal view; end opposite
micropylar protuberance with indistinctly lacunose
papilla.
FIRST-INSTAR LARVA.?Length 0.35-0.75 mm;
maximum breadth 0.10-0.15 mm. Frontoclypeal
length not measured; clypeal arch distinctly more
angular at level of cheliform spot in lateral view
than in third-instar larva; supraspiracular protuber-
ance present with central spinous seta; terminal
peritremal spine present; verriculose subspiracular
protuberance present; dorsal setae and spinules
apparently absent except on abdominal segment 8.
Newly eclosed larva translucent, light yellowish gray;
late first-instar larva same but with greenish-yellow
tinge.
SECOND-INSTAR LARVA.?Length 1.00-3.50 mm;
maximum breadth 0.20-0.50 mm. Frontoclypeal
length not measured. Dorsal setae absent; dorsal
spinules restricted to thorax and abdominal segment
8; abdominal segment 8 with 3-4 annuli of spinules.
Body translucent, light green.
THIRD-INSTAR LARVA.?Length 3.50-5.50 mm;
maximum breadth 0.30-0.60 mm. Frontoclypeal
length 0.42-0.50 mm (Figure 97). Ventral fronto-
clypeal index 2.5-2.8; phragmatal index 1.0-1.2;
bifurcation index 3.6-3.8; clypeal-arch index 1.7-
2.0. Clypeal arch sloping gradually at 10?-15? and
slightly convex at level of cheliform spot. Mouth-
hook beak and base lengths subequal; maximum
mouth-hook base thickness about 1.8 times that of
beak; mouth-hook light spot small, discal. Prothorax
and metathorax moderately spinulose; dorsal setae
and spinules restricted to thorax and abdominal
segment 8; abdominal segment 8 without ventral,
transverse row of setulae, but with 3-4 annuli of
spinules. Body translucent, light green.
PUPARIUM.?Length 2.75-4.00 mm; maximum
breadth 0.65-1.00 mm; fusiform (Figure 113).
Puparial length:minimum breadth 15.0-23.0; maxi-
mum breadth: minimum breadth 4.2-5.0; anal-plate
index 3.5-4.0. Prothoracic end semicircular or sub-
triangular in ventral view; head-lobe scar circular to
obovoid; maximum puparial breadth in metathorax;
prothorax distinctly rugulose ventrally and laterally;
anal plate subrectangular with anterior margin
straight to slightly convex. Empty puparium trans-
lucent, light yellowish brown. Pupal color as in H.
bilobifera.
TYPE.?Holotype male, USNM 70532.
TYPE-LOCALITY.?Samburg, Reelfoot Lake, Ten-
nessee, 36?22.9' N, 89?21.3' W (VIII-11-1962, D.
L. Deonier).
PARATYPES.?112 (48c" d1, 64? ? ) from 24 local-
ities :
California: Davis (X-21-1954, A. Grigarick), 13
(UCD); Maxwell, Colusa County (VIII-12-1954, A.
Grigarick), IS (UCD); Vidal, San Bernadino (IV-no
date-1948, R. Coleman), 19 (USNM).
District of Columbia: Washington (VIII-13-1923, W.
McAtee), 1$ (USNM).
Florida: Cape Sable (XII-18-1949, C. Sabrosky), 1$
(USNM).
Iowa: Goose Lake, Hamilton County (VII-2-1961, D.
L. Deonier), 4<J $ (USNM and ISC); Siewers Springs
State Park, Decorah (IX-9-1961, D. L. Deonier), 29 9
(USNM); Spring Lake, Greene County (IX-9-1961, D. L.
Deonier), 3 <$ $, 49 9 (USNM and ISC).
Kansas: Kansas University Natural History Reservation
near Lawrence, Douglas County (VI-7-1963, D. L.
Deonier), 1 3 , 29 9 (USNM and ISC); Leavenworth
County Lake (VIII-27-1962, D. L. Deonier), 3$ <J, 29 9
(USNM and ISC); Marais des Cygnes Wildlife Refuge,
Linn County (IX-5-1961, D. L. Deonier), 5$S, 9 9 9
(USNM and ISC).
Michigan: Cheboygan County (VII-26-1946, C. Berg),
1$ (USNM).
Minnesota: Biological Station, Itasca State Park (VI-
26-1963, D. L. Deonier), 1$ (USNM); Bohall Trail Bog,
Itasca State Park (VI-25-1963, D. L. Deonier), 19
(USNM); Headwaters of LaSalle Creek, Itasca State Park
NUMBER 6 8 59
(VIII-6-1963, D. L. Deonier), 29 9 (USNM); Mississippi
River near north entrance Itasca State Park (VII-9-1963,
D. L. Deonier), 19 (USNM).
Mississippi: Dickinson's Pond, Lamar County (VII-24-
1962, D. L. Deonier), 1?, 19 (DLD); Lake Shelby State
Park, Forrest County (VII-24-1962, D. L. Deonier), \$,
29 9 (USNM and ISC).
Ontario: Marmora (VIII-7-1952, J. McAlpine), 2$ $,
19 (CNC), (IX-8-1952, E. Smith), 3 <$ 3 (CNC and
USNM), (IX-9-1952, J. McAlpine), 4$ $, 1 9 ( CNC and
USNM) ; Ottawa (VII-2-1947, G. Shewell), 4 3 $ ( CNC).
Tennessee: Brewer's Bar Ditch, Reelfoot Lake, 36?27.1'
N, 89?21.2' W (VIII-10-1962, D. L. Deonier), 89 9
(USNM and ISC), (VIII-14-17-1962, D. L. Deonier), 1 9
(DLD); Miller's Camp, Reelfoot Lake, 36?24.3' N, 89?20'
W (VIII-12-1962, D. L. Deonier), 5$ $, 79 9 (USNM
and ISC); Samburg, Reelfoot Lake, 36?22.9' N, 89?21.3'
W (VIII-11-16-1962, D. L. Deonier), 9$ $, 169 9
(USNM and ISC).
Texas: Galveston (IX-13-1953, M. Wheeler), 1,5, 19
(USNM).
IMMATURE SPECIMENS EXAMINED.?45 from 3
localities:
Tennessee: Samburg, Miller's Camp, and Brewer's Bar
Ditch, Reelfoot Lake (collected and reared by D. L.
Deonier).
REMARKS.?According to Berg (1950, p. 391):
"Cresson designated only the flies reared from P.
alpinus {P. tenuifolius) collected at Nigger Creek,
Cheboygan County, in his type series, and he identi-
fied the others from other Potamogeton species as a
variety of H. ascita." Since I discovered several speci-
mens of H. discursa among material determined by
Cresson as H. ascita, Cresson may have suspected the
existence of a cryptic species when he designated
some as a variety of H. ascita.
Adult. Habitats recorded: leaves of Potamogeton
gramineus, P. nodosus, P. epihydrus, Nuphar advena,
Nymphaea tuberosa, Nelumbo lutea, Zizania aqua-
tica, and Oryza sativa.
At Dickinson's Pond, Lamar County, Mississippi,
the male of a pair observed in copulation held an
attitude almost perpendicular to the long axis of the
female's abdomen.
Egg and larva. The incubation period ranged from
3 to 5 days for 26 eggs. This is approximate, for I
collected the egg masses a short time after they were
deposited. Examination immediately after collection
revealed the eggs to be in an early embryonic stage.
I collected the egg masses on floating leaves of
Potamogeton gramineus at Samburg, Reelfoot Lake,
Tennessee. I collected several egg masses on stems
and leaves of Nelumbo lutea and Alisma sp. in the
same locality. I did not determine the species, but
circumstances indicated they were H. discursa. Three
larvae passed the first stadium in three days. Since
I collected several adults in the vicinity of water
primrose, Jussiaea repens, I supplied leaves of this
plant to first-instar larvae of H. discursa. They did
not mine in them. I did not measure the second
larval stadium, and I obtained only an approximate
measurement of 6 days for the third stadium of one
larva, since I only fixed the time of molting within
two days.
Puparium. The puparial phase ranged from 7 to
10 days for four specimens. The larvae pupariated
in the leaf axil, on the abaxial side of floating leaves,
and on the adaxial side of submerged leaves.
Host plants. I found larvae and puparia only in
Potamogeton gramineus and P. nodosus. In addition,
I examined Alisma sp., Heteranthera dubia, Sagit-
taria australis, Jussiaea repens, and Zizaniopsis
miliacea for larvae and puparia of this species.
Parasites. I reared one braconid of undetermined
species from a puparium and observed an undeter-
mined parasitic larva within a late pupa.
Hydrellia flavicoxalis Cresson
FIGURE 62
Hydrellia flavicoxalis Cresson, 1944b, p. 167.?Wirth, 1965,
p. 744 [catalog listing].
DIAGNOSIS.?Palpus moderate yellow; 7-8 aristal
rays; vertex index 7.0-10.0; body length:wing length
0.9; male tergum 5 broadly rounded posteriorly;
4?5 mesokatepisternals; 2 basal coxals; thoracic
pleuron light gray. Male length 2.38 mm; female
2.64 mm. Male terminalia as in Figure 62.
HEAD.?Face yellowish gray or light yellowish
brown; 5-6 pfa; epistomal index 1.4-1.8; mesofacial
index 2.0-2.5; vertex index 7.0-10.0; A-index 1.7-
1.9; ocular index 5.0-6.0. Palpus moderate yellow;
7-8 aristal rays; antenna and most of parafrontale
dark brown; frontal vitta and fronto-orbital area
moderate olive-brown; 14-16 postoculars.
THORAX.?Ppn usually yellowish gray; mesonotum
moderate brown; 4 adc and 4 pdc; pleuron light
gray except upper posterior third of mesanepistemum
moderate olive-brown; 4-5 mesokatepisternals (1
macrochaetous) ; 2 basal coxals (1 macrochaetous);
60 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
legs light-gray pruinose except dark brown tarsi.
Wing length 2.60-2.81 mm; veins light yellowish
brown; 6-8 setae on basal end of costa; 9-10 dorsal
and 11-14 anterior interfractural costals; costal-
section ratios: I I : I 2.5-3.2; I I I : IV 2.6-3.0; V:IV
3.5-3.9; M 1 + 2 index 1.3-1.5.
ABDOMEN.?Terga 1-3 or 2-A moderate brown
medially, light gray laterally and ventrally. Male
terminalia: median third of posterior margin of
sternum 5 straight; anterolateral margin of sternum
5 prominent and roundly right-angled; copulobus
slightly angular posteriorly (about 100? from lateral
corner), irregularly notched and irregularly setose.
Postgonite bent anterolaterad and postgonite uncus
straight; distal two-thirds of distiphallus uniform in
breadth. Anterior margin of fused surstyli narrowly
and deeply notched medially to about midlength of
structure, and prominently lobate laterally; B-index
about 2.0; C-index 6.0-8.0.
TYPE.?Holotype male, ANSP 6666.
TYPE-LOCALITY.?Walden, Jackson County, Colo-
rado (VII-27-1938, M. T. James).
SPECIMENS EXAMINED.?1 9 with same collection
data as for holotype; these two are the only known
specimens of this species. Walden, Colorado, is in
North Park, a high basin isolated by a ring of moun-
tain ranges having several peaks from 11,000 to
12,000 feet high. Thus, this species may very well
have a very small distribution.
REMARKS.?This is a member of the H. griseola
species group as indicated by the male terminalia
and the habitus.
Hydrellia floridana, new species
FIGURE 31
DIAGNOSIS.?Palpus dark brown; 6-7 aristal rays;
ocular index 9.0 or more; wing length 1.32-1.53
mm; face light gray or yellowish gray; parafrontale
mostly velvety dark brown; lower third of postocular
region light gray, upper two-thirds moderate olive-
brown; male mid tibia expanded. Male length 1.19-
1.39 mm; female 1.36-1.60 mm. Male terminalia as
in Figure 31.
HEAD.-?Face light gray or yellowish gray; 4 pfa;
epistomal index 1.2-1.4; mesofacial index 1.8-2.1;
vertex index 4.5-5.5; A-index 2.0-2.3; ocular index
9.0-15.0. Palpus dark brown; 6-7 aristal rays;
antenna and frontal vitta dark brown; parafrontale
(except moderate olive-brown fronto-orbital area)
velvety dark brown; 13-15 postoculars.
THORAX.?Ppn and scutellum bronzed moderate
olive-brown; remainder of mesonotum dark gray;
3-4 adc and 2 pdc; pleuron mostly moderate olive-
brown; coxae dark gray and tarsi dark brown; re-
mainder of legs light-gray pruinose; male mid tibia
expanded. Wing length 1.32-1.53 mm; veins light
brown; 5-7 setae on basal end of costa; 5-6 dorsal
and 6-8 anterior interfractural costals; costal-section
ratios: I I : I 2.0-2.4; I I I : IV 3.1-3.6; V . IV 3.4-3.7;
Mi ?
 2 index 1.6-2.0.
ABDOMEN.?Terga semiglossy dark gray. Male
terminalia: median third of posterior margin of
sternum 5 concave; anterolateral margin of sternum
5 obtusangular (115?-125?); posterolateral angle
of copulobus roundly 45? from medial corner;
copulobus regularly setose. Postgonite directed
anteriad (bent slightly anterolaterad) ; postgonite
uncus absent; pregonite projecting farther anteriad
than postgonite; distiphallus transversely expanded
proximally and near slightly convex apex; disti-
phallus lamellate ventrally and ventrally bicarinate
preapically. Anterior margin of fused surstyli un-
dulate (convex medially) ; length of fused surstyli
less than half the breadth; C-index 1.0-1.3.
TYPE.?Holotype male, USNM 70533.
TYPE-LOCALITY.?Lacoochee, Florida (VIII-18-
1930, P. W. Oman).
PARATYPES.?53 (25^0*, 28? $ ) from 13 local-
ities :
Florida: Baxter (IX-no date-1954, M. Wheeler), 5<$ $,
1 9 (USNM, ISC, and MRW); DeFuniak Springs, Walton
County (VIII-1-1962, D. L. Deonier), 1$, 1$ (USNM
and ISC); Lake Worth (VIII-8-1951, W. Wirth), 34 6,
59 9 (USNM); Orlando (IV-18-1956, W. Wirth), \$
(USNM); Winter Garden (HI-6-1942, A. Melander), 1$
(USNM).
Georgia: Waycross (VI-30-1953, M. Wheeler), 2$$,
29 9 (USNM and MRW).
Louisiana: Bethany (IX-2-1952, M. Wheeler), IS
(MRW).
Mississippi: Bluff Creek, Vancleave, Jackson County
(VI-21-1962, D. L. Deonier), 19 (ISC); Dickinson's
Pond, Lamar County (VII-24-1962, D. L. Deonier),
39 9 (DLD); gravel pit near U.S. Highway 90, sec. 20,
T.7 south, R.5 west, Jackson County (VI-9-1962, D. L.
Deonier), 19 (ISC), (VI-17-1962, D. L. Deonier), 4$ $,
29 9 (USNM and ISC); Saucier (VI-13-1953, M.
Wheeler), 1$ (MRW); Vancleave Road, Jackson County,
30?25.1' N, 88?46' W (VI-23-1962, D. L. Deonier), 6$ $,
7 9 9 (USNM and ISC); Vancleave Road, Jackson County,
NUMBER 6 8 61
30?28' N, 88?43' W (VI-28-1962, D. L. Deonier), 2$ &,
29 9 (USNM, ISC, and DLD), (VII-7-1962, D. L.
Deonier), 39 9 (USNM and ISC).
REMARKS.?Males of this species can be distin-
guished from the other members of the H. prudens
species group by their extremely short fused surstyli.
Adult. Habitats recorded: leaves of Nymphaea
odorata; stems of Juncus repens, Eleocharis wolfii,
E. mammilata, and E. tuberculosa.
Hydrellia formosa Loew
FIGURE 43
Hydrellia formosa Loew, 1861, pp. 355-356; 1862, p. 154;
1864, p. 94.?Osten Sacken, 1878, p. 202.?Becker, 1896,
p. 269 [bibliographic listing].?Howard, 1900, p. 593
[biology].?Aldrich, 1905, p. 626.?Jones, 1906, p. 185
[catalog listing].?Kahl, 1917, p. 385 [biology].?Johnson,
1925, p. 271.?Cresson, 1931, p. 104; 1936, p. 259; 1938,
p. 33; 1944b, pp. 163, 172.?Deonier, 1964, p. 115;
1965, pp. 500-506 [ecology].?Wirth, 1965, p. 744
[catalog listing].
DIAGNOSIS.?Palpus moderate yellow; 8-11 aristal
rays; body length: wing length 1.0-1.2; ocellar
absent; frons (except vertical sockets and ocellar
triangle) velvety black; notopleuron, supra-alar area,
and scutellum velvety brownish black; thoracic
pleuron contrastingly velvety brownish black and
light gray. Male length 1.27-1.53 mm; female 1.39-
1.79 mm. Male terminalia as in Figure 43.
HEAD.?Face light yellowish brown, light gray, or
white; 3-4 pfa; no sfa; epistomal index 1.1-1.5;
mesofacial index 2.0-2.8; vertex index 6.0-13.0;
A-index 2.0-2.4; ocular index 5.5-7.5. Palpus mod-
erate yellow; 8-11 aristal rays; antennal segment
3 mostly moderate yellow, 1 and 2 mostly dark brown;
frons (except glossy dark-brown ocellar triangle and
sockets of verticals) velvety black; ocellar absent;
ll-14postoculars.
THORAX.?Ppn and mesoscutum glossy brownish
black; notopleuron, supra-alar area, and scutellum
velvety brownish black; 2-3 adc and 2 pdc; upper
posterior two-thirds of mesanepisternum, entire
mesanepimeron and laterotergite light gray; re-
mainder of pleuron velvety brownish black; legs
light-gray pruinose except moderate-yellow trochan-
ters, tarsi, and at least distal third of mid and hind
tibiae. Wing length 1.36-1.96 mm; veins light yel-
lowish brown; 4-7 setae on basal end of costa; 5-7
dorsal and 5-9 anterior interfractural costals; costal-
section ratios: I I : I 2.4-2.7; I I I : IV 3.4-4.2; V: IV
3.4-4.0; M i .
 2 index 1.6-2.0.
ABDOMEN.?Terga semiglossy brownish black.
Male terminalia: median third of posterior margin
of sternum 5 concave; anterolateral margin of ster-
num 5 roundly obtusangular; copulobus acutangular
posteriorly (rounded 75?) ; posterior half of copu-
lobus irregularly setose. Postgonite bent laterad;
postgonite uncus straight, directed anterolaterad;
distiphallus gradually constricted at midlength, with
semicircular apex. Anterior margin of fused surstyli
broadly concave; B-index about 2.0; C-index 1.8-
2.2.
TYPE.?Holotype female, MCZ 11153.
TYPE-LOCALITY.?Pennsylvania (collector: Osten
Sacken).
SPECIMENS EXAMINED.?587 (217cfcf, 370? ? )
from 90 localities east of the Rocky Mountains:
Alabama: Chattahoochee State Park, Houston County
(111-24-1954, G. Steyskal), 1$, 1 $ .
Arkansas: Calion, Union County (IX-3-1952, M.
Wheeler), 1 $ ; Rison, Cleveland County (IX-3-1952,
M. Wheeler), 4$ $, 19-
Connecticut: Redding (V-23-1936, A. Melander), 1 $ ;
Storrs (VI-8-1931, A. Melander), 1 <$ ; southwest corner of
Sleeping Giant State Park, near Hamden (VII-29-1961,
J. Laffoon), 1 3 , (VII-30-1961, J. Laffoon), 2$ S, 29 9 .
District of Columbia: Rock Creek Park, Washington
(VI-15-1913, R. Shannon), 19 , (VIII-14-1931, J.
Aldrich), 1 9 ; Washington (VIII-17-1913, J. Aldrich),
5 9 9, (VIII-16-1952, J. Laffoon), IS.
Florida: Baxter (IX-no date-1954, M. Wheeler), 1 9 ;
DeFuniak Springs, Walton County (VIII-1-1962, D. L.
Deonier), 1 9 ; Fort Ogden, DeSoto County (IV-9-1952,
J. Vockeroth), 1 $ ; Silver Springs, near Ocala, Marion
County (IV-5-1953, W. Mason), 1 6\ 49 9 ; Torreya State
Park (IV-28-1952, O. Peck), 19 .
Georgia: Clayton, Rabun County, 2,000 feet (V-18-1926,
J. Bradley), 1 $ ; Waycross (VI-30-1953, M. Wheeler),
IS, 39 9 .
Indiana: LaFayette (VII-14-1915, J. Aldrich), 1 9 ,
(VII-4-1916, J. Aldrich), 2$$, 29 9 , (VI-25-1916, J.
Aldrich), 23 3, 29 9 , (VI-29-1916, J. Aldrich), 1 9 ,
(VII-7-1916, J. Aldrich), 1 9 , (VIII-31-no year, J.
Aldrich), 1 9 ; Lake County (VII-25-1950, W. Kwolek),
1 9 , (IX-15-1950, W. Kwolek), 1 9 : Valparaiso (VI-29-
1954, D. Cable), 1 $ .
Iowa: Fraser Dam, Boone County (IX-4-1960, D. L.
Deonier), 1 <$, 2 9 9 ; Siewer's Springs State Park, Dccorah
(IX-9-1961, D. L. Deonier), 1<J, 19 ; 3 miles east-south-
east of Waterville, Allamakee County (VIII-2-1960, D. L.
Deonier), 3$ <$, 19 .
Kansas: 1.5 miles south of Bonner Springs, Johnson
County (VIII-28-1962, D. L. Deonier), 19 .
Louisiana: Lake Providence (VII-14-1953, W. Wirth),
62 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
1 3 , 59 9 ; White Sulphur Springs (IX-no date-1954, M.
Wheeler), 3 3 5 , 3$ 5 .
Maine: Kennebec Point (VHI-no date-1900, G. Clapp),
Maryland: Cabin John (VI-4-1916, R. Shannon), 1 $ ;
Catoctin Furnace (VIII-9-1920, J. Aldrich), 29 9 ; Chesa-
peake Beach (VII-3-1924, J. Malloch), 1 9 ; (VIH-no
date-1926, J. Aldrich), 1 9 ; Glen Echo (VIII-12-no
year, J. Aldrich), 89 9 , (VIII-22-1922, J. Malloch), I $ ;
Pennyfield Lock of Chesapeake and Ohio Canal (VII-5-
1953, A. Stone), 2 5 5 , 59 9 ; Plummer's Island, Potomac
River (VIII-5-1913, R. Shannon), 1 9 , (IX-1-1913, W.
McAtee), 1 9 ; near Plummer's Island, Potomac River
(V-17-1914, R. Shannon), 1 9 , (X-10-1914, R. Shannon),
1 9 , (V-22-1915, R. Shannon), 2 5 5 , 39 9 ; Prince
Georges County (VII-4-1954, C. Sabrosky), 15-
Massachusetts: Boston (V-no dates, A. Melander), 1$ ,
(Vl-no dates, collector unknown), 1 3 ; Woods Hole
(VII-22-no year, A. Sturtevant), 19.
Michigan: Detroit (VI-19-1938, G. Steyskal), 1 9 ,
(IX-6-1942, G. Steyskal), 1 5 , (VII-3-1944, G. Steyskal),
29 9 ; Grand Rapids (IX-13-1937, collector unknown),
29 9 ; Midland County (VII-23-1948, R. Dreisbach), 19-
Mississippi: Bellefontaine Point, Jackson County (VII-
10-1962, D. L. Deonier), 19 ; Bluff Creek, Vancleave,
Jackson County (VI-18-1962, D. L. Deonier), 1 9 ;
Dickinson's Pond, sec. 3, T. 4 north, R. 14 west, Lamar
County (VII-11-1962, D. L. Deonier), 115 5 , 399 9 ,
(VII-24-1962, D. L. Deonier), 13 $ 5 , 27 9 9 ; Highway
59, George County, 30?45.7' N, 88?45.1' W, (VII-24-1962,
D. L. Deonier), 1 9 ; Lake Shady, Lamar County (VII -
11-1962, D. L. Deonier), 5 5 3 , 13 9 9, (VI-29-1962,
D. L. Deonier), 3 3 5 , 3 9 9 ; gravel pit near U.S. Highway
90, sec. 20, T. 7 south, R. 5 west, Jackson County (VI-9-
1962, D. L. Deonier), 25 3 ; Saucier (VI-13-1953, M.
Wheeler), 3 9 9 .
Missouri: Atherton (VIII-21-1915, C. Adams), 1 9 ,
(IX-27-1915, C. Adams), 1 9 , (VI-25-1916, C. Adams),
25 3 , 19 .
New Jersey: Trenton (VII-11-1914, collector unknown),
1 3 ; Vineland (VH-no date-1954, M. Wheeler), 19-
New York: Bear Mountain (V-18-1941, A. Melander),
1 3 , 1 9 ; Cold Spring Harbor, Long Island (VII?no dates.
A. Melander), 23 3 ; Ithaca, campus (IX-24-1925, P
Babing), 1 9 ; Ithaca (VI?no dates, collector unknown)
1 9 ; Lancaster (VII-25-1946, L. Beamer), 1 3 , (VII-
25-1946, R. Beamer), 1 3 ; 1 mile northwest of Pine Lake,
Fulton County, 1,600 feet (VIII-5-1961, D. L. Deonier),
1 9 .
North Carolina: Asheville (V-3-1957, J. Vockeroth),
1 3 ; Clingman's Dome, Great Smoky Mountains National
Park, 6,300-6,642 feet (V-20-1957, J. Vockeroth), 1 3 ,
1 9 , (V-28-1957, J. Vockeroth), 29 9 ; Highlands, 3,800
feet (V-8-1957, J. Vockeroth), 19 , (V-10-1957, J.
Vockeroth), 1 3 , (V-13-1957, J. Vockeroth), 19 , (VII-
23-1957, J. Vockeroth), 1 9 ; Highlands, Macon County,
35?8.2' N, 83? 11.3' W, 3,850 feet (VI-28-1958, J.
Laffoon), 1 9 .
Nova Scotia: Truro (no dates-1913, R. Matheson), 1 3 .
Ohio: Rome (VII-19-1946, R. Beamer), 1 3 .
Ontario: Ottawa (VIII-31-1908, J. Fletcher), 19 ,
(VI-3-1958, J. Vockeroth), 1 3 , 5 9 9 , (VIII-26-1959,
J. Vockeroth), 1 3 .
Pennsylvania: Alleghany County (VII-10-no year, H.
Smith), 2 3 3 ; Harmarville, Alleghany County (VI1-2 3-
1916, H. Kahl), 1 3 , 29 9 ; Hartstown, Crawford County
(VII-4-1921, H. Kahl), 19 ; Hills Station (VIII-30-1905,
H. Kahl), 1 9 ; Lansdalc (VII-3-1910, E. Cresson, Jr.),
2 3 3 , 1 9 ; Ohiopyle, Fayette County (VII-26-1905, H.
Kahl), 1 3 , (VII-28-1905, H. Kahl), 1 3 , 49 9 , (VIII -
7-1905, H. Kahl), 363 5 , 529 9 , (VIII-10-1905, H.
Kahl), 255 5 , 159 9 , (VIII-11-1905, H. Kahl), 55 3 ,
89 9, (VHI-no date-1907, H. Kahl), 345 5 , 3 3 9 9 ;
Pittsburgh (VIII-29-1908, H. Kahl), 2 3 5 , (VI-28-1909,
H. Kahl), 1 9 ; Point Pleasant (V-30-1914, E. Cresson,
Jr.), 1 5 ; Swarthmore (V-13-1906, E. Cresson, Jr .) , 1 9 ,
(VII-1-1908, E. Cresson, Jr.), 1 5 , 69 9 , (VI-20, VII-18,
VIII-22-1909, E. Cresson, Jr.), 8 9 9 , (V- l , VII-31-1910,
E. Cresson, Jr.), 1 5 , 39 9 , (V-16, VI-27-1920, E. Cres-
son, Jr.), 2 3 3 , 29 9 ; Westmoreland County (no dates,
H. Kahl), 1 3 , 3 9 9 .
Quebec: Lac Phillipe, 45?37' N, 76? W (VIII-22-1959,
J. Vockeroth), 19 .
South Carolina: Aiken (VI-13-1957, J. Vockeroth),
1 9 ; Cross Anchor (VII-3-1953, M. Wheeler), 2 9 9-
Tennessee: Clarksville (VII-2-1939, A Hardy), 1 9 ;
Knoxville (VIII-18-1939, D. Hardy), 19 , (IV-16-no
year, collector unknown), 1 9 ; Reelfoot State Park, Reel-
foot Lake, 36?21.1' N, 89?25.5' W (VIII-16-1962, D. L.
Deonier), 7 3 5 , 109 9 .
Texas: Carthage (IX-2-1952, M. Wheeler), 1 3 , 49 9 ;
Palmetto State Park, near Ottine (VII-10-1950, M.
Wheeler), 19-
Virginia: Alexandria (VI-14-1952, W. Wirth), 1 9 ;
Falls Church (VII-4-1950, W. Wirth), 1 9 , (VI I I -8 -
1950, W. Wirth), 1 9 , (VI-30-1951, W. Wirth), 1 9 ;
(VII-22-1951, W. Wirth), 19 , (VI-9-1954, W. Wirth),
19 , (VII-15-1954, W. Wirth), 1 5 , 1 9 , (VH-no date-
1954, M. Wheeler), 1 9 , (VI-30-1960, W. Wirth), 19 ,
(VII-24-1960, W. Wirth), 1 9 , (VIII-6-1960, W. Wirth),
19 , (VIII-8-1960, W. Wirth), 1 5 ; Great Falls (VII-
21-1962, G. Steyskal), 1 9 ; Holmes Run, Falls Church
(VI-20-1960, W. Wirth), 1 3 , (VII-1-1960, W. Wirth),
19 , (VII-3-1960, W. Wirth), 1 9 , (VII-31-1960, W.
Wirth), 1 9 ; Maywood, Alexandria County (VII-9-
1922, M. McAtee), 1 9 ; Pimmit Run (VI-3-1923, J.
Aldrich), 1 9 ; Potomac River at Scott Run, Fairfax County
VI-7-1955, C. Sabrosky), 29 9 ; Saltville, Smyth County
(V-4-1962, W. Wirth), 13-
West Virginia: Cranberry Glades, Pocahontas County
(VII-16-1955, C. Sabrosky), 29 9 ; Fairmont (VI-22-
1908, collector unknown), 1 3 .
REMARKS.?This species is the most readily dis-
tinguishable of Nearctic Hydrellia. It and H.
notata constitute a distinct group.
Adult. Habitats recorded: mats of Hydrochloa
NUMBER 6 8 63
caroliniensis, Hypericum punctatum, and Eragrostis
hypnoides; leaves of Sporobolus sp. (tidal marsh),
Pontederia cordata, and Nelumbo lutea; stems of
Eleocharis acicularis; moist lawn, riffle rocks, limnic
wrack, open sewer, old human feces, and light trap.
Kahl (1917) stated that he found H. formosa
very abundant on moist lawns in Fayette County,
Pennsylvania. Howard (1900) reported finding a few
adults on human feces.
Host plants. There is one unconfirmed rearing of
this species from wheat. I believe larvae of this spe-
cies may prefer gramineous hosts, for I collected
many from mats of water grass and Eragrostis
hypnoides.
Hydrellia gladiator, new species
FIGURES 24, 71
DIAGNOSIS.?Palpus moderate yellow; 5-8 aristal
rays; face planate; vertex index 5.0-5.4; ocular
index 3.6-4.8; male abdomen slightly convex pos-
teriorly in lateral view; female cercus rounded
basally and acute apically; antennal segment 3 at
least partly moderate yellow; apicodorsal antennal
prominent; wing length 2.21-2.45 mm. Male length
1.96 mm: female 2.30 mm. Male terminalia as in
Figure 24; female as in Figure 71.
HEAD.?Differing mainly from H. bilobifera in
the following: mesofacial index 2.0-2.5; vertex in-
dex 5.0-5.4; ocular index 3.6-4.8.
THORAX.?Differing mainly from H. bilobifera in
the following: wing length 2.21-2.45 mm.
ABDOMEN.?Terga semiglossy dark gray medially,
light gray laterally and ventrally. Male terminalia:
median third of posterior margin of sternum 5 deeply
recessed and congruent with distiphallus; antero-
lateral margin truncate and prominent at about 130?
angle from lateral margin of copulobus; anterolateral
margin with dorsal digitiform process projecting
anteromediad (covered by sternum 4 ) ; copulobus
slightly convex posteriorly (partly or wholly obscured
by loose fascicula of 6-8 strong macrochaetae);
medial margin of copulobus peninsulate (narrow digi-
tiform lobe projecting posteriad medial to postgonite)
and serially setulose; remainder of copulobus irregu-
larly setose. Postgonite bent mediad and acuminate
postgonite uncus anteromediad; distiphallus only
slightly constricted proximally and preapically, with
notched apex and lamellate on proximal two-thirds;
most of basiphallus exposed ventrally. Anterior mar-
gin of fused surstyli deeply concave medially; B-index
about 1.4; C-index 0.3-0.5. Syntergum 9 + 1 0 slightly
convex posteriorly. Female terminalia: tergum 8
much longer than sternum 8; cercus irregularly setose
dorsally and laterally, triangular with nearly semi-
circular base and about 1.6-1.7 times as long as
wide (lateral view). Segment 7 with group of 4
seriated lateral setae on posterior margin; median
spermatheca similar to that of H. discursa (Figure
70).
TYPE.?Holotype male, USNM 70534.
TYPE-LOCALITY.?Florida, reared from Vallisneria
sp. leaves from unspecified locality (XII-28-1957,
D. M. Wood).
PARATYPES.?1 $ with same data as holotype
(USNM). These are the only known specimens of
this species.
REMARKS.?This species is most similar to H.
ascita, but even here there are marked differences in
the male and female terminalia.
Hydrellia griseola (Fallen)
FIGURES 1, 3-6, 10-11, 13, 26, 130, 131, 133-137, 139-142
Notiphila griseola Fallen, 1813, pp. 250-251.?Zetterstedt,
1846, pp. 1849, 1869-1871.?Lilljeborg, 1861, pp. 205-
215; figure 2 [adult], 3 and 4 [immatures; biology].?
Becker, 1896, p. 167.
Hydrellia comniums Robineau-Desvoidy, 1830, p. 790.
Hydrellia griseola.?Macquart, 1835, p. 523.?Walker, 1856,
p. 345.?Loew, 1860, pp. 17, 22, 46; 1870, p. 7.?
Schiner, 1864 [at least in part], pp. 246-247.?Kawall,
1867, pp. 120-121 [biology].?Stein, 1867, pp. 395-397
[biology].?von Frauenfeld, 1869, p. 603 [biology].?
Brischke, 1883, p. 107.?Gobert, 1887, p. 41.?Bczzi,
1894, pp. 340-341; 1921, p. 7.?Kowarz, 1894, p. 65.?
Maragues y de Manzanas, 1894, p. 86.?Becker, 1896,
pp. 171, 180-181, pi. 4, fig. 15; 1903, p. 171; 1919, pp.
203-204; 1926, p. 68.?van de Wulp and de Meijere,
1898, p. 132.?Strobl, 1900, p. 1; 1904, p. 564.?de
Meijere, 1902, p. 685 [biology]; 1939, p. 163.?Marchal,
1903, p. 237 [biology].?Jones, 1906, p. 186 [catalog
listing].?Lampa, 1906, p. 19 [biology].?Lameere, p.
580.?Griinberg, 1910, pp. 275, 278.?Linnaniemi, 1913,
p. 45 [biology].?Sorauer and Reh, 1913, pp. 408-409
[biology].?Moreley and Atmore, 1915, p. 162 [biology].?
Hcring, 1922, p. 36 [biology]; 1925, p. 538 [biology];
1926, p. 180 [ecology]; 1937 [for many page references
see index of these volumes; ecology]; 1951 [for many
page references see index of the volume; ecology]; 1957a
[for many page references see index of the volume;
ecology].?Frost, 1924, pp. 94, 169 [biology].?Wilke,
64 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
1924, pp. 172-179 [biology].?Grimshaw, 1925, p. 19
[biology].?Hendel, 1926 [for many page references see
index of the volume; ecology]; 1933, p. 52.?Kreuter,
1927, pp. 92-98 [biology].?Collin, 1928, p. 129.?
Cresson, 1932, pp. 3, 4, 5-8, 9, 16-17, 22; 1944b, pp.
166?167, 173 [includes var. hypoleuca, obscuriceps, and
scapularis]; 1947, p. 38.?Frey, 1933, p. 83.?Vayssiere,
1933, pp. 86-87 [biology].?Seguy, 1934, p. 430; 1950,
pp. 286, 290 [biology]; 1951, pp. 96-97, pi. 5, fig. 52
[biology].?Goetghebuer, 1942, p. 8.?Grensted, 1944, p.
202.?Wahlgren, 1947, pp. 77-79 [biology].?Berg, 1950,
p. 375.?Bertrand, 1954, pp. 490-491 [biology].?Kato,
1955, pp. 11, 13 [morphology].?Wirth and Stone, 1956,
p. 469.?Hennig, 1958, p. 665, fig. 332 [morphology].?
Dahl, 1959 [for many page references see index of the
volume; ecology]; 1961, p. 44; 1964, pp. 179-187; 1967,
p. 416 [ecology].?Grigarick, 1959, pp. 1-80 [biology and
ecology].?Harrison, 1959, p. 223.?Tsacas, 1959, p.
129; 1960, p. 243.?Glick, 1960, p. 12 [ecology].?
Timon-David, 1961, pp. 228-230.?Darby, 1962, pp. 1,
137 [biology].?Fulmek, 1962, p. 37 [ecology].?LeBerre
et al., 1962, pp. 151-160 [biology].?Deonier, 1965, pp.
500, 506 [ecology].?Wirth, 1965, p. 744 [catalog listing].
Hydrellia hypoleuca Loew, 1862, p. 151.?Osten Sacken,
1878, p. 202.?Becker, 1896, p. 269 [index listing]; 1903,
pp. 171, 172.?Aldrich, 1905, p. 627 [catalog listing].?
Jones, 1906, p. 186 [catalog listing].?Washburn, 1906,
p. 80.?Cresson, 1918, p. 49.?Johnson, 1925, p. 271.?
Sturtevant, 1926, p. 10, pi. 3, fig. 26 [female internal
genitalia].
Hydrellia obscuriceps Loew, 1862, p. 152.?Osten Sacken,
1878, p. 202.?Becker, 1896, p. 269 [index listing]; 1903,
p. 171.?Slosson, 1902a, p. 8.?Aldrich, 1905, p. 627
[catalog listing].?Jones, 1906, p. 185 [catalog listing].
Hydrellia scapularis Loew, 1862, p. 153.?Osten Sacken,
1878, p. 202.?Becker, 1896. p. 269 [index listing]; 1903,
p. 171.?Coquillett, 1900, p. 461; 1904, p. 75.?Slosson,
1902b, p. 320.?Aldrich, 1905, p. 627 [catalog listing].?
Jones, 1906, pp. 160, 185 [catalog listing].?Malloch,
1915, pp. 345-346 [biology].?Cresson, 1918, p. 49.?
DeOng, 1922, p. 432 [biology].?Johnson, 1925, p. 272.
Hydrellia griseola var. hypoleuca.?Johannsen, 1935, pp.
56-57.
Hydropota griseola.?Kloet and Hincks, 1945, p. 396.
Hydrellia griseola var. scapularis.?Lange et al., 1953, pp.
8-9 [biology and ecology].
DIAGNOSIS.?Palpus moderate yellow; prementum
glossy brownish black; 5-7 aristal rays; body length:
wing length 0.8; fore tarsus dark brown; most of
mesonotum and abdominal dorsum moderate brown;
thoracic pleuron and side and venter of abdomen
light gray; epistomal index 1.2-1.8; costal section
I I : I 2.6-2.8. Male length 1.70-2.38 mm; female
1.96-2.81 mm. Male terminalia as in Figure 26.
HEAD.?Face light gray, moderate yellow, or dark
brown; 4-6 pfa; epistomal index 1.2-1.8; mesofacial
index 2.1-2.5; vertex index 7.0 or more; A-index
1.8-2.2; ocular index 4.0-8.0. Palpus moderate
yellow; 5-7 aristal rays; prementum glossy brownish
black; antenna and most of parafrontale dark brown;
frontal vitta and fronto-orbital area moderate olive-
brown; 11-15 postoculars.
THORAX.?Ppn usually light gray; mesonotum
moderate brown; 3-5 adc and 2-3 pdc; pleuron
light gray; legs light-gray pruinose except dark-
brown tarsi. Wing length 2.09-3.40 mm; veins
usually light brown; 6-8 setae on basal end of costa
proper; 8-11 dorsal and 9-12 anterior interfractural
costals; costal-section ratios: I I : I 2.6-2.8; I I I : IV
2.1-3.8; V:IV 3.4-4.2; Mi ?
 2 index 1.2-1.6.
ABDOMEN.?Terga 1-4 moderate brown medially,
light gray laterally and ventrally; tergum 5 usually
mostly light gray. Male terminalia: median third of
posterior margin of sternum 5 concave; anterolateral
corner of sternum 5 projecting and obtusangular;
posteromedial margin of copulobus concave; copu-
lobus irregularly setose. Postgonite bent anterolaterad
and postgonite uncus laterad; distal two-thirds or
so of distiphallus uniform in breadth; basiphallus
with black, spinous, lateral process. Anterior margin
of fused surstyli broadly emarginate; B-index about
2.0; C-index 6.0-9.0.
EGG.?Length 0.59?0.71 mm; maximum breadth
0.15-0.17 mm. Chorion very similar to that of H.
ischiaca (Figure 76), white, corrugate, the ridges
occasionally anastomosing and flanked by regular
perpendicular striae; sculpturing slightly less distinct
ventrally. Micropylar protuberance infundibulate,
concealed in dorsal view by a hoodlike chorionic
projection.
FIRST-INSTAR LARVA.?Length 0.33-1.13 mm;
maximum breadth 0.07-0.15 mm. Clypeal arch
obtusangular and slightly concave in lateral view;
frontoclypeal length 0.18-0.23 mm. With heavily
sclerotized supraspiracular asteriform process of 2-4
spinous projections surrounding a translucent, blunt
seta. Newly eclosed larva translucent, light yellowish
gray; late first-instar larva opaque, with yellow
tinge from labial glands and fat body.
SECOND-INSTAR LARVA.?Length 0.82-3.15 mm;
maximum breadth 0.13-0.65 mm. Frontoclypeal
length 0.28-0.33 mm. Color and other characters
similar to those in first-instar larva except supra-
spiracular asteriform process with 4-7 spinous pro-
jections (Figure 82).
NUMBER 6 8 65
THIRD-INSTAR LARVA.?Length 1.67-5.10 mm;
maximum breadth 0.23-0.75 mm. Frontoclypeus
very similar to that of H. ischiaca (Figure 100)
except dorsal phragmatal ramus black; frontoclypeal
length 0.43-0.55 mm. Ventral frontoclypeal index
3.4-3.6; phragmatal index 0.9-1.0; bifurcation
index 3.4-3.6; clypeal-arch index 1.6-1.8. Clypeal
arch slightly concave and angled about 30? in rela-
tion to lower frontoclypeal margin. Mouth-hook beak
slightly longer than base; maximum mouth-hook
base thickness about 2.0 times that of beak. First
two thoracic segments densely spinulose; abdominal
segment 8 with 4-5 annuli of spinules and ventral,
transverse row of 6 setulae; supraspiracular asteri-
form process absent. Body opaque, with yellow tinge.
PUPARIUM.?Length 3.25-5.00 mm; maximum
breadth 0.75-1.25 mm; subcylindrical. Puparial
length: minimum breadth 10.0-16.0; maximum
breadth: minimum breadth 3.0-5.2; anal-plate index
2.0-3.0. Prothoracic end semicircular in ventral
view; head-lobe scar circular; maximum puparial
breadth 5.0 times or less than diameter of head-lobe
scar and 1.8-2.0 times maximum prothoracic
breadth; anal plate subelliptical, with anterior mar-
gin straight or slightly concave. Empty puparium
translucent, light yellowish brown.
TYPES.?Notiphila griseola Fallen: Fallen appar-
ently based his original description of the species on
several syntypes. R. G. Dahl (Halsingfors, Sweden,
in litt., 1965) informed me that Fallen's collection
has been distributed among the University of Lund,
National Museum of Natural History in Stockholm,
and the British Museum (Natural History). TYPE-
LOCALITY: Fallen and his students probably collected
many of the syntype specimens near Kivik, in the
Skanor District of southern Sweden, between 1810
and 1813.
Hydrellia communis Robineau-Desvoidy: Robineau-
Desvoidy very probably used only syntypes for this
binomen. Most of them are probably in the Paris
Museum (Natural History). TYPE-LOCALITY: prob-
ably in the vicinity of Paris.
Hydrellia hypoleuca Loew: Two syntype females,
(MCZ 11158). TYPE-LOCALITY: "Middle States"
(Loew, 1862, recorded Osten Sacken as collector).
Hydrellia obscuriceps Loew: Syntype male and
female, (MCZ 11157). TYPE-LOCALITY: "Middle
States" (Loew, 1862, stated that Osten Sacken col-
lected the type specimens).
Hydrellia scapularis Loew: Syntypes in MCZ. One
female (MCZ 11155) examined. TYPE-LOCALITY:
"Middle States" (Loew, 1862, recorded Osten Sacken
as collector).
ADULT SPECIMENS EXAMINED.?Locality data for
the more than 5,000 adults examined are too
numerous for detailed presentation. Instead, these
data are summarized in Map 1.
IMMATURE SPECIMENS EXAMINED.?30 from 4
localities:
Iowa: Ames (collected and reared by D. L. Deonier);
Siewer's Springs State Park, near Decorah (collected and
reared by D. L. Deonier).
Minnesota: Mississippi River at Sucker Creek, Clear-
water County (collected and reared by D. L. Deonier) ;
Rapid River Logging Camp, Hubbard County (collected
and reared by D. L. Deonier).
DISTRIBUTION.?Authors have recorded this spe-
cies from all zoogeographic regions except the
Oriental and Australian Regions. I found it the most
abundant and widespread species in the Nearctic
Region. Workers have collected it at 285 feet below
sea level (Death Valley, California) and 9,700
feet above sea level (Mount Timpanogos, Utah) ;
January-December.
REMARKS.?I have studied specimens of the types
listed except those of Notiphila griseola Fallen and
Hydrellia communis Robineau-Desvoidy. In addition,
I have studied several Palaearctic specimens deter-
mined by Mik, Schiner, Loew, and Becker. The varia-
tions of facial color in H. griseola led many early
workers astray. Sibling species are now a possible
source of confusion. I found three such species in
this research (H. flavicoxalis, H. spinicornis, and
H. rixator), and other workers probably will find
additional ones.
Adult. Habitats recorded: leaves of oats, straw-
berries, grass, Salix sp., sagebrush, Leersia oryzoides,
Polypogon monspeliensis, Oryza sativa, Zizania
aquatic a, Glyceria grandis, Echinochloa crusgalli,
Nasturtium officinale, Polygonum scabrum, celery,
potatoes, and Nuphar advena; flowers of Eupatorium
perfoliatum, choke-cherry, dandelion, wild plum,
clover, Heracleum lanatum, Ranunculus aquatilis,
and R. longirostris; sea beach, tamarack bog, cow
pasture, lawn, limnic wrack, hot spring, gravel and
sandbars, mud flat, stream-riffle rocks, algal mat,
fronds of Lemna minor, sedge meadow, reed marsh,
stems of Eleocharis sp., fronds of Riccia sp., and mats
of Eragrostis hypnoides.
66 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
MAP 1.?Distribution of Hydrellia griseola. Each dot represents one or more localities within
the county or district.
Glick (1960) collected 500 adults at 200 feet alti-
tude in an aerial survey over parts of Louisiana,
Arkansas, and Mississippi. He collected very few of
any other species of Hydrellia. This can possibly be
correlated with the greater dispersal potential of
H. griseola, viz., factors such as their larger wings
and frequent high-population densities.
Grigarick (1959) found the adult stadium to be
108 days (male) and 139 days (female) at 55?-81?
F. for specimens collected in July, 130 days (male)
and 135 days (female) at 50?-74? F. for October
specimens, and 54 days (male) and 49 days (female)
at 70? F. for February specimens. These represented
maxima. He fed the specimens 10 percent sucrose
and 5 percent yeast hydrolysate solution; therefore,
his ranges of the maxima for the adult stadium prob-
ably should be regarded as physiological rather than
ecological, especially in consideration of the findings
of Le Berre et al. (1961), who found adult longevity
to be directly correlated with the chemical composi-
tion of the food. Also, Grigarick showed in detail the
effect of temperature and atmospheric saturation
deficit on the physiological stadium. The percent
mortality varied from 0 in 48 hours at 0.3 mm Hg
saturation deficit and 58? F., to 50 in 48 hours at
0.5 mm Hg and 80? F., and then to 100 in 3 hours
at 31.9 mm Hg saturation deficit and 100? F. With
adults under constant high humidity, he obtained
the following thermal limits to activity:
110? to 114C F:
107? to 110? F:
104? to 107? F:
heat death
heat paralysis
dropping
NUMBER 6 8 67
98? to 104? F: heat rest
52? to 92? F: normal locomotion
40? to 50? F: slow walking
Of 12 field-collected adults kept at 29? F. and sup-
plied with food, 50 percent died after 34 days.
Considering these and other factors, the mean eco-
logical stadium may be 30 to 40 days in most
Nearctic habitats.
I observed adults attacking and feeding on adult
chironomids, adult tipulids, adult psyllids, and adult
and immature collembolans. The collembolans were
free when captured, but the others were trapped in
the surface film. One adult fly pulled a small adult
chironomid from the surface film and carried it off
in a short flight.
Laurence (1952) listed as observed prey of H.
griseola in Great Britain a symphypleonan species
and an arthropleonan species (Collembola), a spe-
cies of Braconidae (Hymenoptera), and the follow-
ing Diptera: Sciara sp., an orthocladiine species,
Bibio marci, Bibio sp., Tachydromia agilis, Leptocera
crassimana, Musca autumnalis, and Sarcophaga sp.
Laurence found both sexes feeding on some or all of
these insects. He stated that the canalicular teeth of
the labella are carnassial in Hydrellia. These teeth
appeared similar to those in several other Hydrellia
species examined. Grigarick observed adults feeding
on collembolans, psyllids, mayflies, dragonflies, and
adults of their own species. He saw adults searching
over the surface film and grasping and manipulating
various floating objects. Adults readily fed on banana
flakes, fish meal, raw hamburger, and yeast hydrolysate
with 10 percent sucrose solution presented in the field.
In my survey, the gut contents of freshly killed speci-
mens were greenish yellow and granular. Examina-
tion of sections of 125 adults revealed granular con-
tents, some of which were macrophytic cells and
diatoms. Culture of the gut contents from 15 adult
males and 15 adult females show the presence of
some aquatic yeasts.
I observed some instances of agonistic behavior in
adults. In one of these, the adults adopted a defen-
sive posture in the presence of Ochthera mantis
(Ephydridae), and some made short rushes toward
the latter. I did not gather enough data to establish
the importance of O. mantis as a predator on H.
griseola adults, but in a few instances it stalked some
adults and very probably (as observation on H.
cruralis populations showed) caught and fed on
them. Grigarick considered lycosid spiders as impor-
tant predators of adults. He witnessed several cap-
tures by these spiders. In June and July, he caught
more lycosid spiders than H. griseola adults in float-
ing traps. Predators of several kinds can prey more
readily on freshly emerged adults and ovipositing
females.
In one of my observations, a pair copulated for
15 minutes. During this time three other males at-
tempted to mate with the female, first grasping her
head, then her wing, and finally grasping the copu-
lating male. When the third one intruded, the copu-
lating male dismounted and the newcomer tried to
mount the female, but she rejected him. Grigarick
found copulation time ranged from 1 to 50 minutes.
Both male and female exhibited wing scissoring as
the conspicuous epigamic behavior. Mating occurred
on both floating and erect leaves during daylight
and even in overcast weather. In Grigarick's labora-
tory, pairs of the same age copulated as early as 3
days and as late as 70 days after emergence. The
same pairs sometimes mated repeatedly in one day,
and some mated through five consecutive days. One
male inseminated at least three females. One female
deposited viable eggs 93 days after isolation.
In my observations and those of Grigarick, females
preferred to oviposit on floating leaves, but they
would oviposit on leaves about 20 cm above the
water. The female deposited eggs in loose aggrega-
tions parallel with the long axis of the leaf blade.
Balachowsky and Mesnil (1935) reported that
females oviposited on the basal part of the leaf blade
of young barley. This is probably the preferred site
on emergent plants because of shade and micro-
environmental humidity. Grigarick found 52 eggs
on one leaf blade and observed oviposition in the
field every month of the year at temperatures above
50? F. In his laboratory, females started ovipositing
5 days after emergence while caged with males of
the same age at temperatures from 54? to 72? F.
Oviposition rates at 72? F. for field-collected females
were 18 eggs in 2.5 hours, 28 in 24 hours, 40 in 48
hours, and 70 in 5 days. The maximum number of
eggs per female varied from 73 in 52 days at 58? F.
to 199 in 60 days at 72? F. No oviposition occurred
at 43? F.
In one experiment, Grigarick fed 5 percent yeast
hydrolysate to a female that had been isolated and
fed 10 percent sucrose for 108 days after she last
68 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
oviposited. She did not oviposit during isolation,
but she laid viable eggs 3 days after taking the yeast
hydrolysate. Le Berre et al. (1961) reported the
following experimental data on oviposition:
1. mean survival of 2 days for females given only water;
2. females survived long enough to lay several eggs
when fed sucrose or honey in water;
3. female survival and fecundity increased distinctly with
a milk diet;
4. total number of eggs per female on milk diet was 88;
total number on milk and sucrose or honey was 116
and 119;
5. delays in oviposition were directly related to the
chemical composition of the food;
6. uninseminated females laid fewer eggs than insemi-
nated females.
EGG.?Grigarick reported incubation periods of
1.9 days at 90? F., 2.3 at 80?, 2.9 at 72?, 7.5 at 58?,
and 17.8 days at 50? F. Of 80 eggs exposed to 29?
F. and then removed to 80? F., 80 percent hatched
after 24 hours exposure, 20 percent hatched after
48 to 120 hours exposure, and none hatched after
148 hours exposure to 29? F. Normal incubation
periods obtained for submerged eggs. Unfertilized
eggs collapsed soon after deposition.
LARVA.?According to Grigarick, each larval sta-
dium varied from 2 to 3 days at 80? F. The larval
stadia totaled 6.9 days at 90? F., 7.6 at 80?, 8.1 at
72?, 21.1 at 58?, and 41.1 days at 50? F. These fig-
ures were based on the time 50 percent completed
development. Larvae eclosed from the micropylar
end of the egg after slitting the chorion with their
mouth-hooks. Grigarick observed some larvae that
remained within the opened chorion from minutes
to hours. He stated that generally the larvae begin
mining shortly after eclosion or even after partial
eclosion. I think this would be so only under optimal
conditions. First-instar larvae required 2-2.5 hours
at 80? F. for making and entering a new mine, while
third-instar larvae required only 0.5-1.5 hours for
this task at 80? F. With mining third-instar larvae
under constant high humidity, Grigarick obtained
the following thermal limits to activity:
112? to 114? F:
104? to 1110 F:
95? to 103? F:
50? to 90? F:
36? to 50? F:
heat death
heat paralysis
agitated movement
normal mining and locomotion
quiescence or very slow movement
In one mortality experiment in which ten larvae of
each instar under constant high humidity were ex-
posed to 100? F., Grigarick obtained 70, 80, and 100
percent mortality in first, second, and third instars
after 24 hours exposure and uniformly 100 percent
after 48 hours exposure. In a similar experiment, at
29? F., he obtained 20, 70, and 80 percent mortality
in first, second, and third instars after 24 hours
exposure and 80, 100, and 100 percent mortality
after 48 hours exposure. That these last results do
not agree with the fact of overwintering of larvae
in subboreal and boreal habitats strongly suggests
that harmful low temperatures are countered by the
larvae entering diapause.
In my observations and in those of Grigarick,
Lilljeborg (1861), Balachowsky and Mesnil (1935),
and others, the larvae mined mostly in leaves, espe-
cially those in, on, or near water; however, I found
many in leaf sheaths and several in the culm proper.
Contrary to Hering's (1951, p. 203) statement that
"The short blotch-mine of such species [H. griseola
and H. butomi] always remains above water", I
often found larvae and puparia in submerged plant
tissues, especially in Zizania aquatica. I found as
many as six first-instar larvae in a wild-rice leaf and
Grigarick found as many as 15 to 30 first-instar larvae
in one grass leaf. Wilke (1924) reported finding
commonly over 40 larvae mining in one barley leaf.
PUPARIUM.?The time between pupariation and
adult emergence varied from 8 to 9 days for three
larvae in my laboratory. Grigarick obtained the fol-
lowing temperature-related variations in length of
the puparial phase (called pupa by Grigarick) : 5.0
days at 90? F., 6.1 at 80?, 7.0 at 72?, 18.0 at 58?, and
33.8 days at 50?. At each of these temperatures, a
higher saturation deficit increased the puparial
phase or prevented emergence. Exposing new puparia
to 28? F. for 5 days followed by removal to 72? F.
resulted in only 50 percent emergence. No adults
emerged after 10 days exposure of puparia to 28? F.
In the observations of most authors, the late third-
instar larvae often sought a new mine for puparia-
tion. These mines varied much in size and were
usually blotch-shaped. Pupariation occurred toward
the center of the mine where the larva anchored its
spiracular peritremes in the plant tissue. Attachment
to plant tissue is not necessary for pupariation, for
Grigarick observed pupariation on moist soil, sand,
blotting paper, and glass; Stormer and Kleine (1911)
and Kuwayama (1955) found puparia in soil; and
Burghele (1959a) collected thousands of puparia
NUMBER 6 8 69
from the bottom of small, shallow, ice-covered pools
in November, January, and February at Ghencea,
Yugoslavia.
HOST PLANTS.?In Table 1 are listed 40 grami-
neous genera reported as containing host plants for
this species. Additionally, various authors have re-
ported hosts in 20 nongramineous genera. Most of
the latter probably represent incidental puparial
hosts; however, I found all three larval instars and
puparia in Nasturtium officinale, and Taylor (1928)
reported miners of H. nasturtii [as H. ranunculi] in
that plant species. I observed that the larvae mined
in leaves and moved up through linear mines in the
stem cortex between successively younger leaves.
They usually pupariated in the leaf axils. Because of
the usual rapid growth of water cress and perhaps
other factors, the larvae seemed not to disturb or
harm the plants permanently.
Hydrellia griseola has attained its economic status
because of its gramineous preference. Intermittent
infestations of barley and oats in Europe prompted
much of the early biological investigation of this fly.
Lilljeborg (1861) reported heavy infestations of
late-sown barley near coastal lowlands in Sweden.
Stormer and Kleine (1911), Wilke (1924), and
Grimshaw (1925) reported similar infestations of
barley in northern Europe. Balachowsky and Mesnil
(1935) reported that H. griseola developed more
rapidly in barley than in oats and had almost no
mortality in barley but up to 42 percent mortality in
oats. DeOng (1922) first reported the serious infes-
tations of domestic rice in California. In 1953, Lange
et al. estimated the fly species destroyed 10 to 20
percent of the California rice crop with a probable
loss of $16 million. Grigarick (1959) listed the fol-
lowing four conditions that increase susceptibility to
infestation of rice with H. griseola: (1) low soil fer-
tility and low temperatures weakening plants and
retarding growth; (2) strong wind and wave action
keeping plants down; (3) excessive water depth,
through rain or mismanagement, retarding growth;
(4) dense algal mats retarding emergent growth of
plants. These conditions, by restraining rapid plant
emergence, kept the rice in positions favorable for
the rice leaf miner.
Grigarick estimated a maximum of 11 overlapping
generations of H. griseola in the Sacramento Valley.
Two generations occurred in January-April on spe-
cies of Polypogon, Avena, Hordeum, and other wild
grasses in rain pools and irrigated rice fields and
three generations in May-June on rice and asso-
ciated grasses, mainly young Echinochloa crusgalli.
Four slightly overlapping generations occurred in
June-September on Polypogon monspeliensis and
Echinochloa crusgalli in cool water inlets to rice
fields and two generations in September-January
on several late-summer grass species along streams
and canals and on fall grasses in fall rain pools.
Primarily because of high temperatures, larvae in-
fested rice plants only rarely after the middle of July.
Parasites. Forty-three species of Hymenoptera re-
ported as having been reared from H. griseola are
listed in Table 2. Grigarick reported Chorebus
aquaticus, Opius hydrelliae, and Halticoptera sp. as
the most abundant species, overall. He found that
the parasite-density maximum immediately followed
the host-density maximum. The first generation of
H. griseola had 50 to 60 percent parasitism. The
third generation (first generation on rice) had 1.0
to 2.5 percent parasitism, but some later generations
had 80 to 87 percent parasitism by July. Opius
hydrelliae and Chorebus aquaticus showed the high-
est densities at that time.
Hydrellia harti Cresson
FIGURES 15, 69
Hydrellia harti Cresson, 1936 [in part], p. 262; 1944b, pp.
170, 175.?Deonier, 1964, pp. 115, 125, fig. 4; 1965,
pp. 500, 506 [ecology].?Wirth, 1965, p. 744 [catalog
listing].
DIAGNOSIS.?Palpus moderate yellow; 6-8 aristal
rays, face planate; vertex index 5.0-6.0; male
abdomen often slightly bilobate posteriorly; female
cercus less than 1.5 times as long as wide; antennal
segments dark brown; apicodorsal antennal promi-
nent. Male length 1.79-1.96 mm; female 1.70-2.31
mm. Male terminalia as in Figure 15; female as in
Figure 69.
HEAD.?Differing mainly from H. bilobifera in the
following: vertex index 5.0-6.0; antennal segment
3 dark brown; fronto-orbital area and frontal vitta
darker than most of paraf rontale.
THORAX.?Differing mainly from H. bilobifera in
the following: 3^4 adc and 2 pdc; legs mostly light-
gray pruinose except moderate-yellow tarsi; 5-7
dorsal and 8-11 anterior interfractural costals; wing
length 1.70-1.97 mm.
70 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
ABDOMEN.?Terga semiglossy dark gray medially,
light gray laterally and ventrally. Male terminalia:
median third of posterior margin deeply, rectangu-
larly recessed and congruent with distiphallus;
anterolateral margin of sternum 5 rounded through
about 160? with conspicuous dorsal, rounded process
projecting anterolaterad; copulobus slightly convex
posteriorly, without macrochaetae, with seriated
setulae on middle third of medial margin, and with
mediolateral triangular lobe. Postgonite bent mediad
and postgonite uncus anteriad; distiphallus con-
stricted proximally and preapically and bicarinate
ventrally; carinae partly lamellate; most of basi-
phallus exposed ventrally. Anterior margin of fused
surstyli deeply and broadly concave; B-index about
1.5; C-index 0.4-0.6. Syntergum 9 + 1 0 often incon-
spicuously bilobate posteriorly. Female terminalia:
tergum 8 much longer than sternum 8; cercus irregu-
larly setose dorsally and laterally, triangular with
trapezoid, hooked base, and less than 1.5 times as
long as wide (lateral view). Segment 7 with group
of about 7-8 seriated lateral setae on posterior mar-
gin; median spermatheca similar in size and shape
to that of H. discursa (Figure 70).
TYPE.?Holotype male, ANSP 2078.
TYPE-LOCALITY.?Havana, Illinois (IX-19-1895,
C. A. Hart) .
SPECIMENS EXAMINED.?47 (38o*d\ 9 ? $ ) from
11 localities:
California: Laguna Canyon, Orange County (IX-no
date-1922, J. Needham), 1 3 .
Iowa: Spring Lake, Greene County (IX-19-1961, D. L.
Deonier), 1 3 , 39 9.
Kansas: Kansas University Natural History Reservation
near Lawrence, Douglas County (VI-7-1963, D. L.
Deonier), 2 3 $.
Mexico: Ciudad Mante, Tamaulipas (VII-7-1960, H.
Hovvden), 1 $.
Nebraska: Glen (VHI-no date-1960, W. Rapp), 1$ .
Ontario: Grand Bend (VII-10-1939, G. Shewell), 1 3 ;
Marmora (VIII-7-1952, J. McAlpine), 43 3 , (VIII-25-
1952, E. Smith), 43 3 , (IX-8-1952, E. Smith), 33 3 ,
(IX-9-1952, J. McAlpine), 5 3 3 ; Midland (VIII-18-
1955, J. Chillcott), 1 3 ; Ottawa (VII-2-1952, J.
McAlpine), 1$.
Quebec: Perkin's Mills (VIII-25-1949, G. Shewell),
133 3 , 5 ? 9 .
Rhode Island: Providence (IX-16-no year, collector
unknown), 1 $ .
REMARKS.?Adult habitats recorded: leaves of
Potamogeton nodosus, P. epihydrus, and Nuphar
ad vena; limnic wrack and sedge meadow.
Hydrellia idolator, new species
FIGURE 21
DIAGNOSIS.?Palpus moderate yellow; 8-10 aristal
rays; vertex index 4.5-5.5; ocular index 13.0-17.5;
wing length 1.53-2.13 mm; prementum pale yellow;
antennal segment 3 moderate yellow (contrasting
with 1 and 2). Male length 1.50-1.70 mm; female
1.45-1.53 mm. Male terminalia as in Figure 21.
HEAD.?Face pale yellow; 6-7 pfa; epistomal
index 1.3-2.0; mesofacial index 2.6-3.5; vertex
index 4.5-5.5; A-index 1.5-2.2; ocular index 13.0-
17.5. Palpus moderate yellow; 8-10 aristal rays;
prementum pale yellow; apicodorsal antennal promi-
nent; lower third of postocular area light gray, upper
two-thirds light brown; frons (except light-brown
fronto-orbital area) dark gray; 13-15 postoculars.
THORAX.?Ppn and mesonotum moderate brown;
3-4 adc and 2 pdc; pleuron light gray except upper
fifth light brown; legs light-gray pruinose except
trochanters, tibiae, and tarsi moderate yellow. Wing
length 1.53-2.13 mm; veins light brown; 5-7 setae
on basal end of costa; 6-8 dorsal and 6-10 anterior
interfractural costals; costal-section ratios: I I : I 2.0-
2.5; I I I : IV 2.8-3.2; V: IV 3.3-3.8; M, *2 index
1.3-1.7.
ABDOMEN.?Terga semiglossy dark brown medi-
ally, but with alternate light-gray and dark-brown
lateral wedges and light-gray ventral lobes. Male
terminalia: median third of posterior margin of
sternum 5 deeply concave and congruous with disti-
phallus; anterolateral margin of sternum 5 smoothly
rounded; copulobus roundly acutangular posteriorly
(about 60? from lateral corner) and irregularly
setose; 2 falciform lateral projections from above
copulobus. Postgonite bent slightly anterolaterad and
postgonite uncus directed anterolaterad; distiphallus
uniformly tapering to ovoid apex; basiphallus not
nearly covered laterally by surstyli. Anterior margin
of fused surstyli narrowly notched medially (for
about one-fifth of length); B-index about 5.0;
C-index 1.6-2.0.
TYPE.?Holotype male, CNC 10896.
TYPE-LOCALITY.?Ottawa, Ontario, Canada (VII -
2-1947, G.E. Shewell).
PARATYPES.?4 (1 d\ 3 ? $ ) from 4 localities:
Mississippi: Bellefontaine Road, Jackson County (VI-
12-1962, D. L. Deonier), 13 (ISC); gravel pit near
U.S. Highway 90, sec. 20, T. 7 south, R. 5 west, Jackson
County (VI-17-1962, D. L. Deonier), 19 (USNM).
NUMBER 6 8 71
Ontario: Ottawa (VIII-24-1938, G. Shewcll), 19
(USNM).
Quebec: Norway Bay (VIII-24-1938, G. Shewell), 19
(ISC).
REMARKS.?This species is very easily confused
with H. atroglauca, and since no specimen of the
latter was available for male terminalia study, con-
specificity is possible. H. atroglauca, however, has a
moderate orange-yellow prementum, an ocular index
of 7.0-12.0, and 8-10 dorsal and 10-13 anterior
interfractural costals.
Adult. Habitats recorded: mats of Hydrochloa
caroliniensis and sedges.
Hydrellia insulata, new species
FIGURE 46
DIAGNOSIS.?Palpus moderate yellow; prementum
glossy brownish black; 8-10 aristal rays; mesofacial
index 2.2-2.6; antenna velvety dark brown; thoracic
pleuron moderate olive-brown except anterior fourth
of mesanepistemum, and mesokatepisternum light
gray; costal section V:IV 3.5-4.5. Male length 1.70-
2.30 mm; female 2.04-2.47 mm. Male terminalia as
in Figure 46.
HEAD.?Face light yellowish brown or light gray;
4-6 pfa; epistomal index 1.4-1.8; mesofacial index
2.2-2.6; vertex index 8.5-12.5; A-index 2.0-2.4;
ocular index 5.5-6.5. Palpus moderate orange-
yellow; prementum glossy brownish black; 8-10
aristal rays; antenna velvety dark brown; fronto-
orbital area and frontal vitta moderate olive-brown;
most of parafrontale dark brown; 13-17 postoculars.
THORAX.?Ppn and postalar bridge densely mod-
erate olive-brown pruinose; mesoscutum densely
strong yellowish-brown pruinose; 3-4 adc and 2 pdc;
pleuron moderate olive-brown except anterior fourth
of mesanepistemum, and mesokatepisternum light
gray; legs light-gray or moderate olive-brown
pruinose except dark-brown tarsi. Wing length 1.79?
2.47 mm; veins dark brown; 6-5 setae on basal end
of costa; 8-11 dorsal and 10-13 anterior interfrac-
tural costals, costal-section ratios: II:I 2.0?2.4;
III:IV 2.8-3.4; V:IV 3.5-4.5; Mi ?
 2 index 1.2-
1.6.
ABDOMEN.?Terga dark gray medially, light brown
laterally, and light gray ventrally. Male terminalia:
median third of sternum 5 concave; anterolateral
margin of sternum 5 acutangular (about 75?);
posteromedial margin of copulobus concave and
undulate with seriated setae; remainder of copulobus
usually nonsetose. Postgonite concealed and directed
posteriad, with postgonite uncus bent anterolaterad;
distiphallus uniform in breadth to papillate apex.
Anterior margin of fused surstyli with narrow, shal-
low notch medially in broad notch; B-index about
3.0; C-index 3.0-3.6.
TYPE.?Holotype male, USNM 70535.
TYPE-LOCALITY.?Plummer's Island, Potomac
River, Maryland (IX-1-1962, K. V. Krombein).
PARATYPES.?14 (7cTd\ 7 9 2 ) from 1 locality:
Maryland: Plummer's Island, Potomac River (VIII-3?
1962, K. Krombein), 4$$, 39 9 , (VHI-7-1962, K.
Krombein), 2$ $, 3 9 9 , (IX-1-1962, K. Krombein), 1$,
19 (USNM, ISC, and DLD).
Hydrellia ischiaca Loew
FIGURES 44, 76, 82, 100, 122, 126, 131
Hydrellia ischiaca Loew, 1862, pp. 150-151.?Becker, 1896,
p. 269 [index listing].?Jones, 1906, p. 185 [catalog list-
ing]?Johnson, 1925, p. 271.?Cresson, 1944b, pp. 165,
173.?Deonier, 1964, p. 116; 1965, pp. 500, 506 [ecology].
?Wirth, 1965, p. 744 [catalog listing].
DIAGNOSIS.?Palpus moderate yellow; 8-10 aristal
rays; body length:wing length 0.8-0.9; fore coxa
and tibia moderate yellow; 2-3 black anterior apical
setae on hind tibia; thoracic pleuron light gray ex-
cept upper fourth of mesanepistemum light yellowish
brown. Male length 1.53-2.04 mm; female 1.75-
2.23 mm. Male terminalia as in Figure 44.
HEAD.?Face moderate yellow or yellowish gray;
5-7 pfa; epistomal index 1.4-1.6; mesofacial index
2.2-2.5; vertex index 7.5-12.0; A-index 1.6-2.0;
ocular index 6.5-7.5. Palpus moderate yellow; 8-10
aristal rays; antenna and most of parafrontale dark
brown; frontal vitta and fronto-orbital area light
gray with green reflection; 13-16 postoculars.
THORAX.?Ppn, most of anterior part of meso-
notum, and notopleuron light gray; remainder of
mesonotum moderate brown; 3-4 adc and 2 pdc;
pleuron light gray except upper fourth of mesane-
pistemum light yellowish brown; 1-2 basal coxals
(1 macrochaetous); legs light-gray pruinose except
moderate-yellow coxae, trochanters, fore tibia, mid
and hind tarsi, and dark-brown fore tarsus; 2-3
black anterior apical setae on hind tibia; 2 black
anterior basal setae on hind tarsus. Wing length
72 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
1.77-2.43 mm; veins dark brown or moderate olive-
brown; 6-8 setae on basal end of costa; 6-8 dorsal
and 7-12 anterior interfractural costals; costal-
section ratios: I I : I 2.7-3.0; I I I : IV 2.1-2.6; V:IV
3.0-3.9; M i .
 2 index 1.4-1.6.
ABDOMEN.?Terga moderate brown medially and
light gray laterally and ventrally. Male terminalia:
median third of posterior margin of sternum 5
prominent (roundly right-angled); posteromedial
margin of copulobus angular from medial hooklike
corner; copulobus with seriated setae on postero-
medial and medial margins, otherwise bare. Post-
gonite bent anterolaterad and postgonite uncus
laterad; distal two-thirds of distiphallus uniform in
breadth. Anterior margin of fused surstyli with a
narrow, shallow U-shaped notch medially; spicate
projection posterolaterally on surstyli; B-index about
2.0; C-index 5.0-8.0.
EGG.?Length 0.59-0.71 mm; maximum breadth
0.15-0.17 mm. Chorion (Figure 76) white, cor-
rugate, with longitudinal ridges occasionally anas-
tomosing and flanked by regular perpendicular striae.
Sculpturing distinct dorsally, indistinct ventrally.
Micropylar protuberance infundibulate, concealed
dorsally by hoodlike chorionic projection.
FIRST-INSTAR LARVA.?Length 0.33-1.10 mm;
maximum breadth 0.07-0.15 mm. Clypeal arch dis-
tinctly obtusangular and slightly concave in lateral
view; frontoclypeal length 0.18-0.23 mm. With
heavily sclerotized supraspiracular asteriform process
of 2-4 spinous projections surrounding a translucent,
blunt seta. Newly eclosed larva translucent, yellowish
gray; late first-instar larva opaque with yellow tinge
from labial glands and developing fat body.
SECOND-INSTAR LARVA.?Length 0.80-3.15 mm;
maximum breadth 0.15-0.65 mm. Frontoclypeal
length 0.28-0.33 mm. Other characters similar to
those in first-instar larva except supraspiracular
asteriform process with 4-7 spinous projections
(Figure 82).
THIRD-INSTAR LARVA.?Length 1.75-5.10 mm;
maximum breadth 0.30-0.90 mm. Frontoclypeal
length 0.43-0.55 mm; dorsal phragmatal ramus
partly hyaline (Figure 100). Ventral fronto-clypeal
index 3.4-3.6; phragmatal index 0.9-1.0; bifurca-
tion index 3.4-3.6; clypeal-arch index 1.6-1.8.
Clypeal arch angled 30?. Mouth-hook beak slightly
longer than base; maximum mouth-hook base thick-
ness about 2.0 times that of beak. Supraspiracular
asteriform process absent; prothorax and mesothorax
densely spinulose; abdominal segment 8 with ven-
tral, transverse row of 6 setulae and 4-5 annuli of
spinules. Body opaque, with yellow tinge.
PUPARIUM.?Length 3.35-5.00 mm; maximum
breadth 0.75-1.15 mm; subcylindrical (Figure 122).
Puparial length: minimum breadth 15.0-18.0; maxi-
mum breadth: minimum breadth 4.0-6.0; anal-plate
index 2.0-3.0. Prothoracic end semicircular in ven-
tral view; head-lobe scar circular; maximum
puparial breadth 5.0 times or less than diameter of
head-lobe scar and 1.8-2.0 times maximum pro-
thoracic breadth; anal plate subelliptical, with
anterior margin straight or slightly concave. Empty
puparium translucent light yellowish brown.
TYPE.?Holotype female, MCZ 11154.
TYPE-LOCALITY.?"Middle States." Loew (1862)
stated that Osten Sacken was collector.
ADULT SPECIMENS EXAMINED.?328 (136 cf d, 192
$ $ ) from 91 localities:
Alaska: Valdez (VII-15-1948, G. Marks), 19.
California: Lake Tahoe (VIII-11-1940, L. Lipovsky),
2? 9.
Connecticut: Colebrook (VIII-28-1941, A. Melander),
1$ ; Redding (VI-3-1933, A. Melander), 1$ .
Georgia: Holcomb Creek (VIII-1-1957, W. Richards),
IS; Villa Rica (VII-5-1953, M. Wheeler), 1$ .
Indiana: Lafayette (VIII-1-1916, J. Aldrich), 1 $ ,
(VIII-18-1916, collector unknown), 1 $.
Illinois: Peoria (VIII-26-1917, J. Aldrich), \$.
Iowa: Ames, Story County (VI-22-1960, D. L. Deonier),
2 S S, 1 9 ; Ames, Izaak Walton League Reserve, Story
County (VI-9-1960, D. L. Deonier), IS, (VII-1-1960,
D. L. Deonier), 2$ S, 3 9 9 , (VII-7-1960, D. L. Deonier),
19 , (VTI-12-1960, D. L. Deonier), 1<$, 29 9, (VII-17-
1960, D. L. Deonier), 1 9 , (VII-18-1960, D. L. Deonier),
19 , (VIII-31-1960, D. L. Deonier), IS; Lake Odessa,
Louisa County (VIII-9-1960, D. L. Deonier), 2 9 9 ; Ledges
State Park, Boone County (V-16-1947, J. Laffoon), (VII -
10-1960, D. L. Deonier), 1 S ; Little Wall Lake, Hamilton
County (VIII-27-1960, D. L. Deonier), 1 9 ; Pilot Knob
State Park, Hancock County (VI-15-1960, D. L. Deonier),
1 S ; Siewer's Spring State Park, Decorah (IX-9-1960, D. L.
Deonier), \$; Spring Lake, Greene County (IX-19-1961,
D. L. Deonier), 29 9 ; White Pine Hollow State Park,
Dubuque County (VIII-2-1960, J. Laffoon), 2$ S ; 4
miles northeast of Wapello, Louisa County (VIII-9-1960,
D. L. Deonier), 2$ $, 29 9 ; 3 miles southeast of Water-
ville, Allamakee County (VIII-2-1960, D. L. Deonier),
IS, 49 9 .
Maine: Pittston (VIII-3-1930, A. Melander), IS, 19 .
Maryland: Baltimore (V-16-1938, E. Fisher), 1,5, 1 9 ;
Beltsville (V-21-1922, J. Malloch), 19 ; Glen Echo (VII-
20-1924, J. Malloch), 1 9 ; Plummer's Island, Potomac
NUMBER 6 8 73
River (VIII-5-1913, R. Shannon), 1 9 , (IX-5-1915, W.
McAtee), 19 ; near Plummer's Island, Potomac River
(V-22-1915, R. Shannon), 1 9 , (IX-2-1915, R. Shannon),
1 9 .
Massachusetts: Brookline (VI-17-no year, C. Johnson),
1 9 ; New Bedford (no dates, A. Melander), 1,$.
Michigan: Isabella County (IX-16-1955, R. Dreisbach),
1 9 ; Midland County (VIII-16-1936, R. Dreisbach), 1 9 ;
Roscommon County (VII-1-1950, R. Dreisbach), 1 9 ;
Vineyard Lake, Jackson County (VII-26-1954, G. Steyskal),
It.
Minnesota: Biological Station, Itasca State Park ( V I -
26-1963, D. L. Deonier), 3$ 3 , 7 9 9 ; Bohall Lake, Itasca
State Park (VIII-17-1963, D. L. Deonier), 1 3 ; Bohall
Trail Bog, Itasca State Park (VIII-17-1963, D. L.
Deonier), 2 3 3 , 3 9 9 ; Douglas Lodge Bay, Lake Itasca,
Itasca State Park (VI-20-1963, D. L. Deonier), 9 3 3 ,
159 9 ; headwaters of LaSalle Creek, Itasca State Park
(VIII-6-1963, D. L. Deonier), 1 9 ; Kabekona Creek, Hub-
bard County (VIII-8-1963, D. L. Deonier), 1 3 , 1 9 ; Iron
Corner Lake, Itasca State Park (VI-18-1963, D. L.
Deonier), 1 3 , 2 9 9 ; Mississippi River, sec. 34, T. 145 north,
R. 36 west, Clearwater County (VII-8-1963, D. L. Deonier)
1 3 , 1 9 ; Mississippi River near north entrance Itasca State
Park (VII-9-1963, D. L. Deonier), 2 9 9 ; Mississippi
River, 150 yards north of Highway 31, Clearwater County
(VII-9-1963, D. L. Deonier), 1 3 , 4 9 9 ; Mississippi River
at Sucker Creek, Clearwater County (VII-9-1963, D. L.
Deonier), 1 3 , 8 9 9 ; Rapid River Logging Camp, Hubbard
County (VII-2-1963, D. L. Deonier), 303 3 , 349 9 ;
Squaw Lake, Itasca State Park (VIII-14-1963, D. L. Deo-
nier), 1 3 , 1 9 ; Sucker Creek, 100 yards north of Highway
31, Clearwater County (VII-23-1963, D. L. Deonier), 1 9 ;
Two Island Lake, Itasca State Park (VI-28-1963, D. L.
Deonier), 2 9 9 ; 6.5 miles east of Waubun (VIII-18-1963,
D. L. Deonier), 2 3 3 , 2 9 9 ; west side across from Bio-
logical Station, Lake Itasca, Itasca State Park (VII -28-
1963, D. L. Deonier), 1 3 , 3 9 9 -
New Hampshire: Mount Washington (no dates?1917, A.
Slosson), 1 9 .
New York: Dryden Lake, Tompkins County (VII I -8 -
1961, D. L. Deonier), 1 9 ; Ithaca (VI11-28-1894, collec-
tor unknown), 1 3 ; vicinity of Six-Mile Creek, Ithaca
(VIII-9-1961, J. Laffoon), 13 ; McLean (VII-7-1919, C.
Johnson), 1 9 ; New York (V-31-1923, A. Sturtevant), 1 9 ;
east foot of West Notch Mountain, Hamilton County, 43?
20.6' N, 74?37.2' W at 1,900 feet (VIII-6-1961, D. L. Deo-
nier), 1 3 , 1 9 .
North Carolina: Clingsman Dome, Great Smoky Moun-
tains National Park, 6,300-6,800 feet (V-28-1957, J. Vocke-
roth), 1 9 ; Forney Ridge, Great Smoky Mountains National
Park (VI-26-1941, A. Melander), 1 9 ; Highlands, Macon
County, 3,800 feet (V-10-1957, J. Vockeroth), 2 9 9 , ( V I -
3-1957, J. Vockeroth), 1 3 ; Highlands, Macon County,
3,000 feet (VI-1-1957, W. Mason), 2 3 3 , 1 9 ; Looking
Glass Peak, Pisgah National Forest (VII-19-1957, W. Rich-
ards), 1 3 ; Wayah Gap, Macon County, 4,000 feet (VII I -
16-1957, J. Chillcott), 1 3 ; Wilson's Gap, Highlands, 3,100
feet (V-12-1957, J. Vockeroth), 1 3 , (V-25-1957, J.
Vockeroth), 1 3 , 1 9 ; 3 miles southwest of Glenville, Jack-
son County, 3,700 feet (VI-19-1958, J. Laffoon), 2 3 3 .
Nova Scotia: Jordan Falls (VIII-5-1958, J. Vockeroth),
19 ; Lockeport (VIII-9-1958, J. Vockeroth), 19 ; Mount
Uniacke (VIII-5-1958, J. Vockeroth), 2 3 3 , 2 9 9 .
Ontario: Ancaster (VI-26-1955, O. Peck), 1 3 ; Mar-
mora (VII-5-1951, J. McAlpine), 1 9 , (VIII-4-1952, J.
McAlpine), 1 9 ; Normandale, 42?42' N, 80?19' W ( V -
27-1956, J. Vockeroth), 1 3 ; Ottawa (VIII-8-1923, C.
Curran), 1 9 .
Pennsylvania: Alleghany County (VH-no dates, H.
Smith), 1 3 ; Castle Rock (V-28-no year, collector un-
known), 1 3 ; Jack Run, Alleghany County (VI-14-1908,
collector unknown), 1 3 , 1 9 ; Ohiopyle, Fayette County
(VII-22-1905, H. Kahl), 1 9 , (VII-28-1905, H. Kahl),
1 3 , (VIH-2-1905, H. Kahl), 1 3 , 4 9 9 , (VIII-7-1905,
H. Kahl), 2 3 3 , (VIII-11-1905, H. Kahl), 1 9 , (VHI-no
date-1907, H. Kahl), 8 3 3 , 3 9 9 ; Point Pleasant ( V - 3 0 -
1914, collector unknown), 1 9 ; Swarthmore (V-28-1907,
collector unknown), 1 9 , (VH-no date-1908, collector un-
known), 1 3 , 1 9 , (VI-13-1909, E. Cresson, Jr.), 1 3 ,
(VIII-21-1910, E. Cresson, Jr.), 1 9 ; Westmoreland
County (VHI-no date-1907, H. Kahl), 6 3 3 , 159 9 ; 2
miles north of Narberth, Montgomery County ( IX-14-
1915, E. Cresson, Jr), 1 3 -
Quebec: Laniel (VII-3-1944, A. Brooks), 1 9 ; Old
Chelsea (VIII-31-1958, J. Vockeroth), 1 9 ; Rigaud (VIII -
3-1919, J. Ouellet), 1 3 -
South Carolina: Cross Anchor (VII-3-1953, M. Wheel-
er), 6 3 3 , 6 9 9 .
Tennessee: Greenbrier Cove, Great Smoky Mountains Na-
tional Park, 2,000 feet (V-18-1957, J. Vockeroth), 2 9 9 ;
Knoxville (VIII-17-1939, R. Beaman), 1 3 ; Shelby Forest
State Park (IX-3-1952, M. Wheeler), 1 3 .
Texas: Huntsville (IV-11-1952, M. Wheeler), 1 3 -
Virginia: Fairfax (VII-19-1954, M. Wheeler), 1 3 ;
Falls Church (V-26-1951, W. Wirth), 4 3 3 , 2 9 9 , ( V I I -
15-1954, W. Wirth), 1 3 , 1 9 ; Maywood, Alexandria ( V -
21-1922, W. McAtee), 1 3 , 2 9 9 , (VII-9-1922, W. Mc-
Atee), 1 9 ; Reddish Knob, Augusta County (VIII -29-
1953, W. Wirth), 2 3 3 , 5 9 9 .
West Virginia: Cranberry Glades (VII-16-1955, C. Sa-
brosky),2 3 3 .
I M M A T U R E S P E C I M E N S E X A M I N E D . ? 3 6 0 collected
and reared by D. L. Deonier from 9 localities:
Minnesota: Biological Station, Lake Itasca, Itasca State
Park; Douglas Lodge Bay, Lake Itasca, Itasca State Park;
Mississippi River, sec. 34, T. 145 north, R. 36 west, Clear-
water County; Mississippi River at Sucker Creek, Clearwater
County; Mississippi River, 150 yards north of Highway 31,
Clearwater County; Rapid River Logging Camp, Hubbard
County; Sucker Creek, 100 yards north of Highway 31,
Clearwater County; 6.5 miles east of Waubun.
North Carolina: Harris Lake, Highlands, Macon County.
REMARKS.?Despite the obvious dissimilarities be-
tween the male terminalia, this species must be
74 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
placed in the H. griseola species group. The imma-
ture stages are nearly identical in H. griseola and
H. ischiaca, both in morphology and bionomics. The
adults differ mainly in color characters.
Adult. Habitats recorded: leaves of Zizania
aquatica, Glyceria grandis, G. obtusa, Leersia
oryzoides, Potamogeton natans, P. gramineus, P.
nodosus, Nuphar advena, Nymphaea tuberosa, and
Nasturtium officinale; sedge meadow and mud flat.
Considering the close relationship of H. griseola
and H. ischiaca, one could suppose that the data on
the physiological stadium of H. griseola would apply
fairly well to H. ischiaca. If this is true, then the
ecological adult stadium of this species must be con-
siderably shorter than the physiological one, for I
observed no end-of-summer buildup in population
density. The one density maximum occurred during
July, in the region of Itasca State Park, Minnesota.
This could be explained by a greater emergence of
adults at that time resulting from ecological effects
on oogenesis, oviposition rate, egg viability, and de-
velopment rate, or it could have been due to slight
generation overlap. I tentatively estimated the mean
ecological adult stadium of this species in subboreal
and boreal habitats to be between 20 and 30 days.
This species, like most of its congeners, is poly-
phagous. Contrary to Berg's (1950) report on the
leaf feeding of H. cruralis, I did not observe H.
ischiaca feeding on the leaves in its microhabitat.
Normally, the adult diet consisted of periphytic
microbiota such as yeasts, Chrysophyta, and Pro-
tozoa, nectar, and insect tissue fluids (especially of
other Hydrellia specimens). During one observation,
a female H. ischiaca probed a dead microcrustacean.
After feeding for a few seconds, the female repeat-
edly touched her abdominal venter to the micro-
crustacean. In another pertinent observation, 40
adults that emerged on 21 July were caged on 22
July in a large culture dish containing clean rice
leaves in tap water. On 23 July most of the flies
were walking over the surface film and the leaves,
and two specimens were probing and feeding on
other dead specimens in the surface film; all speci-
mens died by 26 July, without ovipositing.
I observed little distinct epigamic behavior in this
species. In some observations the male circled the
female closely, and in others the male moved to and
fro or sideways in front of the female before attempt-
ing to mount. In one observation, copulation lasted
for 10 minutes. The female walked intermittently
during copulation. Modes of species and sex recog-
nition fail occasionally, for I observed a male H.
nobilis attempting to mount a female H. ischiaca.
The latter responded by quickly advancing upon the
protagonist and striking out with its fore tarsi. On
another occasion, a male H. nobilis tried to mount a
male H. ischiaca. I observed agonistic behavior in
H. ischiaca several times. In one instance, a female
advanced toward a specimen of Chorebus aquaticus
ovipositing on a rice leaf. On another occasion, a
male struck at a female H. nobilis. This could pos-
sibly have been misdirected epigamic behavior.
Although it is probable, as in some congeners, that
the females oviposit on stubble, twigs, and plants
other than hosts, I observed ovipositing and found
egg masses only on Z. aquatica and the grasses
Glyceria grandis, striata, and obtusa. As in Gri-
garick's (1959) observations of H. griseola, the
females showed notable preference for floating leaves
of wild rice (especially the leaf apices) as oviposition
sites. At the end of the summer, when floating leaves
were uncommon, females oviposited on the leaves
hanging nearest to the water surface.
Egg. I collected and observed the eggs only on the
upper leaf surfaces. Of 20 egg masses collected and
isolated for rearing, the mean number of eggs per
mass (an irregularly spaced aggregation) was 8 and
the range 3-11. It is possible that more than one
female contributed to some of the egg masses.
The incubation period ranged from 2 to 7 days in
the laboratory for 47 eggs in a temperature range of
58? to 75? F. The mean period was 4 days. The
ecological incubation period probably ranges from
2 to 4 days in subboreal and boreal habitats.
Larva. The first larval stadium ranged from 3 to
5 days with a mean of 4 for 20 larvae in a tem-
perature range of 61?-81? F. The newly hatched
larvae moved about actively and soon attempted to
mine into the substrate leaf. Many of these first
attempts were unsuccessful and the larvae randomly
crawled or sank to nearby leaves where they at-
tempted to mine. Occasionally, I found as many as
six first-instar larvae in the same leaf. The second
larval stadium ranged from 3 to 4 days with a mean
of 4 days for 11 larvae in a temperature range of
64?-80? F. The third larval stadium ranged from
3 to 9 days, with a mean of 5 days for 9 larvae in a
temperature range of 67?-81? F.
NUMBER 6 8 75
Any one larva usually required more than one leaf
for development through the three instars because
extensive mining depleted the mesophyll and dam-
aged many veins. A leaf mined by a third-instar
larva was usually yellowish and the apical half often
completely devoid of mesophyll.
Puparium. The puparial phase ranged from 4 to
20 days, with a mean of 7 for 51 puparia in a tem-
perature range of 58?-82? F. The time from larval
eclosion to adult emergence range from 13 to 29
days for 20 specimens. The time from oviposition to
adult emergence ranged from 15 to 35 days for 15
specimens. The third-instar larvae finished puparia-
tion within 8 hours. The puparia were readily
apparent in the partially mined leaves. Rarely, I
found puparia in the stems. Often the puparia had
their spiracular peritremes embedded in the plant
epidermis, but some were lying unattached between
the leaf surfaces. I found puparia oriented in various
attitudes to the leaf axis, but commonly they were
parallel to it. In one leaf, I found six puparia, two
of which were contiguous and parallel.
Host plants. On the basis of frequency and dis-
tribution of larvae, I found the preferred host plant
to be Zizania aquatica during summer in the region
of Itasca State Park, Minnesota. Glyceria grandis
and G. obtusa were the only other species in which
I found larvae during summer. I reared H. ischiaca
from G. obtusa and G. grandis collected from areas
0.5-20 miles from Z. aquatica. I found one puparium
in a floating leaf of Potamogeton natans near in-
fested Z. aquatica, and a third-instar larva pupari-
ated in a floating leaf of P. natans in the laboratory.
Since both of these lacked extensive feeding mines,
I considered them incidental puparial hosts. The
negative results of repeated searches for H. ischiaca
in a stand of Potamogeton richardsonii in a strong
current about 1 meter from a stand of heavily in-
fested rice growing in quiet water supported this
assumption.
I measured the percent of infestation of Z. aqua-
tica with H. ischiaca in three samples from Lake
Itasca, Minnesota. In the first sample taken at the
west shore, opposite the Biological Station, 28 July,
33 percent of 23 plants showed obvious infestation.
Of the infesting immatures, 90 percent were in parts
on or above the water surface. The second sample
of 36 plants collected from a lush stand at the same
locality on 13 August had 50 percent infested. The
third sample of 27 plants taken 19 August from a
sparse stand of Z. aquatica and Scirpus sp., exposed
to the prevailing westerly wind and waves near the
Biological Station, had 89 percent infestation.
Earlier in the season, on 20 June, at Douglas Lodge
Bay, Lake Itasca, I found eggs on the floating leaves
of 13 of a sample of 25 rice plants and larvae in 20
plants of the same sample. On 2 July at Rapid
River Logging Camp, Hubbard County, Minnesota,
I found an infestation of about 70 percent in a
sample of about 50 plants from one small riverside
stand.
Parasites. From 132 puparia reared or collected,
50 hymenopterans emerged. The time from host
pupariation to emergence of the adult hymenopteron
ranged from 6 to 42 days. The mean time in the
laboratory was 8 days. During the early spring of
1967, 12 braconids and one pteromalid emerged from
37 puparia reared from second- or third-instar
larvae.
I reared the following Hymenoptera from H.
ischiaca puparia: Chorebus aquaticus, Chorebidella
bergi, Chaenusa, new species (all Braconidae), and
Trichopria columbiana (Diapriidae). There has
been some conjecture that Trichopria columbiana
is a hyperparasite. I reared a single specimen of this
species under the following circumstances: a third-
instar larva collected as a second instar on 9 July,
pupariated, and on 15 July the puparium containing
a late, white, fly pupa was transferred to a large test
tube. On 27 August, when the diapriid emerged,
an examination showed a pupal exuviae (not that of
the fly host) within the original puparium.
On 28 July at the west shore of Lake Itasca,
opposite the Biological Station, I collected 28 adult
Chorebus aquaticus in about 5 minutes from a stand
of Z. aquatica. This indicated the high-population
density of this parasite. At this time also, I observed
a specimen of C. aquaticus ovipositing repeatedly
for 5 minutes in a rice leaf free of eggs and larvae of
Hydrellia. Chaenusa sp. (probably new) parasitized
three of four H. ischiaca puparia reared from eggs
collected only 2 days after they were deposited. The
single specimen of Opius sp. emerged anteroven-
trally from the puparium.
Overwintering. The grasses Glyceria grandis and
striata serve as overwinter hosts for several species
of Hydrellia, including H. griseola and H. ischiaca.
Some Glyceria grandis pass the winter as relatively
76 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
short, green shoots submerged in moderately swift,
shallow streams. If these streams freeze partly or
completely, the grass and the Hydrellia larvae appar-
ently tolerate the condition. In 1967 I collected 14
second- and 30 third-instar larvae in these grasses
from the date of lake thaw (15 April) to date of
first wild adult collected (23 May). Burghele (1959a)
reported finding viable puparia of griseola on the
bottoms of small, ice-covered pools in Rumania. My
searches through ten large samples of old rice stubble
revealed no larvae or puparia.
Many samples of infested grasses were from three
small streams, all within 1,000 meters of wild-rice
beds in Lake Itasca. The rapid infestation of young
rice in the lake is thereby understandable.
Hydrellia itascae, new species
FIGURES 28, 102, 110, 129
DIAGNOSIS.?Palpus moderate orange-yellow; 6-9
aristal rays; body length:wing length 1.0-1.1; an-
tenna velvety dark brown; apicodorsal antennal
prominent; 2 basal coxals; costal section I I : I 1.4?
2.2; 12-14 dorsal and 14-16 anterior interfractural
costals. Male length 2.89-3.06 mm; female 2.98-
3.23 mm. Male terminalia as in Figure 28.
HEAD.?Face light yellowish brown; 6-7 pfa; 2
sfa; epistomal index 1.1-1.4; mesofacial index 1.5-
1.9; vertex index 4.8-6.2; A-index 1.4-1.8; ocular
index 4.8-6.2. Palpus moderate orange-yellow; 6-9
aristal rays; apicodorsal antennal prominent; an-
tenna velvety dark brown; frons dark brown; 15-17
postoculars.
THORAX.?Ppn semiglossy dark gray; mesonotum
semiglossy dark brown; 3-4 adc and 3-4 pdc;
pleuron yellowish gray except upper fifth of mesane-
pimeron and anterior half of mesokatepisternum
light brown; 2 basal coxals (1 macrochaetous); legs
light-gray pruinose except dark-brown tarsi. Wing
length 2.72-3.15 mm; veins dark brown; 10-11
setae basal end of costa; 12-14 dorsal and 14-16
anterior interfractural costals; costal-section ratios:
I I : I 1.4-2.2; I I I : IV 2.5-2.9; V:IV 3.0-3.5; Mi*s
index 1.1-1.5.
ABDOMEN.?Terga semiglossy dark brown medi-
ally, light gray laterally and ventrally. Male ter-
minalia: median third of posterior margin of sternum
5 concave; anterolateral margin of sternum 5 roundly
right-angled; copulobus angular posteromedially
(about 100? from lateral corner) and irregularly
setose, with verrucate anteromedial lobe. Postgonite
directed anterolaterad; postgonite uncus absent;
distiphallus uniform in breadth (all but apex cov-
ered by fused surstyli). Anterior margin of fused
surstyli deeply cleft medially; B-index about 0.8.
THIRD-INSTAR LARVA.?Body of larva not exam-
ined. Frontoclypeal length 0.50-0.60 mm; cheliform
spot arms removed from clypeal arch by their length
(Figure 102). Ventral frontoclypeal index 2.2-2.8;
phragmatal index 0.8-0.9; bifurcation index 5.8-
6.5; clypeal-arch index 1.8-2.2. Clypeal arch slightly
concave along angle of about 25?-30? in relation to
lower frontoclypeal margin. Mouth-hook beak and
base lengths subequal; maximum mouth-hook base
thickness about 1.5 times that of beak.
PUPARIUM.?Length 4.50-5.00 mm; maximum
breadth 1.00-1.50 mm; subcylindrical; suddenly
tapering from abdominal segment 7 posteriad (Fig-
ure 110). Prothoracic end slightly convex in ventral
view; head-lobe scar obovoid or subcircular; maxi-
mum puparial breadth 5.5-6.0 times maximum
breadth of head-lobe scar and 2.3 times or less
than maximum prothoracic breadth; prothorax with
many ventral rugulae; lateral spinules inconspicuous
in midlength of puparium; abdominal segment 8
without ventral, transverse row of setulae and with
only few nonannular, distinct spinules; anal plate
subrectangular, with anterior margin slightly convex.
Empty puparium opaque moderate brown.
TYPE.?Holotype male, USNM 70536.
TYPE-LOCALITY.?University of Minnesota Biolog-
ical Station, Lake Itasca, Itasca State Park, Clear-
water County, Minnesota (VI-26-1963, D. L.
Deonier).
PARATYPES.?16 (4cf o*, 12 $ $ ) from 2 localities:
Minnesota: Douglas Lodge Bay, Lake Itasca, Itasca State
Park (VI-20-1963, D. L. Deonier), \$ (DLD); Prof
Green Trail, Lake Itasca, Itasca State Park (VII-25-1963,
D. L. Deonier), 1$ (DLD); University of Minnesota Bio-
logical Station, Lake Itasca, Itasca State Park (VI-20-
1963, D. L. Deonier), 5 ? 9 , (VI-26-1963, D. L. Deonier),
99 9 (USNM and ISC).
IMMATURE SPECIMENS EXAMINED.?Nine, from
Douglas Lodge Bay, Lake Itasca, Itasca State Park,
Minnesota (collected and reared by D. L. Deonier).
REMARKS.?This species may be confused with
H. manitobae in a macroscopic field examination.
Adult. Habitats recorded: leaves of Zizania aqua-
tica and Nuphar advena.
NUMBER 6 8 77
Host plants. I reared adults from nine puparia
found in Potamogeton zosteriformis.
Parasites. I reared Aphanta sp. (probably new)
and Chorebidella bergi (all Hymenoptera: Braconi-
dae) from three of the puparia.
Hydrellia lata Cresson
Hydrellia lata Cresson, 1944b, p 165.?Wirth, 1965, p. 744
[catalog listing].
DIAGNOSIS.?Palpus moderate yellow; prementum
light gray; 8-10 aristal rays; ocular index 6.5-8.0;
mesofacial index 1.5-1.9; apicodorsal antennal
prominent; costal section II:I 2.4-2.8; 10-12 an-
terior interfractural costals. Female length 2.30-2.72
mm.
HEAD.?Face light yellowish brown or yellowish
gray; 5-6 pfa; 1-2 sfa; epistomal index 1.1-1.4;
mesofacial index 1.5-1.9; vertex index 4.2-S.6;
A-index 1.5-1.9; ocular index 6.5-8.0. Palpus mod-
erate yellow; 8-10 aristal rays; prementum light
gray; apicodorsal antennal prominent; antenna dark
brown; fronto-orbital area and frontal vitta mod-
erate brown; most of parafrontale velvety dark
brown; 15-18 postoculars.
THORAX.?Ppn and mesonotum moderate brown;
2-5 adc and 2 pdc; pleuron light gray, with some
light-brown areas on mesanepistemum and mesokate-
pisternum; legs light-gray pruinose except dark-
brown tarsi. Wing length 2.82-3.06 mm; veins light
brown; 6-7 setae on basal end of costa; 6-9 dorsal
and 10-12 anterior interfractural costals; costal-
section ratios: II:I 2.4-2.8; III:IV 2.5-3.0; V:IV
3.0-3.4; Mi ?
 2 index 1.3-1.8.
ABDOMEN.?Terga 1-4 semiglossy dark brown
dorsally; tergum 5 dark-brown pruinose dorsally;
ventral tergal lobes light gray.
TYPE.?Holotype female, ANSP 6655.
TYPE-LOCALITY.?Nasel River, Pacific County,
Washington (VII-15-1922, A. L. Melander).
SPECIMENS EXAMINED.?4? $ from 3 localities:
Alaska: King Salmon, Naknek River (VII-31-1952, W.
Mason), 1 9.
Manitoba: Eastern Creek near Churchill (VII-9-1952, J.
Chillcott), 19 .
Washington: Nasel River, Pacific County (VII-15-1922,
A. Melander), 2 9 9 .
REMARKS.?According to Cresson (1944b), this
species is apparently closely related to the H.
proclinata species group. I think that the male will
be found to be very similar to that of either H.
platygastra or H. serena.
Hydrellia luctuosa Cresson
FIGURES 64, 92, 107
Hydrellia luctuosa Cresson, 1942, p. 78; 1944b, pp. 170, 171.
?Berg 1950, pp. 375, 376, 377, 384, 385 (pi. 2, fig. 1),
387, 388, 389 (pi. 3, fig. 1), 396 [biology, morphology, and
taxonomy of immatures].?Deonier, 1964, p. 117; 1965,
p. 500, 506 [ecology].?Wirth, 1965, p. 744 [catalog
listing].
DIAGNOSIS.?Palpus dark brown or black; 6-8
aristal rays; ocular index 4.6-5.4; face dark brown,
centrally protuberant; male mid tibia expanded;
9-12 anterior interfractural costals; body generally
dark olive-brown. Male length 1.79-1.97 mm;
female 1.87-2.04 mm. Male terminalia as in Figure
64.
HEAD.?Face dark brown and centrally protuber-
ant; 4-5 pfa; 0-3 dorsally inclined sfa; epistomal
index 1.0-1.3; mesofacial index 1.4-1.9; vertex
index 3.5-7.0; A-index 1.7-2.0; ocular index 4.6-
5.4. Palpus dark brown or black; 6-8 aristal rays;
antenna brownish black; fronto-orbital area and
frontal vitta moderate olive-brown; most of para-
frontale velvety black; 14-16 postoculars.
THORAX.?Ppn and mesonotum moderate or dark
olive-brown; 3-4 adc and 3--4 pdc; pleuron
moderate olive-brown except light-gray tinge on
mesokatepisternum and mesanepimeron; legs very
sparsely light-gray pruinose except dark-brown tarsi;
male mid tibia expanded. Wing length 1.70-2.13
mm; veins dark brown; 5-8 setae on basal end of
costa; 6-8 dorsal and 9-12 anterior interfractural
costals; costal-section ratios: II:I 2.1-3.6; III:IV
2.4-2.8; V:IV 3.0-3.4; M, ?
 2 index 1.2-1.5.
ABDOMEN.?Terga densely moderate olive-brown
pruinose. Male terminalia: median third of posterior
margin of sternum 5 concave and congruent with
apex of distiphallus; anterolateral margin of sternum
5 about 100? angular; copulobus acutangular
posterolaterally; copulobus about twice as broad
posteriorly as anteriorly and regularly setose, with
2-3 distinct fusiform macrochaetae on posterolateral
corner. Postgonite bent anteriad and postgonite
uncus laterad; distiphallus tapering distally and
78 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
carinate ventrally. Anterior margin of fused surstyli
ovoid; B-index about 1.2; C-index 1.8-2.3.
SECOND-INSTAR LARVA.?Length 1.75-3.20 mm;
maximum breadth 0.20-0.40 mm. Frontoclypeal
length 0.26-0.31 mm. Other characters similar to
those in third-instar larva except body light yellowish
gray.
THIRD-INSTAR LARVA.?Length 3.00-4.20 mm;
maximum breadth 0.40-0.60 mm. Frontoclypeal
length 0.40-0.55 mm; feeding apparatus mostly dark
(Figure 92). Ventral frontoclypeal index 5.0-5.5;
phragmatal index 1.2-1.3: bifurcation index 3.6-
4.0; clypeal-arch index 1.8-2.0. Clypeal arch sloping
at about 20? except for slight hump dorsal to labial
gland orifice. Mouth-hook beak only about 0.9 of
base length; maximum mouth-hook base thickness
about 2.0 times that of beak; mouth-hook with light
spot. Prothorax and mesothorax sparsely spinulose;
prothorax distinctly rugulose in ventral view; abdom-
inal segment 8 without ventral, transverse row of
setulae, but with 3 annuli of spinules. Body trans-
lucent, with bright green pigment showing through.
PUPARIUM.?Length 2.65-3.50 mm; maximum
breadth 0.85-1.10 mm; fusiform (Figure 107).
Puparial length: minimum breadth 8.8-10.0; maxi-
mum breadth: minimum breadth 2.6-3.6; anal-plate
index 2.5-3.0. Prothoracic end nearly semicircular
in ventral view; head-lobe scar subcircular, its maxi-
mum breadth distinctly less than minimum puparial
breadth; maximum puparial breadth in abdomen;
anal plate subelliptical, with anterior margin slightly
convex. Empty puparium translucent, light brown.
TYPE.?Holotype male, ANSP 6620.
TYPE-LOCALITY.?Be.-sey Creek, Cheboygan Coun-
ty, Michigan (VIII-14-1941, C. O. Berg).
ADULT SPECIMENS EXAMINED.?131 (39o"cf,
94$ ? ) from 22 localities:
Iowa: South Pond, Ledges Park, Boone County (VII-
10-1960, D L. Deonier), 13, 19 ; Springbrook State Park,
Gutbrie County (VI-23-1961, D. L. Deonier), \$ ; Spring
Lake, Greene County (IX-19-1961, D. L. Deonier), 1 3 ,
3 9 9 .
Kansas: Kansas University Natural Reservation, near
Lawrence, Douglas County (VI-10-1963, D. L. Deonier),
33 <J ,599 .
Michigan: Bessey Creek, Cheboygan County (VIII-14-
1941, C. Berg), 59 9 ; Cheboygan County (VII-15-1946,
C. Berg), 19 , (VII-26-1946, C Berg), 23 3 , 169 9 ,
(VII-21-1947, C. Berg), 14 , (VIII-4-1947, C. Berg),
23 3 , 1 9 , (VIII-7-1947, C. Berg), 1 3 , 1 9 ; Douglas
Lake, Cheboygan County (VII-3-1941, C. Berg), 1 9 ;
Nigger Creek, Cheboygan County (VIII-21-1941, C. Berg),
1$.
Minnesota: Bohall Lake, Itasca State Park (VIII-17-
1963, D. L. Deonier), 1 3 , 1 9 ; Chamber's Creek, Itasca
State Park (VII-18-1963, D. L. Deonier), 53 3 , 219 9 ;
Douglas Lodge Bay, Lake Itasca, Itasca State Park (VII-
20-1963, D. L. Deonier), 5$ 3, 89 9 ; Eagles Nest (VI-
25-1959, W Balduf), 1 9 ; Headwaters of LaSalle Creek,
Itasca State Park (VIII-6-1963, D. L. Deonier), 1 9 ;
Mississippi River near north entrance Itasca State Park
(VII-9-1963, D. L. Deonier), 4 3 3 , 6 9 9 ; Mississippi
River at Sucker Creek, Clearwater County (VII-9-1963,
D. L. Deonier), 1 9 ; Squaw Lake (VIII-14-1963, D. L.
Deonier), 23$; Two Island Lake (VI-28-1963, D. L.
Deonier), 19 , (VIII-B-1963, D. L. Deonier), 53 3,
119 9 ; 6.5 miles east of Waubun (VIII-18-1963, D. L.
Deonier), 1 3 , 3 9 9 .
Ontario: Dundas Marsh (no dates-1948, W. Judd), 1 3 ;
Marmora (VHI-7-1952, J. McAlpine), 23 3, 39 9 ;
Ottawa (VI1-2-1947, G. Shewell), 19 .
Quebec: Mount Albert (VII-24-1954, J. Martin), 2 9 9.
IMMATURE SPECIMENS EXAMINED.?42 from 3
localities:
Michigan: Cheboygan County (collected and reared by
C.Berg).
Minnesota: Douglas Lodge Bay, Lake Itasca, Itasca State
Park; Squaw Lake, Itasca State Park (all collected and
reared by D. L. Deonier).
REMARKS.?This species probably belongs to the
H. prudens group.
Adult. Habitats recorded: leaves of Sparganium
chlorocarpum, Nymphaea tuberosa, and Potamogeton
natans; stems of Eleocharis palustris. I found 11
specimens congregated on a floating Scirpus stem.
On Two Island Lake, Itasca State Park, Minnesota,
I collected 17 specimens by the lighted-receptacle
method and found several specimens resting on
clumps of Eleocharis palustris. The adult population-
density of this species increased by perhaps 50 per-
cent 24 hours after a 20-cm rise in this lake.
In the laboratory, an adult of Lispe sp. killed and
fed on a female H. luctuosa. In the field, a female
H. luctuosa tried to feed on a chironomid captured
by an adult H. pulla, but the latter quickly repulsed it.
Larva and puparium. The time from just after
eclosion from the egg to the first molt was 10 days
for two larvae. The second larval stadium ranged
from 4 to 6 days for five larvae. The third larval
stadium ranged from 2 to 6 days and the puparial
phase from 6 to 10 days for eight specimens. The
total time from just after eclosion from the egg to
adult emergence ranged from 22 to 32 days. I found
NUMBER 6 8 79
only first-instar larvae at Douglas Lodge Bay, Lake
Itasca, 20 June, and only puparia at Squaw Lake,
Itasca State Park, 14 August. These data, along with
the observation of a sudden rise in adult population-
density, may indicate distinct generations in this
species.
Host plants. I found immatures only in
Potamogeton zosteriformis, but Berg (1950) found
the order of preference to be Potamogeton alpinus,
P. zosteriformis, P. richardsonii, P. amplifolius, and
P. natans.
Parasites. Berg (1950) listed Trichopria colum-
biana (Diapriidae) and Dacnusa sp. (Braconidae)
as parasites of the puparia of H. luctuosa.
Hydrellia manitobae, new species
FIGURE 48
DIAGNOSIS.?Palpus dark brown; 6-8 aristal rays;
vertex index 3.5-4.5; wing length 3.06-3.40 mm;
thoracic pleuron mostly light gray; 11-14 anterior
interfractural costals; 2 basal coxals; male mid tibia
moderately expanded. Male length 2.50-3.16 mm;
female 2.89-3.18 mm. Male terminalia as in Figure
48.
HEAD.?Face light-brown pruinose over dark gray;
4-6 pfa; 4-5 sfa; epistomal index 1.2-1.5; meso-
facial index 1.8-2.2; vertex index 3.5-^.5; A-index
1.3-1.7; ocular index 4.5-8.2. Palpus dark brown;
6-8 aristal rays; antenna and frons (except mod-
erate olive-brown fronto-orbital area) dark brown;
13-18 postoculars.
THORAX.?Ppn and mesonotum light gray; 4-5
adc and 2?4 pdc; pleuron light gray except anterior
half of mesokatepisternum, laterotergite, and
mesomeron dark gray; 1-2 mesokatepisternals (1
macrochaetous); 2 basal coxals (1 macrochaetous);
legs light-gray pruinose except dark-brown tarsi;
male mid tibia moderately expanded. Wing length
3.06-3.40 mm; veins light yellowish brown; 7-10
setae on basal end of costa; 9-11 dorsal and 11-14
anterior interfractural costals; costal-section ratios:
II:I 2.4-2.8; III:IV 2.8-3.2; V:IV 3.8-4.4; Mi ?
 3
index 1.0-1.5.
ABDOMEN.?Terga semiglossy dark gray. Male
terminalia: median third of posterior margin of
sternum 5 concave; anterolateral margin roundly
right-angled; copulobus nearly truncate posteriorly
and irregularly setose. Postgonite bent anterolaterad
and postgonite uncus posterolaterad; distiphallus
constricted at midlength and carinate ventrally.
Anterior margin of fused surstyli convex with medial
indentation; B-index about 1.8; C-index 0.8-1.3.
TYPE.?Holotype male, USNM 70537.
TYPE-LOCALITY.?LaTrappe, Quebec, Canada
(VII-14-1934, J. Ouellet).
PARATYPES.?45 (8o*cf, 37? ? ) from 16 locali-
ties:
Alaska: Anchorage (VI-19-1921, J. Aldrich), 19
(ANSP); King Salmon, Naknek River (VII-31-1952, W.
Mason), 1? (CNC), (VIII-11-1952, W. Mason), 19
(USNM).
Alberta: One Four (VI-1-1956, O. Peck), 1 9 (USNM).
Idaho: Sand Point (VIII-27-1918, A. Melase), 19
(USNM).
Maine: Princeton (VII-1-1912, collector unknown), 19
(ANSP).
Manitoba: Farnsworth Lake near Churchill (VII-14?
1952), J. Chillcott), 19 (ISC); Fort Churchill (VI-23-
1952, J. Chillcott), 1 9 (CNC); Whitewater Lake (VI-22-
1958, R. Bird), 39 9 (USNM and ISC).
Ontario: Marmora (VIII-6-1952, Boyle), 16\ 29 9
(CNC); Ottawa (Vll-no date-1915, C. Johnson), IS
(ISC).
Quebec: Missisquoi Bay (VII-28-1936, G. Shewell),
3 5 S, 49 9 (CNC, USNM, and ISC); Rigaud (VII-27-
1926, J. Ouellet), 1$, 1 9 (ISC); Rupert House (VII-30-
1949, E. LeRoux), 15 , 19 (USNM and DLD).
Saskatchewan: Willows (VI-19-1955, J. Vockeroth),
IS, 109 9 (CNC, USNM, and ISC); Val Marie (VI-5-
1955, J. Vockeroth), 69 9 (CNC, USNM, and ISC), (VI-
13-1955, A. Brooks), 19 (CNC).
Wyoming: Yellowstone National Park (VII-16-1923, A.
Melander), 19 (ISC).
REMARKS.?This species is very similar to H.
definita in habitus, except for its larger size and
more robust mid legs. It is also similar in many
respects to H. tibialis.
Hydrellia melanderi, new species
FIGURE 51
DIAGNOSIS.?Palpus dark brown; 6-8 aristal rays;
vertex index 5.0-6.8; all fronto-orbitals anteriorly
inclined; 8-11 dorsal and 9-12 anterior interfrac-
tural costals; thoracic pleuron with light-gray area.
Male length 1.70-1.87 mm; female 1.87-2.30 mm.
Male terminalia as in Figure 51.
HEAD.?Face light gray or yellowish gray; 4-6
medially inclined pfa; 6-9 medially inclined sfa;
epistomal index 1.0-1.5; mesofacial index 2.0-2.2;
80 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
vertex index 5.0-6.8; A-index 1.5-1.8; ocular index
4.5-5.2. Palpus dark brown; 6-8 aristal rays;
antenna dark brown; frons moderate olive-brown;
fronto-orbitals anteriorly inclined; 13-16 postoculars.
THORAX.?Ppn and mesonotum glossy dark grayish
green; 3-A adc and 2 pdc; pleuron light gray except
moderate brown on upper third or so of mesanepis-
ternum; legs dark brown except light-gray coxae.
Wing length 2.04-2.80 mm; veins dark brown; 6-7
setae on basal end of costa; 8-11 dorsal and 9-12
anterior interfractural costals; costal-section ratios:
11:1 2.5-3.0; III:IV 2.3-3.6; V:IV 3.4-3.8; Mi +
 2
index 1.1-1.5.
ABDOMEN.?Most of terga glossy dark grayish
green. Male terminalia: median third of posterior
margin of sternum 5 very slightly concave with small
medial indentation; anterolateral margin of sternum
5 obtusangular (about 110?) and covered by sternum
4; copulobus truncate posteromedially, with promi-
nent anteromedial lobe and anterolateral lobe and
irregularly setose except for seriated setae on
posteromedial margin. Postgonite bent slightly an-
teriad; postgonite uncus almost straight; distiphallus
transversely expanded proximally and near papillate
apex; basolateral bladelike process projecting from
basiphallus dorsal to surstyli. Anterior margin of
fused surstyli broadly and shallowly concave with
medial indentation; B-index about 1.8; C-index
3.0-4.0.
TYPE.?Holotype male, USNM 70538.
TYPE-LOCALITY.?Sardine Creek, Mono County,
California, 8,500 feet (VII-11-1951, A. T. McClay).
PARATYPES.?64 (180*0*, 4 6 ? $ ) from 33 locali-
ties:
Arizona: Oak Creek Canyon, 6,000 feet (VHI-no dates,
F. Snow), 1$ (USNM); Rustler Park, Chiricahua Moun-
tains (VIII-27-1933, collector unknown), 19 , (X-18-
1958, M. Adachi), 1 3 , 1$ (USNM and DLD).
California: Barton Store (IX-3-1950, A. Melander), 1 9
(USNM); Big Bear Lake (V-24-1935, A. Melander), 19
(ANSP); Carson Pass (IX-11-38, M. Crazier), 1 3 (ISC);
Cuyamaca Park (VI-5-1945, A. Melander), 19 (USNM);
Echo (VIII-10-1940, L. Kuitert), 29 9 (USNM); Garden
Valley (V-3-1952, E. Schlinger), 19 (USNM); Green
Valley (VII-26-1944, A. Melander), 39 9 (USNM); Her-
key Camp (V-14-1950, A. Melander), 1 3 , 3 9 9 (USNM);
Jenks Lake (VII-14-1950, A. Melander), 29 9 (USNM),
(IX-7-1950, A. Melander), 13 (USNM); Keen Camp
(VI-7-1942, A. Melander), 1 9 (USNM); Mammoth Lakes
(VII-29-1940, L. Lipovsky), 13 , 19 (USNM), (VII-29-
1940, D. Hardy), 1 3 , 3 9 9 (USNM and ISC); Mono Lake,
Mono County (VI-7-1948, W. Wirth), 13 , 19 (USNM);
Pacific (VIII-9-1940, D. Hardy), 13 (USNM); Palm
Canyon, Borego (V-5-1945, A. Melander), 19 (USNM);
Riverside (II-3-1935, A. Melander), 13 (USNM); Sar-
dine Creek, Mono County, 8,500 feet (VI1-3-1951, collec-
tor unknown), 19 (USNM), (VII-6-1951, collector un-
known), 29 9 (ISC); South Fork of Santa Ana River
(VII-29-1942, A. Melander), 29 9 (USNM and ISC);
Strawberry, Tuolumne County (VI-19-1951, A. McClay),
1 3 (UCD); Tuolumne Meadows (VII-1-1940, D. Hardy),
33 3 (USNM and ISC); Upper Santa Ana River (VI-4-
1950, A. Melander), 1 3 (DLD).
Colorado: Electra Lake (VI-28-1919, collector un-
known), 19 (USNM); Ohio Pass Road, Gunnison County
(VIII-26-1961, D. L. Deonier), 19 (ISC).
Mexico: Mexico City (V-no date-1888, collector un-
known), 2 9 9 (AMNH and ISC); 6 miles west of El Salto,
Durango State (VIII-31-1957, J. Schaffner), 19 (ISC);
Temecula (111-10-1950, M. Wheeler), 19 (ISC).
Nevada: Crystal Springs, Lincoln County (VI-21-1953,
A. Gurney), 23 3 (USNM and ISC); Pyramid Lake (XI-
5-1938, T. Aitkin), 19 (USNM).
New Mexico: Mescalero, Otero County (VI-27-1947, L.
Beamer), 19 (USNM); Penisco River, Mayhill (VI-22-
1953, W. Wirth), 19 (USNM); Ruidoso (VI-26-1940, R.
Beamer), 13 (USNM).
Oregon: Crane Hot Springs, 25 miles southeast of Burns
(IX-11-1962, K. Goeden), 29 9 (KG), (11-23-1963, K.
Goeden), 79 9 (USNM and ISC).
REMARKS.?The few physiognomic differences be-
tween H. melanderi and H. proclinata were not
explored before this revision. It is very probable that
more members of this species group exist in western
localities. For additional remarks, see H. proclinata.
Adult. Habitats recorded: Eleocharis sp., Equise-
tum sp., and margins of hot springs.
Hydrellia morrisoni Cresson
FIGURES 8, 16, 104, 121
Hydrellia morrisoni Cresson, 1924, p. 162; 1931, p. 105;
1944b, pp. 168, 171.?Johnson, 1925, p. 271.?Deonier,
1964, pp. 117, 124, fig. 11; 1965, pp. 500, 506 [ecology].?
Wirth, 1965, p. 744 [catalog listing].
DIAGNOSIS.?Palpus moderate brown; 9-11 aristal
rays; vertex index 4.0-6.5; wing length 2.72-3.03
mm; thoracic pleuron yellowish gray except anterior
half of mesokatepisternum and mesomeron; male
mid and hind tibiae expanded. Male length 2.46-
2.55 mm; female 2.21-2.89 mm. Male terminalia as
in Figure 16.
HEAD.?Face light gray or yellowish gray; 4-6
pfa; epistomal index 1.0-1.4; mesofacial index 1.5-
2.0; vertex index 4.0-6.5; A-index 1.4-1.8; ocular
NUMBER 6 8 81
index 5.5-8.5. Palpus moderate brown; 9-11 aristal
rays; antenna and most of parafrontale moderate
brown; frontal vitta semiglossy dark gray; fronto-
orbital area strong yellowish brown; 13-17
postoculars.
THORAX.?Ppn and mesonotum densely strong
yellowish-brown pruinose; 3-4 adc and 2 pdc;
pleuron yellowish gray except anterior half of
mesokatepisternum and mesomeron; legs, except
tibiae and tarsi, densely light-gray pruinose; tibiae
very sparsely light-gray pruinose over dark brown;
tarsi dark brown; male mid and hind tibiae ex-
panded (Figure 8). Wing length 2.72-3.03 mm;
veins light yellowish brown; 6-10 setae on basal end
of costa; 7-8 dorsal and 8-11 anterior interfractural
costals; costal-section ratios: I I : I 2.0-2.6; III:IV
3.0-3.4; V:IV 3.3-4.0; Mj ?
 2 index 1.2-1.6.
ABDOMEN.?Terga dark gray. Male terminalia:
median third of posterior margin of sternum 5 evenly
and moderately concave; anterolateral margin of
sternum 5 roundly right-angled; copulobus roundly
angular posteriorly (40?-50? from lateral corner)
and irregularly setose except large setae seriated on
posterior margin. Postgonite bent anteromediad and
postgonite uncus slightly laterad; distiphallus taper-
ing to retuse apex. Anterior margin of fused surstyli
broadly emarginate; B-index about 1.2; C-index
0.7-1.0.
THIRD-INSTAR LARVA.?Body of larva not exam-
ined. Frontoclypeal length 0.60 mm; entire feeding
apparatus dark (Figure 104). Ventral frontoclypeal
index 2.8; phragmatal index 0.72; bifurcation index
4.8; clypeal-arch index 1.5. Clypeal arch angled
about 10?-15? in relation to lower frontoclypeal
margin. Mouth-hook beak and base lengths equal;
maximum mouth-hook base thickness about 1.8
times that of beak.
PUPARIUM.?Length 5.35 mm; maximum breadth
1.30 mm; fusiform (Figure 121). Puparial length:
minimum breadth about 24.0; maximum breadth:
minimum breadth 6.2; anal-plate index 2.5. Pro-
thoracic end nearly semicircular in ventral view;
head-lobe scar subcircular; prothorax relatively
smooth ventrally; prothorax and mesothorax sparsely
spinulose; abdominal segment 8 without ventral,
transverse row of setulae, but with 3-4 annuli of
spinules; anal plate subrectangular, with anterior
margin slightly convex; spiracular peritremes sub-
terminal. Empty puparium translucent, light yellow-
ish brown.
TYPE.?Holotype male, USNM 43453.
TYPE-LOCALITY.?White Mountains of New Hamp-
shire (Morrison).
ADULT SPECIMENS EXAMINED.?45 (17c? cf,
28$ $) from 20 localities:
Alaska: Lower Yukon River (VII-5-1951, C. Berg), 1 3 .
Iowa: Pilot Knob State Park, Hancock County (VI-15-
1960, D. L. Deonier), 1 9 ; 3 miles east southeast of Water-
ville, Allamakee County (VIII-2-1960, D. L. Deonier),
33 3,29 9.
Manitoba: Deepdale (VIII-1-1937, R. Beamer), 19.
Massachusetts: Concord (VII-19-1961, W. Wirth),
2 3 3 , 1$.
Michigan: Dickinson County (VII-1-1955, R. Dreis-
bach), 19; Midland County (VI-10-14-1951, R. Dreis-
bach),13.
Minnesota: Houston County (V-27-1940, W. Connell),
43 3 , 59 9 ; Mississippi River, sec. 34 T. 145 north, R.
36 west, Clearwater County (VII-8-1963, D. L. Deonier),
13, 19; Sucker Creek, 100 yards north of Highway 31,
Clearwater County (VII-23-1963, D. L. Deonier), 19.
New Hampshire: Pinkham Notch (VII-9-1931, J. Al-
drich), 13, 69 9 ; Stinson Lake, White Mountains (VII-
23-1961, W. Wirth), 19; White Mountains (no dates,
Morrison), 1 3 .
New Mexico: Jemez Springs (VII-4-1953, W. Wirth),
39 9.
New York: Bergen (VII-24-1946, L. Beamer), 1 $ ;
Ithaca, Inlet Valley (V-7-1957, B. Foote), 13.
North Carolina: Bubbling Spring Creek, 5,100 feet (VII-
17-1957, J.Chillcott), 19.
Ontario: Algonquin Park (VI-26-1955, C. Sabrosky),
19; Ottawa (VI-12-1946, G. Shewell), 19.
Quebec: Great Gaspe Valley (VII-20-1931, J. Aldrich),
13.
Tennessee: Indian Gap, 5,200 feet (VII-23-1957, W.
Richards), 19, (VII-24-1957, W. Richards), 19, (VIII-
3-1957, C.Hines), 13-
Washington: 15 miles west of Kettle Falls (VII-14-1960,
W. Rapp), 13.
IMMATURE SPECIMENS EXAMINED.?7 from 4 locali-
ties:
Minnesota: Headwaters of LaSalle Creek, Itasca State
Park; Sucker Creek, south of Highway 31, Clearwater
County; Sucker Creek, north of Highway 31, Clearwater
County (all collected and reared by D. L. Deonier).
New York: Ithaca, Inlet Valley (collected and reared
by B. Foote).
REMARKS.?The holotype of this species is in very
poor condition.
Adult. Habitats recorded: leaves of Glyceria
grandis and Sagittaria latifolia; reed marsh and
sedge meadow.
82 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Larva. Third-instar larvae were found along with
larvae of other Hydrellia overwintering in Glyceria
grandis shoots in fast, shallow streams of the Lake
Itasca, Minnesota region in April and May 1967.
It is not known if this plant is a summer host, but
this is probable.
Puparium. The time from pupariation to emer-
gence ranged from 8 to 18 days with a mean of 11
for 8 puparia in a temperature range of 55?-75? F.
No puparia were found prior to the middle of May
when low air temperatures were still prevalent.
Hydrellia nobilis (Loew)
FIGURE 36
Psilopa nobilis Loew, 1862, p. 229; 1872, p. 92.?Osten
Sacken, 1878, p. 201.?Becker, 1896, p. 270 [bibliographic
listing].?Aldrich, 1905, p. 625 [catalog listing].?Jones,
1906, p. 180 [catalog listing].
Hydrellia nobilis.?Cresson, 1931, p. 105; 1944b, pp. 164,
173.?Deonier, 1964, p. 116.?Wirth, 1965, p. 744 [catalog
listing].
DIAGNOSIS.?Palpus moderate yellow; 8-13 aristal
rays; vertex index 6.0-7.5; ocular index 17.0-26.0;
wing length 2.33-3.06 mm; apicodorsal antennal
prominent; tibiae moderate yellow. Male length
2.19-2.46 mm; female 2.55-3.13 mm. Male ter-
minalia as in Figure 36.
HEAD.?Face moderate yellow; 5-8 pfa; epistomal
index 1.3-1.8; mesofacial index 2.2-3.0; vertex
index 6.0-7.5; A-index 1.7-2.2; ocular index 17.0-
26.0. Palpus moderate yellow; 8-13 aristal rays;
apicodorsal antennal prominent; antennal segment
3 moderate yellow; antennal segments 1 and 2,
fronto-orbital area, and frontal vitta moderate
brown; most of parafrontale velvety black; 16-19
postoculars.
THORAX.?Ppn and mesonotum semiglossy dark
gray; 4-5 adc and 2 pdc; pleuron light gray except
dark gray upper fifth of mesanepisternum; legs light-
gray pruinose except moderate-yellow tibiae and
tarsi. Wing length 2.33-3.06 mm; veins light brown;
7-8 setae on basal end of costa; 6-11 dorsal and
9-13 anterior interfractural costals; costal-section
ratios: I I : I 2.2-2.6; I I I : IV 2.6-3.2; V:IV 3.6-4.0;
M i .
 2 index 1.3-1.7.
ABDOMEN.?Terga dark brown medially, light
gray laterally and ventrally. Male terminalia: median
third of posterior margin of sternum 5 concave;
anterolateral margin of sternum 5 acutely prominent
(30? angle) ; copulobus slightly convex posteriorly,
notched on medial margin at midlength, and irregu-
larly setose except on medial margin. Postgonite
bent posteriad and then anteriad; postgonite uncus
short, straight; distiphallus undulate laterally and
lamellate ventrally; apical basiphallus breadth 2.0
times anterior breadth of fused surstyli. Anterior
margin of fused surstyli emarginate medially; B-index
about 5.5; C-index 2.5-3.5.
TYPE.?Holotype male, MCZ 11143.
TYPE-LOCALITY.?District of Columbia (collector,
Osten Sacken).
SPECIMENS EXAMINED.?96 (37cf d\ 5 9 ? $ ) from
24 localities:
Illinois: Chicago (no dates?1899, collector unknown),
1 3 ; McHenry (VIII -21 -no year, A. Melander), 1 3 .
Indiana: Chesterton ( V I - 2 - 1 9 1 6 , collector unknown),
IS.
Iowa: Siewer's Springs State Park, near Decorah ( I X -
15-1962, D. L. Deonier), 1 3 , 2 9 9 -
Minnesota: Biological Station, Itasca State Park ( V I -
26-1963, D. L. Deonier), 1 $ ; Bohall Lake, Itasca State
Park (VIII -17-1963 , D. L. Deonier), 1 $ ; Bohall Trail
Bog (VIII -17-1963 , D. L. Deonier), 1 1 4 3 , 1 9 9 9 ;
Crookston ( IX-5-1936 , D. Denning), 1 9 ; Douglas Lodge
Bay, Lake Itasca, Itasca State Park (VI -20 -1963 , D . L.
Deonier), 5 3 3 , 4 9 9 ; Iron Corner Lake ( V I - 1 8 - 1 9 6 3 ,
D . L. Deonier), 1 3 , 1 9 ; Itasca State Park (VII -14 -1938 ,
P. Schroeder), 1 9 ; Kabekona Creek, Hubbard County
(VIII -8 -1963 , D. L. Deonier), 2 9 9 ; Mississippi River at
Sucker Creek, Clearwater County ( V I I - 9 - 1 9 6 3 , D. L. Deo-
nier), 1 9 ; Mississippi River near north entrance Itasca
State Park (VII -9 -1963 , D. L. Deonier), 1 3 , 1 9 ; Prof
Green Trail, Itasca State Park ( V I I - 2 5 - 1 9 6 3 , D. L. Deo-
nier), 13 ; Squaw Lake, Itasca State Park ( V I I I - 1 4 - 1 9 6 3 ,
D. L. Deonier), 2 3 3 ; Two Island Lake, Itasca State
Park (VI-28-1963 , D. L. Deonier), 1 3 ; west side across
from Biological Station, Lake Itasca, Itasca State Park
(VII -28-1963 , D. L. Deonier), 1 3 , (VII I -14 -1963 , D. L.
Deonier), 8 3 3 , 2 0 9 9 .
New Jersey: Trenton (VII I -5 -1910 , collector unknown)
1 9 .
New York: Dryden Lake, Tompkins County ( V I I I - 8 -
1961, D. L. Deonier), 2 9 9 ; Ithaca (VI-8 -1937 , collec-
tor unknown), 1 9-
Pennsylvania: 2 miles north of Narberth, Montgomery
County ( IX-9-1915 , collector unknown), 1 3 .
Quebec: l ie Jesus, 4 miles north of Montreal ( V I I I - 1 9 -
1956, J. Laffoon), 1 3 .
Virginia: Alexandria ( V I I - 6 - 1 9 5 2 , W. Wirth), 1 9 .
REMARKS.?Hydrellia nobilis, H. atroglauca, and
H. idolator appear to constitute a species group.
Adult. Habitats recorded: leaves of Potomogeton
NUMBER 6 8 83
gramineus, P. natans, Nuphar advena, Leersia
oryzoides, Zizania aquatica, and Nasturtium officinale.
I found several dead specimens on leaves of P.
natans at Squaw Lake, Minnesota. I made the fol-
lowing observations on behavior of adults. At Lake
Itasca, Minnesota, a male tried to mount a male
H. ischiaca after repeatedly scissoring its wings, and
another male tried to mount a female H. ischiaca
but the latter repulsed it by rapidly striking out with
its fore tarsi. Specimens at Lake Bohall, Itasca State
Park, behaved similarly. At Bohall Trail Bog, a pair
captured while in copulation remained so for 6 min-
utes in the live-capture tube. They copulated again
in the laboratory two days later. At this locality,
another pair approached each other and without any
apparent epigamic behavior, the male mounted
from the side. Shortly afterwards, a second male
approached the pair frontally, crawled onto the pair
and exerted its phallus for a few seconds before
being forced off. It did this eleven times, and several
times when it approached from the rear the copu-
lating female turned quickly to meet it. Despite this
interference, the pair copulated for 19 minutes. Here
also, a male extended its wings straight out from the
thorax and approached a second male, but it stopped
abruptly and folded its wings. Then the second male
extended its wings when faced by another specimen
of H. nobilis. The latter immediately flew away and
then the first male tried to mount the second. In
another case, a male approached and tried to mount
another male. The latter did not meet this advance
with wing extension as it did in an immediately
preceding instance.
Larva. Third-instar larvae were found along with
larvae of other Hydrellia overwintering in Glyceria
striata in slow, shallow streams and lakes of the
Lake Itasca, Minnesota, region in May 1967. It is
not known if this plant is a summer host. Specimens
of this grass were found embedded in ice.
Puparium. The time from pupariation to emer-
gence ranged from 10 to 14 days, with a mean of
11.8 days for 12 puparia in a temperature range of
55?-75? F.
Hydrellia notata, new species
FIGURE 52
DIAGNOSIS.?Palpus moderate yellow; 6-9 aristal
rays; vertex index 7.2-8.7; ocellar absent; 2 anterior
fronto-orbitals; supra-alar area velvety black; tho-
racic pleuron nearly all light gray. Male length
1.33-1.67 mm; female 1.70-2.00 mm. Male ter-
minalia as in Figure 52.
HEAD.?Face light gray or yellowish gray; 4-6
pfa; epistomal index 1.2-1.6; mesofacial index 2.2-
2.5; vertex index 7.2-8.7; A-index 1.4-2.0; ocular
index 8.3-9.0. Palpus moderate yellow; 6-9 aristal
rays; antenna dark brown; parafrontale velvety dark
brown; frontal vitta glossy dark brown; subocular
area to comer of postocular area dark brown; upper
fourth of postocular area dark gray, remainder
light gray; 2 anterior fronto-orbitals and 1 posterior
fronto-orbital; ocellar absent; length of postocellar
about 5.0 times that of posterior fronto-orbital;
10-15 postoculars.
THORAX.?Ppn and mesonotum dark brown except
velvety black supra-alar area, light-brown postalar
area with small light-gray spot, and densely light-
brown pruinose scutellum; 3-4 adc and 2 pdc;
pleuron light gray except brown spot above fore
coxa; legs light-gray pruinose except moderate yellow
tibial apices. Wing length 1.62-1.99 mm; veins
light yellowish brown; 6-7 setae on basal end of
costa; 6-8 dorsal and 8-10 anterior interfractural
costals; costal-section ratios: II:I 2.4-3.8; III:IV
2.8-3.6; V.IV 3.5-3.8; Mi
 + 2 index 1.5-1.8.
ABDOMEN.?Terga dark brown. Male terminalia:
median third of posterior margin of sternum 5 con-
vex; anterolateral margin of sternum 5 obtusangular
(about 100?); copulobus conical, directed postero-
laterad, and irregularly setose, with 4-5 medially
inclined macrochaetae on posteromedial margin.
Postgonite short, bent anteriad; postgonite uncus
fusiform and straight; distiphallus short, almost uni-
form in breadth to slightly convex apex. Anterior
margin of fused surstyli deeply V-clefted medially
to a central gibba; B-index about 2.6; C-index 5.0-
6.5.
TYPE.?Holotype male, USNM 70539.
TYPE-LOCALITY.?Canaan, Connecticut (VIII-31-
1952, A. Stone).
PARATYPES.?5(2cT d\ 3? ? ) from 5 localities:
Florida: Hilliard, Nassau County (VIII-6-1939, R.
Bessemer), 1 <J (ISC); Liberty County (111-22-1954, G.
Steyskal), 19 (USNM).
Georgia: Blackshear, Pierce County (V-10-1911, collec-
tor unknown), 1 9 (CU).
Mississippi: Vancleave Road, Jackson County (VII-7-
1962, D. L. Deonier), 1 <$ (ISC).
84 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
New York: Fish Creek Pond (VIII-13-1941, A. Me-
lander), 1$ (USNM).
REMARKS.?On the basis of several characters,
this very distinct species belongs to the H. formosa
species group.
Adult. Habitats recorded: leaves of Nymphaea
odorata and seepage slope.
Hydrellia notiphiloides Cresson
FIGURES 65, 84,95, 116
Hydrellia notiphiloides Cresson, 1924, p. 162; 1944b, pp.
170, 175.?Wirth, 1965, p. 744 [catalog listing].
DIAGNOSIS.?Palpus moderate yellow; prementum
glossy brownish black; 6-8 aristal rays; ocular index
3.7-4.3; body length:wing length 0.9-1.0; lower
three-fourths of postocular area light gray, upper
fourth brown; thoracic pleuron light gray; costal
section I I I : IV 2.5-3.5. Male length 1.89-2.30 mm;
female 2.13-2.55. Male terminalia as in Figure 65.
HEAD.?Face light gray or pale yellow; 4-6 pfa;
epistomal index 1.0-1.3; mesofacial index 1.3-1.9;
vertex index 3.6-5.5; A-index 1.6-2.8; ocular index
3.7^4.3. Palpus moderate yellow; 6-8 aristal rays;
prementum glossy brownish black; antenna and most
of parafrontale dark brown; fronto-orbital area and
frontal vitta moderate brown; 13-17 postoculars.
THORAX.?Ppn and pleuron light gray; mesonotum
moderate brown; 3-4 adc and 2-3 pdc; legs light-
gray pruinose except dark-brown tarsi. Wing length
2.04-2.72 mm; veins moderate brown; 7-11 setae
on basal end of costa; 6-10 dorsal and 7-11 anterior
interfractural costals; costal-section ratios: I I : I 1.8-
2.2; I I I : IV 2.5-3.5; V:IV 3.0-3.6; Mi ?
 2 index
1.1-1.6.
ABDOMEN.?Terga dark gray or moderate brown.
Male terminalia: median third of posterior margin
of sternum 5 evenly and moderately concave; antero-
lateral margin of sternum 5 roundly right-angled;
copulobus acuminate posteriorly and irregularly
setose. Postgonite bent anteriad and postgonite uncus
posterolaterad and truncate apically; distiphallus
tapering distally. Anterior margin of fused surstyli
shallowly notched medially; B-index about 1.0;
C-index 2.0-2.5.
THIRD-INSTAR LARVA.?Body of larva not exam-
ined. Frontoclypeal length 0.48-0.60 mm; cheliform
spot oblique to dorsal phragmatal ramus (Figure 95).
Ventral frontoclypeal index 3.0-3.5; phragmatal
index 0.9-1.0; bifurcation index 4.5-5.5; clypeal-
arch index 1.6-1.7. Clypeal arch sloping smoothly
along about 20? angle relative to lower frontoclypeal
margin. Mouth-hook beak longer than base; maxi-
mum mouth-hook base thickness about 2.0 times that
of beak (Figure 84).
PUPARIUM.?Length 3.25-4.00 mm; maximum
breadth 1.00-1.25 mm; fusiform (Figure 116).
Puparial length: minimum breadth 15.0-17.0; maxi-
mum breadth: minimum breadth 5.0-6.0; anal-plate
index 1.8-2.4. Prothoracic end semicircular in ven-
tral view; head-lobe scar subcircular; maximum
puparial breadth 7.5-8.0 times maximum breadth
of head-lobe scar and 2.5 times maximum pro-
thoracic breadth; prothorax with many ventral
rugulae; mesothorax and metathorax prominently
rounded laterally; lateral spinules inconspicuous in
midlength of puparium; abdominal segment 8 with-
out ventral, transverse row of setulae and with only
2-3 distinct annuli of spinules (immediately peri-
spiracular); anal plate ovoid, with anterior margin
slightly or distinctly convex. Empty puparium trans-
lucent, light yellowish brown.
TYPE.?Holotype male, OSU (not numbered).
TYPE-LOCALITY.?Cedar Point, Sandusky, Ohio
(VIII-5-1902, collector unknown).
ADULT SPECIMENS EXAMINED.?133 (56cTcf,
77$ $ ) from 29 localities:
Arizona: Granite Delta (VII-3-1950, H. Wright), 1 $ .
British Columbia: Hatzic Lake (VII-26-1953, W. Ma-
son), 1 $.
California: Big Bear Valley (VIII-9-1933, collector un-
known), A$S, 65 9 ; Buena Park (V-19-1914, A. Me-
lander), 2SS, 1 $ ; Carpenteria (VIII-11-1950, A. Me-
lander), 6S S, 15 9 9 ; Davis (VII-21-1955, A. Grigarick),
IS, 2 9 9 , (VII-29-1955, A. Grigarick), 5SS, 8 9 9 ,
(IX-16-1954, A. Grigarick), 19 , (IX-17-1955, A. Gri-
garick), 7$S, 29 9, (IX-21-1957, A. Grigarick), IS,
(IX-22-1955, A. Grigarick), 19 , (IX-27-1955, A. Gri-
garick), 2$S, 49 9 ; Green Valley (VII-26-1944, A.
Melander), 1 $ ; Jenks Lake, San Bernardino (VI I I -3 -
1942, A. Melander), 29 9 , (VII-22-1957, J. Hall), 1 9 ;
Maxwell, Colusa County (VIII-12-1954, A. Grigarick),
7 S S, 69 9 ; Putah Canyon, Yolo County (X-20-1954, A.
Grigarick), 2 S S ; Vidal, San Bernardino County (IV-no
date-1948, R. Coleman), I $ ; Santa Cruz, Santa Cruz
County (VII-17-1940, B. Brookman), 29 9 .
Iowa: Little Wall Lake, Hamilton County (IX-22-1962,
D. L. Deonier), 2$ S, 19 ; Spring Lake, Greene County
(IX-19-1961, D. L. Deonier), 1 9 ; Springbrook State
Park, Guthrie County (VI-23-1961, D. L. Deonier), 1 3 .
NUMBER 6 8 85
Massachusetts: Nantucket (VII?20?year unknown, col-
lector unknown), 1 3 .
Mexico: Tepexpan, Mexico, 6,900 feet (VIII-12-1954, J.
Chillcott), 1 $ .
Michigan: Cheboygan County (VII-21-1947, C. Berg),
1 3 , 59 9 ; Ocqueoc Lake, Presque Isle County (VII-13-
1941, C. Berg), 1 3 .
Minnesota: Bigstone County (VI-5-1938, C. Mickel),
19 ; Chamber's Creek, Itasca State Park (VII-18-1963,
D. L. Deonier), 1 3 .
Nevada: Ely (VIII-13-1940, R. Beamer), 1 ?$.
Ohio: Cedar Point, Sandusky River (VIII-5-1902, col-
lector unknown), 1 3 , 2 9 9.
Ontario: London (VII-31-1958, W. Judd), 19 .
Quebec: Rupert House (VII-25-1949, D. Gray), 3 3 3 ,
49 9 , (VII-26-1949, D. Gray), 1 9 , (VII-29-1949, E.
LeRoux), 1 3 , 19 , (VII-30-1949, E. LeRoux), 1 3 , 49 9 ,
(VIII-7-1949, E. LeRoux), 1 9 .
Saskatchewan: Saskatoon (VHI-no date-1947, collector
unknown), 1 9 ?
Virginia: Colonial Beach (VIII?no date?1916, collector
unknown), 1 9 .
Wyoming: Biscuit Basin, Yellowstone National Park
(VIII-2-1954), A. Melander), 1 9 ; Northwest entrance Yel-
lowstone National Park (VI11-3-1938, A. Melander), 13-
IMMATURE SPECIMENS EXAMINED.?8 from 2
localities:
California: Davis (collected and reared by A. Grigarick).
Michigan: Cheboygan County (collected and reared by
C. Berg).
REMARKS.?Adults of H. notiphiloid.es and H.
caliginosa are so similar that inspection of the male
terminalia often may be necessary. I have not seen
the holotype of this species.
Adult. Habitats recorded: leaves of Oryza sativa
and Potamogeton nodosus; limnic wrack.
Host plants. Grigarick reared this species from
Zannichellia palustris and Polypogon monspeliensis
at Davis, California, September 1955 and 1957, dur-
ing research on H. griseola. Judd (1964) recorded
one adult from an emergence trap floating near
Potamogeton amplifolius in Ontario in July. The
water depth where the trap was anchored ranged
from 5.3 to 5.4 meters.
Hydrellia penicilli Cresson
FIGURE 66
Hydrellia penicilli Cresson, 1944b, pp. 168-169.?Deonier,
1964, p. 116; 1965, pp. 500, 506 [ecology].?Wirth, 1965,
p. 744 [catalog listing].
DIAGNOSIS.?Palpus moderate yellow; 8-10 aristal
rays; vertex index 3.1-4.0; antennal segment 3 partly
moderate yellow; mesonotum and abdomen strong
yellowish brown; thoracic pleuron light gray; 3 black
anterior basal setae on hind basitarsus. Male length
2.60-2.81 mm; female 2.62-3.15 mm. Male ter-
minalia as in Figure 66.
HEAD.?Face yellowish gray; 4-6 pfa; 1-2 sfa;
epistomal index 1.0-1.2; mesofacial index 1.5?1.8;
vertex index 3.1-4.0; A-index 1.6-1.8; ocular index
6.0-8.0. Palpus moderate yellow; 8-10 aristal rays;
antennal segments 1 and 2 and most of parafrontale
moderate brown; antennal segment 3 partly mod-
erate yellow; frontal vitta and fronto-orbital area
yellowish gray or strong yellowish brown; 14-17
postoculars.
THORAX.?Ppn and mesonotum strong yellowish-
brown; 3^4 adc and 2 pdc; pleuron light gray or
strong yellowish brown; 1-2 basal coxals (1 macro-
chaetous); legs light-gray pruinose except dark-
brown tarsi; 3 black anterior basal setae on hind
basitarsus. Wing length 3.06-3.40 mm; veins light
brown; 8-11 setae on basal end of costa; 7-10
dorsal and 9-12 anterior interfractural costals;
costal-section ratios: I I : I 2.5-2.9; I I I : IV 2.5-2.8;
V: IV 3.1-3.9 ; M , t 2 index 1.0-1.4.
ABDOMEN.?Terga and sterna strong yellowish
brown. Male terminalia: median third of posterior
margin of sternum 5 concave; anterolateral margin
of sternum 5 smoothly rounded; copulobus regularly
and densely setulose and truncate posteriorly. Post-
gonite bent anteriad and postgonite uncus slightly
laterad; distal two-thirds of distiphallus uniform in
breadth. Anterior margin of fused surstyli deeply
and narrowly cleft medially to three-fourths of length
of structure; B-index about 1.0; C-index 1.5-2.0.
TYPE.?Holotype male, ANSP 6656.
TYPE-LOCALITY.?Pembroke, Ontario, Canada
(VII-3-1938, A. L. Melander).
SPECIMENS EXAMINED.?ll^cfo" , 9 $ $ ) from
10 localities:
British Columbia: Qualicum (VI-21-1955, G. Shewell),
13-
Iowa: Goose Lake, Hamilton County (VII-14-1960,
D. L. Deonier), 19 .
Maine: Salisbury Cove (VII-15-1923, C. Johnson), 1 9.
Manitoba: Birtle (VIII-13-1928, R. Bird), 2 9 9-
Michigan: Wexford County (VII-4-1952, R. Dreisbach),
1 3 , 19 .
New York: McLean (V-24?1915, collector unknown),
1 9 .
86 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Ontario: Pembroke (VII-3-1938, A. Melander), 29 9 ;
Waubumick (VH-no date-1915, H. Parish), 1 $ .
Quebec: Natashquan (VIII-5-1929, W. Brown), 1 $ ;
Thunder River (VIII-19-1930, W. Brown), 1 5 .
REMARKS.?This species seems closely related to
H. cruralis, H. notiph.iloid.es, and H. cessator.
Adult. Habitat recorded: reed marsh.
Hydrellia personata, new species
FIGURE 49
DIAGNOSIS.?Palpus dark brown; 6-8 aristal rays;
dorsal interfractural costals twice size of anterior
ones; body length:wing length 0.7-0.8; apicodorsal
antennal prominent; frons (except light-brown
ocellar triangle) velvety dark brown; 1 pdc; thoracic
pleuron moderate reddish brown. Male length 1.28?
1.45 mm; female 1.45-1.62 mm. Male terminalia as
in Figure 49.
HEAD.?Face light gray; 3-4 pfa; 1-3 sfa; epis-
tomal index 1.0?1.4; mesofacial index 1.5-2.0;
vertex index 5.5-7.0; A-index 1.3-2.0; ocular index
4.0-5.0. Palpus dark brown; 6-8 aristal rays;
apicodorsal antennal prominent; antenna and frons
(except light-brown ocellar triangle) velvety dark
brown; 12-16 postoculars.
THORAX.?Ppn and mesonotum moderate olive-
brown; 3-4 adc (1-2 macrochaetous) and 1 pdc;
auxiliary apical scutellars usually present; pleuron
moderate reddish brown; legs dark brown except
yellowish-gray trochanters. Wing length 1.53-2.04
mm; veins dark brown; 6-8 setae on basal end of
costa; 3-6 dorsal and 6?9 anterior interfractural
costals (dorsals 2.0 times size of anteriors) ; costal-
section ratios: 11:1 1.8-2.2; III:IV 3.5-4.0; V:IV
3.0-3.6; M i .
 2 index 1.2-1.5.
ABDOMEN.?Terga dark brown. Male terminalia:
median third of posterior margin of sternum 5
broadly concave; anterolateral margin of sternum 5
rounded through 95?-100? angle; copulobus almost
truncate posteriorly and irregularly setose. Postgonite
straight and covered by surstyli; postgonite uncus
straight and aciculate; maximum breadth of pre-
gonite 2.0 times that of postgonite, pregonite extend-
ing further anteriad; distiphallus slightly expanded
at midlength. Anterior margin of fused surstyli
acutely papilliform medially; B-index about 4.5;
C-index 0.4-0.7.
TYPE.?Holotype male, WSC 322.
TYPE-LOCALITY.?O' Sullivan Dam, Grant County,
Washington (X-30-1954; presumed collector, H.
G.Davis).
PARATYPES.?27 (10c* d \ 17 9 ? ) from 9 locali-
ties:
Arizona: Patagonia, Santa Cruz County (VI-27-1953,
W. Wirth), 1$ (USNM).
California: Mono Lake, Mono County (VI-7-1948, W.
Wirth), 19 (USNM); Richmond, Contra Costa County
(VI-16-1948, W. Wirth), 59 9 (USNM); Sunol, Alameda
County (IX-9-1947, W. Wirth), 19 (USNM); Victor-
ville, San Bernadino County (V-2-1953, G. Moran and
R. Schuster), 19 (USNM).
Iowa: Izaak Walton League reserve, near Ames, Story
County (IV-27-1962, D. L. Deonier), 1$, 29 9 (USNM
and ISC); Ames (IV-23-1962, D. L. Deonier), 1 $ (DLD).
Texas: University of Texas Arboretum, Austin (1-20-
1950, presumed collector, M. Wheeler), 2 $ $, 3 9 9 (MRW,
USNM, ISC, and DLD).
Washington: O'Sullivan Dam, Grant County (X-30-
1954, H. Davis), 6$ 3, 3 9 9 (USNM, ISC, and WSC).
REMARKS.?I have not been able to place this
species in any particular group. It differs from the
H. prudens and H. tibialis groups noticeably by its
distinctly wide head, and its normal, unexpanded,
male mid tibia.
Adult. Habitats recorded: sedge meadow and
margin of Mono Lake, California.
Hydrellia platygastra Cresson
FIGURE 38
Hydrellia platygastra Cresson, 1931, pp. 105, 106; 1944b,
pp. 168, 172.?Wirth, 1965, p. 744 [catalog listing].
DIAGNOSIS.?Palpus dark brown; 8-11 aristal rays;
mid and hind tarsi moderate orange-yellow; face
velvety dark brown; thoracic pleuron light brown
except light gray on lower half of mesokatepistemum;
10-14 setae on basal end of costa; male abdomen
compressed. Male length 2.30-2.38 mm; female
2.89-3.06 mm. Male terminalia as in Figure 38.
HEAD.?Face velvety dark brown; 3-4 pfa; 6-9
sfa; epistomal index 1.0-1.2; mesofacial index 1.7-
2.2; vertex index 4.0-7.0; A-index 1.7-2.0; ocular
index 4.5-6.0. Palpus dark brown; 8-11 aristal rays;
apicodorsal antennal prominent; antenna, frontal
vitta, and fronto-orbital area dark-brown pruinose;
most of parafrontale velvety black; 17-18 post-
oculars.
NUMBER 6 8 87
THORAX.?Ppn and mesonoturn brassy moderate
brown; 4-5 adc and 2-3 pdc; pleuron semiglossy
light brown except light gray on lower half of
mesokatepisternum; legs dark brown except dark-
yellow trochanters and moderate orange-yellow mid
and hind tarsi. Wing length 2.89-3.40 mm; veins
dark brown; 10-14 setae on basal end of costa; 6-9
dorsal and 9-11 anterior interfractural costals;
costal-section ratios: I I : I 2.2-2.5; I I I : I V 2.5-3.0;
V:IV 3.0-3.8; Mi ?
 2 index 1.4-1.6.
ABDOMEN.?Terga and sterna dark gray. Male
terminalia: median third of posterior margin of
sternum 5 deeply concave; anterolateral margin of
sternum 5 smoothly rounded on a very obtuse angle;
copulobus curving elliptically posteriorly and regularly
setose. Postgonite bent anteriad and postgonite uncus
straight; distal half of distiphallus shallowly undulate
laterally and lamellate ventrally. Anterior margin of
fused surstyli broadly emarginate; B-index about 2.0;
C-index 0.5-0.8.
TYPE.?Holotype male, ANSP 6483.
TYPE-LOCALITY.?Beaver Creek, Newport, Oregon
(no dates, J. M. Aldrich).
SPECIMENS EXAMINED.?71 (35o*o*, 36 9 9 ) from
24 localities:
British Columbia: Langley (VIII-9-1917, A. Melander),
1,5; Milner (VII-12-1953, W. Mason), 13, (VIII-12-
1953, W. Mason), It, 2 9 9.
California: Willits (V-30-1955, E. Schlinger), 1 9 .
Idaho: Lake Coeur d'Alene (VII-15-1926, E. Nast),
1 9 ; Potlatch (VI-17-1911, collector unknown), 1 $ ; Sol-
dier Creek, Priest Lake (VIII-22-1919, A. Melander), 19 .
Nebraska: Arapahoe (VIII-11-1960, W. Rapp), It.
(This specimen may have been incorrectly labeled, for it is
the sole record east of the Rocky Mountains.)
Oregon: Albany (V-26-1941, Schuh and Gray), 1 9 ;
Beaver Creek, Newport (no dates, J. Aldrich), 39 9 ; Brei-
tenbush Springs, 2,222 feet (VII-6-1934, H. Scullen), 19 ;
Gold Beach (VII-27-1951, collector unknown), 29 9 ;
Hood River (no dates, Childs), 1 3 , 5 9 9 ; Independence
(VII-25-1934, N. Larson), 1 9 ; Marshfield (VI-27-no
year, J. Aldrich), 2$ 3 , 1 9 ; Newport (VI-9-1925, E. Van
Dyke), 13 ; Sear Lake, Fort Lewis, Pierce County ( I X - 1 -
1945, P. Arnaud),2 9 9 .
Washington: Arlington (VII-28-1931, J. Nottingham),
1 9 ; Cusick (VII-14-1960, W. Rapp), 1 9 ; Five-Mile Lake
(VII-4-1935, A. Melander), 1 1 ; Hwaco (Vll-no date-
1917, A. Melander), 1$, 19 , (VIII-27-1917, A. Me-
lander), IS, (VI-28-1925, A. Melander), 2tt; Lake
Stevens, Everett (VIII-5-1917, A. Melander), 1 9 ; Sea-
view (VI-26-1925, A. Spuler), 1 $ ; Tacoma (VIII-27-
1911, collector unknown), 1 3 ; 15 miles west of Kettle
Falls (VII-14-1960, W. Rapp), 193 3 , 119 9.
REMARKS.?Hydrellia platygastra and H. wilburi
have identical male terminalia (as far as I could
determine). The principal differential character is
facial color. Even though these two species are
sympatric to a great extent, I have jusified species
designation for H. wilburi on the absence of inter-
gradational phenotypes in several moderately large
series and on the distinct possibility of ecological
isolation of the two groups. The localities for H.
wilburi are all at higher altitudes than those for
H. platygastra.
Hydrellia proclinata Cresson
FIGURE 57
Hydrellia proclinata Cresson, 1915, pp. 69-70; 1931, p. 107;
1944b, pp. 165-171.?Wirth and Stone, 1956, p. 469.?
Wirth, 1965, p. 744 [catalog listing].
DIAGNOSIS.?Palpus dark brown; 6-8 aristal rays;
all fronto-orbitals anteriorly inclined; vertex index
4.5-7.0; body length:wing length 0.8-1.0; thoracic
pleuron moderate brown with 2 light gray areas.
Male length 1.87-2.21 mm; female 2.30-2.81 mm.
Male terminalia as in Figure 57.
HEAD.?Face light gray; 4-6 medially inclined
pfa; 6-8 medially inclined sfa; epistomal index 1.0-
1.3; mesofacial index 2.0-2.2; vertex index 4.5-7.0;
A-index 2.0-2.2; ocular index 7.0-7.5. Palpus dark
brown; 6-8 aristal rays; antenna, frontal vitta, and
fronto-orbital area moderate brown; most of para-
frontale velvety black; apicodorsal antennal promi-
nent; anterior and posterior fronto-orbitals anteriorly
inclined; 14-17 postoculars.
THORAX.?Ppn light brown, with light-gray tinge;
mesonotum glossy dark grayish green; 3-4 adc and
2-3 pdc; occasional auxiliary apical scutellars;
pleuron moderate brown except middle third of
propleuron and mesanepisternum and lower anterior
corner of mesepimeron light gray (these forming 2
distinct spots) ; legs dark brown except fore coxa
partly light gray. Wing length 2.13-2.89 mm; veins
dark brown; 6-7 setae on basal end of costa; 8-11
dorsal and 9-12 anterior interfractural costals;
costal-section ratios: I I : I 2.2-3.5; I I I : IV 2.6-2.8;
V: IV 3.0-3.8; M, *
 2 index 1.0-1.5.
ABDOMEN.?Most of the terga usually glossy dark
grayish green. Male terminalia: median third of
posterior margin of sternum 5 concave; anterolateral
88 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
margin of sternum 5 acutangular (80?-90?); copu-
lobus distinctly bilobate posteromedially, deeply
emarginate at midlength on medial margin, and
irregularly setose. Postgonite bent anterolaterad and
postgonite uncus laterad; distal half of distiphallus
of uniform breadth (proximal half 3.0-4.0 times as
broad as distal half). Anterior margin of fused
surstyli nearly truncate with only slight medial
indentation; B-index about 2.0; C-index 0.8-1.0.
TYPE.?Holotype male, ANSP 6075.
TYPE-LOCALITY.?Berkeley Hills, Alameda County,
California (IV-20-1908, E. T. Cresson, Jr.).
SPECIMENS EXAMINED.?797 (269cTcf, 5 2 8 $ $ )
from 169 localities:
Alberta: Vermilion Lake, Banff, 4,500 feet (VIII-17-
1925, O.Bryant), 2 9 9 .
Arizona: Chiricahua Mountains (VII?5?1940, D. Hardy),
15 , (VII-3-1947, L. Beamer), 29 9 ; Coconino County
(VIII-1&-1927, R. Beamer), 1 3 ; Hassayampa River,
Wickenburg (VI-29-1953, W. Wirth), 1 9 ; Oak Creek
Canyon, Sedona (VI-29-1953, W. Wirth), 29 9 ; Pinery
Canyon, Chiricahua Mountains, Cochise County, 6,000 feet
(VI-23-1919, W. Stone), 1 3 ; Sunnyside Canyon, Hua-
chuca Mountains (VI1-9-1940, L. Lipovsky), 19 .
British Columbia: Atlin, 2,200 feet (VI-13-1955, H.
Huckel), 1 9 , (VIII-11-1955, B. Gibbard), 1 9 ; Okana-
gan Falls, 2,000 feet (VI-16-1953, J. McGillis), 4 3 3 ,
29 9 ; Oliver, 1,000 feet (VIII-18-1953, J. McGillis), 19 .
California: Adobe Creek, West Stanislaus County (V-6-
1948, J. Mac Swain), 1 <$ ; Asilomar (IX-29-1946, A. Me-
lander), 1 3 , (X-2-1946, A. Melander), 1 3 , 1 9 ; Auburn
(V-12-1953, J. Hall), 1 <$ , 1 9 ; Barton Flats (VIII-3-1942,
A. Melander), 59 9 ; Barton Store (VII-4-1946, A. Me-
lander), 1 3 ; Berkeley Hills, Alameda County ( IV-11-
1908, collector unknown), 19 , (IV-20-1908, collector un-
known), 33 <$, 59 9 ; Big Bear Lake (V-24-1935), A.
Melander), 1 9 ; Big Meadow, 7,200 feet (VII-30-1937,
G. Spurlock), 13 ; Big Pines (VIII-2-1944, A. Melander),
23 3 , 7 9 9 , (VIII-9-1944, A. Melander), 53 3 , 7 9 9,
(V-23-1945, A. Melander), 29 9 ; Big Springs, Shasta
County, 4,000 feet (VI-26-1947, H. Chandler), 1 9 ; Bishop
(VII-28-1940, D. Hardy), 1 3 , 1 9 , (VII-28-1940, L.
Kuitert), 1 3 , (VII-28-1940, L. Lipovsky), IS, 1 9 ;
Bowerman Meadow Trinity County (VI-3-1951, A. Mc-
Clay), 1 9 ; Cajon (VIII-9-1944, A. Melander), 1 9 ;
Campo (VII-18-1940, R. Beamer), 1 9 ; Carson Pass (IX-
11-1938, M. Cazier), 1 $ ; Cassel, Shasta County (VII-15-
1955, E. Schlinger), 19 ; Cedar Pass, Modoc County (V-
15-1948, W. Wirth), 1 9 ; Cisco (VI-5-1940, M. James)
1 S ; Clarksville, Eldorado County (IV-27-1929, H. Keefer),
1 9 ; Clear Lake Oaks, Lake County (IV-19-1954, E.
Schlinger), 1 9 ; Clio, Plumas County (VII-9-1916, H.
Dyar), 1 S ; Coalinga, Fresno County below 500 feet (VI-
1-3-1907, J. Bradley), 19 ; Convict Creek, Mono County
(VI-6-1948, W. Wirth), IS; Cow Creek, Tuolumne
County (VI-25-1951, A. McClay), 29 9 ; Crabtree
Meadow, Tulare County, 10,500 feet (VII-29-1915, col-
lector unknown), IS; Dellecker, Plumas County (VII-4-
1951, E. Schlinger), 1 9 ; Ebbett's Pass, Alpine County
(IX-11-1938, M. Cazier), 23 3 , (VII-22-1957, E. Mez-
ger), 19 , (VIII-24-1957, E. Mezger), 1 9 , (VIII-29-1957,
E. Mezger), 19 ; Echo (VIII-10-1940, D. Hardy), 4$ S,
(VIII-10-1940, L. Kuitert), \S, 49 9 , (VIII-10-1940,
L. Lipovsky), 29 9 ; Echo Lake, Eldorado County (VII -
23-1955, E. Schlinger), 4 3 $, 59 9 ; Felton, Santa Cruz
Mountains (V-20-25-1907, J. Bradley), IS, 1 5 ; Garden
Valley, Eldorado County (V-3-1952, E. Schlinger), 1$,
2 9 9 ; Glacial Point Road, Yosemite National Park (VII -
6-1947, A. Melander), 1 9 ; Grass Lake, Luther Pass, El-
dorado County (VII-24-1955, E. Schlinger), 59 9 ; Green
Valley (VII-26-1944, A. Melander), 1$, 59 9 ; Guatay,
San Diego (IV-10-1950, L. Quate), 1 9 ; Hallelujah Junc-
tion, Lassen County (VII-4-1949, J. MacSwain), 1 9 ,
(VII-16-1953, E. Schlinger), 1 <J ; Hat Lake, Lassen Na-
tional Park (VI1-2 3-1950, P. Arnaud), 19 ; Herkey Camp
(V-14-1950, A. Melander), IS, 1 9 ; near Hobart Mills,
Nevada County (X-10-1952, E. Schlinger), 1 9 ; Hope
Valley, Alpine County (VII-18-1948, L. Quate), 1 9 ;
Idylwild (VI-4-1935, P. Oman), 1 9 ; Jacumba (VII-17-
1940, D. Hardy), 29 9 ; Jenks Lake (VIII-18-1950, A.
Melander), 1 9 ; Keen Camp (VI-7-1942, A. Melander),
2S S, 19 ; Laguna Mountains (VII-6-1929, L. Anderson),
1 9 , (VII-6-1929, R. Beamer), 1 9 ; Lake Alpine ( IX-
11-1938, M. Cazier), 3 3 3 , 29 9 ; Lake City, Modoc
County (X-l 1-1952, E. Schlinger and J. Hall), 1 3 , 3 9 9 ;
Lake Cuyamaca (VII-5-1945, A. Melander), 39 9 ; Lake
Fontanillia, 8,500 feet, Eldorado County (VIII-21-1955,
E. Schlinger), 1$, 1 9 ; Lake Tahoe (VIII-11-1940, E.
Kenaga), 1 9 , (VIII-11-1940, L. Kuitert), 1 9 , (VII I -
11-1940, L. Lipovsky), 3$ S, 39 9 ; LaPosata Creek, San
Diego (IV-10-1950, L. Quate), 1 9 ; Leavitt Meadow,
Mono County (VII-6-1951, A. McClay), 1 9 , ( X - l l -
1952, J. Hall), 19 ; Likeley, Modoc County (VII-6-1951,
A. McClay), 19 , (X-ll-1952, E. Schlinger), 29 9 , (X-
11-1952, J. Hall), IS; Mammoth Lakes (V-29-1940, R.
Beamer), 3SS, 19 , (VII-29-1940, D. Hardy), 32$ S,
99 9 9, (VII-29-1940, E. Kenaga), 1$, 2 9 9 , (VII-29-
1940, L. Lipovsky), 9$ S, 119 9 ; Manzanita Lake, Lassen
National Park (IH-1-1941, J. Fisher), 1 $ ; Marion Moun-
tain Camp, San Jacinto Mountains (VII-1-1952, A. Mc-
Clay), 1 9 ; Mason Creek R. S., Modoc County (X-12-
1952, E. Schlinger), 2 3 3 , 4 9 9 ; Mono Lake, Mono County
(VII-31-1940, E. Kenaga), 1 9 ; Monterey County (VII -5-
1896, collector unknown), 1 3 , 19 , (VII-8-1896, W.
Wheeler), 1 3 , (VII-9-1896, W. Wheeler), 1 3 , 1 9 , (VII -
17-1896, G. Hough), 1 9 ; Morro Bay (VI1-27-1940, A.
Melander), 1 9 ; Ortega Highway, El Cariso Camp (V-26-
1944, A. Melander), 1 3 ; Pacific Grove (V-7-1906, J.
Aldrich), 39 9 , (V-13-1906, J. Aldrich), 19 (Vl-no
dates, W. Mann), 1 9 : Palm Springs (XII-14-17-1917, J.
Bradley) , 2 3 3 , (XI-19-1943, A. Melander), 1 $, 1 9, (XI-
22-1943, A. Melander), 1 3 , 29 9 ; Palomar Mountain
(VI-14-1948, A. Melander), 1 3 , 2 9 9 ; Pinecrest Lookout,
Tuolumne County (VII-7-1948, P. Arnaud), 1 3 , (VII -
30-1948, P. Arnaud), 1 9 ; Putah Canyon, Yolo County
NUMBER 6 8 89
(XI-10-1942, E. Schlinger), 1$ , (XI-6-1954, W. Lange),
1 3 ; Redwood City, San Mateo County (XII-26-1951, P.
Arnaud), 1 9 ; Riverside (XII-22-1934, A. Melander), 1 3 ,
(X-28-1944, A. Melander), 19 ; Sage Hen, 5 miles north-
west of Hobart Mills (VI-20- 1954, M. James), 1 3 , 3 9 9 ,
(VII-2-1954, J. Downey), 1 $ ; San Antonio R. S., Santa
Clara County (VI-27-1953, R. Schuster), 1 $ ; San Ber-
nardino Mountains, Cienaga (IX-28-1947, A. Melander),
1 3 , 1 $ ; San Diego (IV-no date-1913, E. Van Duzee),
1 9 ; San Gabriel River (IX-18-1949, M. Wheeler), 1 9 ,
(1-21-1950, M. Wheeler), 43 3 , 29 9 ; San Jacinto
Mountains (VII-21-1929, R. Beamer), 1 $ ; San Jose
Mission (X-no date-1897, collector unknown), 1 $ ; Sardine
Creek, Mono County (VII-2-1951, A. McClay), 2 9 9 ,
(VII-5-1951, A. McClay), 1 3 , (VII-6-1951, A. McClay),
2 9 9, (VII-11-1951, A. McClay), 1 5 , 2 9 9 , (VII-12-
1951, A. McClay), 19 , (VII-18-1951, A. McClay), 2 9 9 ,
(VII-29-1954, J. Downey), 1 9 ; Sequoia National Park
(VIII-6-1940, L. Kuitert), 1 9 ; Siberian Outpost, Tulare
County, 9,500-10,500 feet (VII-3-1915, collector un-
known), 1 9 ; Sonora Pass, 9,624 feet (VII-12-1951, A.
McClay), 1 9 , (VII-17-1951, A. McClay), 69 9 ; South
Fork, Santa Ana River (VII-29-1942, A. Melander), 1 3 ,
1 9 , (VIII-4-1945, A. Melander), 19 ; South Fork Camp,
Barton Flats (VIII-31-1944, A. Melander), 1 3 , ( I X - 3 -
1944, A. Melander), 1 9 ; South Fork Camp, Santa Ana
River, San Bernardino County (VI-28-1952, R. Beamer
et al.), 1 9 ; Stanford University, Santa Clara County
(XII-29-1897, collector unknown), 1 9 ; Strawberry,
Tuolumne County (VI-15-1951, A. McClay), 1 3 , 1 9 ,
(VI-19-1951, A. McClay), 1 3 , 1 9 , (VI-22-1951, A.
McClay), 29 9 ; Sugarloaf, Barton Flats (IX-17-1945, A.
Melander), 1 3 , 59 9 ; Sugarloaf Mountain (V-l 1-1947,
J. Sperry), 1 9 , (V-30-1947, J. Sperry), 29 9 ; Summit
Lake, Lassen National Park (VII-23-1950, L. Quate),
1 3 ; Temecula, Riverside County (IV-24-1951, E.
Schlinger), 1 9 ; Thousand Springs, San Bernardino Moun-
tains (X-24-1946, A. Melander), 59 $ ; Tioga Pass (VII-
31-1940, D. Hardy), 33 3 , 69 9 , (VII-31-1940, L.
Lipovsky), 29 9 ; Trinity River, Trinity County (VII-13-
1953, A. McClay), 19 , (VII-17-1953, A. McClay), 83 3 ,
139 9 , (VII-18-1953, A. McClay), 1 9 ; Truckee (VII -
5-1936, A. Pritchard), 1,5, (VII-6-1927, E. Van Duzee),
3$ $, 39 9 ; Tuolumne Meadows (VIII-1-1940, R.
Beamer), 1 3 , 29 9, (VII-1-1940, D. Hardy), 83 3 ,
209 9 ; Upper Santa Ana River (V-20-1947, A. Melander),
1 9 , (VI-18-1950, A. Melander), 1 9 , (IX-4-1950, A.
Melander), 2$ $, 1 9 ; Upper Santa Ana River, Cienaga
(V-28-1948, J. Sperry), 1 $ , 49 9 , (V-31-1948, J.
Sperry), 29 9 ; Yosemite Valley (V-22-1908, collector
unknown), 1 3 , 1 9 ; (VIII-1-1940, R. Beamer), 1 3 ,
(VIII-1-1940, D. Hardy), 1 3 , 29 9 , (VIII-1-1940, E.
Kenaga), 1 9 , (VIII-1-1940, L. Kuitert), 19 , (VIII -
1-1940, L. Lipovsky), 2 3 3 , 29 9 ; Yuba Pass, Sierra
County (VIII-20-1953, E. Schlinger), 1 9 ; 3 miles
southeast of Mount Lassen (VIII-8-1955, A. Mueller),
2 9 9 ; 10 miles west of Salinas near Carmel Valley,
Monterey County (IV-4-1953, J. Lattin), 1$.
Colorado: Cerro Summit, 9,500 feet (IX-5-1938, A.
Hardy), 1 9 ; Electra Lake, 37?33' N, 107?48' W, 8,400
feet (VI-28-VII-1-1919, collector unknown), 1 $ ; Emerald
Lake, Gunnison County (VIII-27-1961, D. L. Deonier),
13 ; Glade Peak, Mesa County (VII-8-1953, A. Gurney),
2$$, 2 9 9 , Golden (VI-17-1940, A. Melander), I $ ;
Lindland (VII-27-1938, M. James), 19 ; Ohio Pass Road,
Gunnison County, 38?42.2' N, 1O7?5.2' W (VIII-26-
1961, D. L. Deonier), 433 3, 55 9 9 ; Pingree Park (VIII -
13-1934, C. Sabrosky), 1 9 ; Rocky Mountain Biological
Laboratory, 9,500 feet, Gunnison County (VIII-27-1961,
D. L. Deonier), 5$$, 39 9 ; 3 miles north of Rocky
Mountain Biological Laboratory, Gunnison County (VIII?
27-1961, D. L. Deonier), 23 3, 39 9 ; 1 mile north of
Rocky Mountain Biological Laboratory (VIII-27-1961, D.
L. Deonier), 49 9 .
Idaho: Echo Bay, Coeur d'Alene Lake (VIII-3-1924,
A. Melander), 1 9 ; Juliaetta (no dates, J. Aldrich), 1 9 ;
Moscow Mountain (VIII-23-1908, collector unknown),
\$, 39 9 ; Potlatch (IX-21-1918, A. Melander), 1 3 ,
29 9 ; Priest Lake (IX-3-1919, A. Melander), 19 ; Swan
Lake, Bancock County (VI-19-1952, E. Schlinger), 1 9 ;
Yale (IX-10-1912, J. Aldrich), 19 .
Mexico: 70 miles west of Durango, Durango, 9,000 feet
(V-7-1961, Howden and Martin), 1$, 19 ; 10 miles east
of Toluca, Mexico, 9,000 feet (VIII-17-1954, J. Chillcott),
23 3, 29 9 .
Nevada: Beatty (V-26-1940, G. Bohart), 1 9 ; Double
Spring (VI-21-1916, H. Dyar), 13; Reno (X-20-1915,
H. Dyar), 29 9 .
New Mexico: Jemez Mountains (IX-10-1914, collector
unknown), 1 9 ; Tajique (VI-28-1947, R. Beamer), 19 .
Oregon: Crater Lake (IX-16-1934, A. Melander), 1 9 ;
Crater Lake National Park, near headquarters, 6,600 feet
(VII-30-1930, F. Wynd), 3$$, 89 9 ; Crater Lake
National Park, Pole Bridge Meadows, 6,000 feet (VIII-
15-1930, F. Wynd), 1 9 ; Crater Lake National Park, Sun
Creek Meadow, 6,500-7,000 feet (VIII-26-1930, H.
Scullen), 1 9 ; Fish Lake, Harney County (VII-16-1957,
J. Lattin), 1 9 ; Harney County (VI-29-1958, A. Gurney),
19 ; Pringle Falls, 9 miles west of La Pine, Deschutes
County (VII-13-1957, G. Kraft), 1 9 ; 22 miles north of
Prospect (VI-8-1948, Schuh and Gray), 2 3 .5, 1 9 ; Quartz
Mountain Service Station (VI-18-1934, J. Schuh), 19 .
Saskatchewan: Aftons Lake (IX-4-1940, A. Brooks),
19 .
Texas: Henkes Pond, Kerrville (IV-4-1955, W. Wirth),
1 3 , 1 9 .
Utah: Cedar City (VIII-13-1939, P. Oman), 1 3 , 7 9 9 ;
Echo (X-16-1943, G. Knowlton), 2 $ 6 ; Eden (VI-21-
1937, D. Hardy), 1 9 ; Emigrant Canyon, 7,000 feet,
Wasatch Mountains (VII-8-1911, J. Aldrich), 29 9 ;
Lake Cottonwood Canyon, 8,000 feet (VI-23-1940, A.
Melander), 7$ $, 139 9 ; Leeds (VI-27-1945, G. Knowl-
ton), 29 9 ; Provo Canyon (VIII-15-1940, R. Beamer),
1 9 ; Roberts Pass, Uinta Mountains, Duchesne County,
12,500 feet (VII-9-1946, collector unknown), 1 9 ;
Timpanogos Mountain (VI-25-1940, A. Melander), 23 3 ,
69 9 .
90 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Washington: Blue Mountains, Rose Spring (VI -19 -1921 ,
A. Melander), 1 $ ; Dungeness (VIII -24 -1910 , collector
unknown), 1?$, 1 $ ; Holland (VII -3 -1919 , A. Melander),
1 9 ; Kalama River (VII -21 -1931 , R. Beamer), 1 3 ;
Klickatack River, Glenwood (VI-27-1917 , A. Melander),
1 & ; Mount Rainier National Park, Berkeley Park ( V I I I -
23-1934, A. Melander), 1 9 ; Mount Rainier National Park,
Paradise Park ( V I H - n o date-1917, A. Melander), 1 9 ;
Mount Rainier National Park, Summerland (VIII -29-1934 ,
A. Melander), 2$ $, 4 $ 9 ; Mount Rainier National Park,
Yakima Park (VIII -19-1934 , A. Melander), 2&&, 1 9 ;
Olga (VII -26-1909 , collector unknown), 1 $ ; O'Sullivan
Dam, Grant County (X-30-1954 , H. Davis), 1 9 ; Pullman
(VI-16 -1912 , collector unknown), 2 9 9 , (X-17-1915 , A.
Melander), 1 <J ; Pullman, Lyles Grove (VIII -10-1923 , A.
Melander), 1 9 ; Uniontown (VI-26-1932 , J. Aldrich),
1 9 .
Wyoming: Kemmerer (VIII -14-1950 , M. Wheeler),
1 $ ; Yellowstone National Park, Geyser Basin ( V I I I - 7 -
1919, A. Melander), 1$, 2 9 9 ; Yellowstone National
Park, Norris Basin (VII -26-1923 , A. Melander), 1 9 .
REMARKS.?Except for H. decens, the male of
which is unknown, this species probably could be
confused only with H. melanderi. Hydrellia melan-
deri and H. proclinata differ conspicuously only in
the male terminalia, but the light-gray pleural spots
are fused in H. melanderi and separate in H.
proclinata.
Adult. Habitats recorded: Equisetem sp.,
Eleocharis sp., sedge meadow, and flowers of Prunus
emarginata. An observer found this species swarming
near some black scale infestations of trees at San
Jose Mission, California.
Hydrellia procteri Cresson
FIGURE 20
Hydrellia proctori Cresson, 1934, p. 235.
Hydrellia procteri; Cresson, 1936, p. 257 [emendation of
lapsus calami]; 1944b, pp. 169, 174.?Deonicr, 1964, p.
116; 1965, pp. 500, 506 [ecology].?Wirth, 1965, p. 744
[catalog listing].
DIAGNOSIS.?Palpus moderate yellow; 7-8 aristal
rays; epistomal index 1.0-1.4; wing length 1.70-2.13
mm; thoracic pleuron light gray except light-brown
upper edge of mesanepimeron; distal half of male
hind femur depressed and distal end of hind tibia
expanded slightly. Male length 1.50-2.04 mm;
female 1.70-2.04 mm. Male terminalia as in Figure
20.
HEAD.?Face light gray or yellowish gray; 4-5
pfa; epistomal index 1.0-1.4; mesofacial index 2.0-
2.5; vertex index 4.0-4.4; A-index 1.8-2.6; ocular
index 11.5-14.0. Palpus moderate yellow; 7-8
aristal rays; antenna and frons moderate brown;
13-16 postoculars.
THORAX.?Ppn and mesonotum dark gray; 3-4
adc and 2 pdc; pleuron light gray except light brown
upper edge of mesanepimeron; legs light-gray
pruinose except light-gray coxae and moderate-
yellow tarsi; distal half of male hind femur depressed
and distal end of hind tibia expanded slightly. Wing
length 1.70-2.13 mm; veins dark brown; 6-7 setae
on basal end of costa; 7-8 dorsal and 10-12 an-
terior interfractural costals; costal-section ratios:
I I : I 2.0-2.3; I I I : IV 3.0-3.5; V: IV 3.6-4.2; Mi?s
index 1.5-2.0.
ABDOMEN.?Terga dark brown medially, light
gray laterally and ventrally. Male terminalia: median
third of posterior margin of sternum 5 very slightly
concave (congruent with apex of distiphallus);
anterolateral margin of sternum 5 roundly obtus-
angular (about 100?) ; copulobus lanceolate; pos-
terior half of copulobus irregularly setose, anterior
half mostly bare. Postgonite bent anterolaterad;
postgonite uncus straight; distiphallus constricted
preapically and lamellate ventrally. Anterior margin
of fused surstyli broadly and slightly concave;
B-index about 5.0; C-index 1.0-1.5.
TYPE.?Holotype male, ANSP 6152.
TYPE-LOCALITY.?Bar Harbor, Mount Desert
Island, Maine (V-7-1933, W. Procter).
SPECIMENS EXAMINED.?52 (15 c? d \ 3 7 $ ? )
from 11 localities:
Connecticut: Southwest corner of Sleeping Giant State
Park, near Hamden (VI1-30-1961 , J. Laffoon), 1 9 .
Illinois: Chicago (VIII -29-1896 , W. Wheeler), 1 9 -
Iowa: Goose Lake, Hamilton County (VII -14 -1960 , D.
L. Deonier), 1 ? ; Ledges State Park, Boone County ( V I I I -
16-1960, D. L. Deonier), 1 3 ; Little Wall Lake, Hamilton
County (VIII -13-1960 , D. L. Deonier), 2$$, 8 9 9 ,
(VIII -27-1960, D. L. Deonier), 2 9 9 , ( I X - 2 2 - 1 9 6 2 , D.
L. Deonier), 1$, 1 9 ; Spring Lake, Greene County
( IX-19-1961 , D. L. Deonicr), 4 3 $, 1 2 9 9 ; Springbrook
State Park, west ?/2 sec. 33, T.81 north, R.31 west, Guthrie
County (VII-19-1960, D. L. Deonier), 1 9 .
Kansas: Marais des Cygnes Wildlife Refuge, Linn County
( I X - 5 - 1 9 6 1 , D. L. Deonier), 5$ $, 8 9 9 .
Maine: Bar Harbor, Mount Desert Island ( V I I - 1 5 - 1 9 3 3 ,
W. Procter), 1$ ; Mount Desert Island ( V I I - 2 5 - 1 9 3 5 , W.
Procter), 2 9 9 .
Michigan: Augusta ( X - 4 - 1 9 5 3 , collector unknown), 1 9 .
REMARKS.?This species seems to belong in the
NUMBER 6 8 91
H. crassipes group. Most of the similarities are in
coloration and modification of the male hind leg,
but there are some in the male terminalia. The disti-
phallus is ventrally lamellate in both H. crassipes
and H. procteri, and the posteromedial edge of
sternum 5 is heavily sclerotized in both.
Adult. Habitats recorded: leaves of Potamogeton
nodosus and Nelumbo lutea; limnic wrack, sedge
meadow, mud flat, and riffle rocks.
Hydrellia prudens Curran
FIGURE 18
Hydrellia prudens Curran, 1930, p. 78.?Cresson, 1944b,
pp. 164, 172.?Wirth, 1965, p. 744 [catalog listing].
DIAGNOSIS.?Palpus dark brown; 6-8 aristal rays;
vertex index 3.8-4.5; 1-3 sfa; antenna and para-
frontale not velvety; thoracic pleufon bronzed mod-
erate olive-brown; male mid tibia expanded. Male
length 1.53-1.67 mm; female 1.53-1.70 mm. Male
terminalia as in Figure 18.
HEAD.?Face light brown except lower third or so
light gray; 4-5 pfa; 1-3 sfa; epistomal index 1.0-
1.3; mesofacial index 1.7-2.0; vertex index 3.8-4.5;
A-index 2.0-2.6; ocular index 4.5-6.5. Palpus dark
brown; 6-8 aristal rays; antenna and most of para-
frontale very dark brown; fronto-orbital area and
frontal vitta moderate olive-brown; 12-16 post-
oculars.
THORAX.?Most of thorax and legs bronzed mod-
erate olive-brown; 3?4 adc and 2 pdc; male mid
tibia expanded; thoracic pleuron bronzed moderate
olive-brown. Wing length 1.62-1.87 mm; veins dark
brown; 5-7 setae on basal end of costa; 5-7 dorsal
and 6-8 anterior interfractural costals; costal-section
ratios: II:I 2.0-2.6; III:IV 2.8-3.4; V:IV 2.8-3.8;
Mi ?
 2 index 1.3-1.8.
ABDOMEN.?Terga bronzed moderate olive-brown.
Male terminalia: median third of posterior margin
of sternum 5 slightly concave and more heavily
sclerotized than remainder; anterolateral margin of
sternum 5 right-angled (90?-100?); copulobus not
prominently produced, slightly convex posteriorly
and the whole with 6-9 large scattered setae. Post-
gonite bent distinctly anteromediad then laterad;
postgonite uncus with end bent about 45? laterad;
distiphallus short, triangular, and furrowed ventrally
between 2 carinae. Anterior margin of fused surstyli
moderately concave medially; B-index 0.8-1.0;
C-index 0.8-1.0.
TYPE.?Holotype male, AMNH (not numbered).
TYPE-LOCALITY.?Station for Study of Insects,
Tuxedo, New York (VI-29-1928, C. H. Curran).
SPECIMENS EXAMINED.?7 (3cfo*j 4$ $ ) from
4 localities (Wirth, 1965, listed Idaho, Maine, New
York, and eastern Canada as the known distribution
for this species) :
Massachusetts: Woods Hole ( V I H - n o date-1922, A.
Sturtevant), 1$.
Mississippi: Lake Shady, Lamar County (VI -29 -1962 ,
D. L. Deonier), 1 $.
New York: Station for Study of Insects, Tuxedo ( V I -
29-1928, C. Curran), 1$, 3 5 $ .
Pennsylvania: Ohiopyle, Fayette County ( V I H - n o date-
1907, H. Kahn) , 1 $ .
REMARKS.?Cresson (1944, p. 164) listed Hydrellia
johnsoni Cresson as a full synonym of H. prudens.
I found this to be incorrect after examination of the
male terminalia in at least two specimens labeled by
Cresson as paratypes of H. johnsoni. The exact status
of H. johnsoni must remain undetermined until the
holotype terminalia can be studied.
Adult. Habitat recorded: leaves of Nuphar advena.
Hydrellia pulla Cresson
FIGURES 61, 106, 114
Hydrellia pulla Cresson, 1931, p. 108; 1944b, pp. 170,
175.?Berg, 1950, pp. 375, 376, 377, 379, 382, 383 (pi.
1, fig. 4), 384, 385 (pi. 2, fig. 5) , 386, 387, 388, 389 (pi.
3, fig. 5), 390, 396. [biology, morphology, and taxonomy
of immatures].?Judd, 1964, p. 412. [biology].?Wirth,
1965, p. 744 [catalog listing].
DIAGNOSIS.?Palpus moderate orange-yellow; 7?9
aristal rays; pfa and sfa nonseriated (4-7 of each);
body length: wing length 0.9-1.0; face carinate
and conically prominent on lower half; 2-3 basal cox-
als. Male length 2.38-2.55 mm; female 2.55-3.15
mm. Male terminalia as in Figure 61.
HEAD.?Face light gray, carinate and conically
prominent on lower half; 4-7 nonseriated pfa; 2-5
sfa more or less parallel to pfa; 2 sfa, ventrally
directed and above pfa; epistomal index 1.0-1.4;
mesofacial index 1.7-2.0; vertex index 4.5-6.5;
A-index 1.6-1.9; ocular index 4.3-5.2. Palpus mod-
erate orange-yellow; 7-9 aristal rays; antenna,
fronto-orbital area, and frontal vitta light brown;
92 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
most of parafrontale velvety dark brown; 14-20
postoculars.
THORAX.?Ppn and mesonotum dark brown; 3-4
adc and 2 pdc; pleuron light gray except strong
yellowish brown on posterior half of mesanepisternum
and on mesokatepisternum and laterotergite; 2-3
basal coxals (1 macrochaetous) ; legs sparsely strong
yellowish-brown pruinose except dark-brown tarsi.
Wing length 2.21-3.06 mm; veins dark brown; 9-14
setae on basal end of costa; 7-11 dorsal and 9-13
anterior interfractural costals; costal-section ratios:
11:1 2.3-3.6; IILIV 2.6-3.8; V:IV 3.4-4.6; M, ?
 2
index 1.2-1.6.
ABDOMEN.?Terga dark brown medially and light
gray laterally and ventrally. Male terminalia: median
third of posterior margin very slightly concave;
anterolateral margin of sternum 5 obtusangular
(about 110?); copulobus acutangular posteriorly,
verrucate anteromedially, and irregularly setose.
Postgonite bent laterad; postgonite uncus buttonlike;
pregonite breadth 2-3 times postgonite breadth;
distiphallus tapering distally and ventrally bicarinate.
Anterior margin of fused surstyli slightly concave;
B-index about 1.0; C-index 3!0-4.0.
EGG.?Length 0.57-0.65 mm; maximum breadth
0.18-0.22 mm. Chorion indistinctly corrugate, with
longitudinal ridges anastomosing infrequently; end
opposite micropylar protuberance acute. Micropylar
protuberance infundibulate, but long and visible in
dorsal view. This description in part after Berg
(1950).
FIRST-LARVAL INSTAR.?Length 1.00-1.80 mm;
maximum breadth 0.18-0.25 mm. Frontoclypeal
length 0.18-0.22 mm. Opaque, yellowish gray. This
description in part after Berg (1950).
SECOND-IN STAR LARVA.?Length 1.80-3.50 mm;
maximum breadth 0.25-0.65 mm. Frontoclypeal
length 0.28-0.35 mm. This description in* part after
Berg (1950).
THIRD-INSTAR LARVA.?Length 3.35-6.00 mm;
maximum breadth 0.80-1.20 mm. Frontoclypeal
length 0.50-0.66 mm; feeding apparatus hyaline
except mouth-hook and cheliform spot (Figure 106).
Ventral frontoclypeal index 4.8-5.2; phragmatal
index 1.2-1.4; bifurcation index 3.5-3.8; clypeal-
arch index 1.8-2.0. Clypeal arch angled 20?-25?
in relation to lower frontoclypeal margin. Mouth-
hook beak distinctly longer than base; maximum
mouth-hook base thickness about 2.0 times that of
beak; mouth-hook light spot small, elliptical. Pro-
thorax and mesothorax sparsely spinulose; abdom-
inal segment 8 without ventral, transverse row of
setulae, but with many irregular dark spinules ven-
trally. Body opaque yellowish gray.
PUPARIUM.?Length 3.35-5.25 mm; maximum
breadth 0.90-1.50 mm; subcylindrical (Figure 114).
Puparial length: minimum breadth 15.0-16.0; maxi-
mum breadth: minimum breadth 3.5?4.5; anal-plate
index 2.5-3.0. Prothoracic end slightly convex or
slightly triangular in ventral view; head-lobe scar
subcircular; anal plate crescentric, with anterior
margin distinctly convex. Empty puparium trans-
lucent, light brown.
TYPE.?Holotype female, ANSP (not numbered).
TYPE-LOCALITY.?Spencer Lake, New York (VI-
30-1907, collector unknown). Labeled "loaned
property of Cornell."
ADULT SPECIMENS EXAMINED.?84 (17 c? cf,
6 7 ? ? ) from 15 localities:
British Columbia: Elk Lake, Vanonov Island (VII?11-
1934, A. Melander), 19 .
Michigan: Douglas Lake, Cheboygan County (VII-17-
1941, C. Berg), 1 <$, 1 $ ; Huron River, Washtenaw County
(V-15-1941, C. Berg), 1 $ ; Ocqueoc Lake, Presque Isle
County (VII-13-1941, C. Berg), 1 $ ; Third Sister Lake,
Washtenaw County (IV-22-1940, C. Berg), 1 9 , (V-20-
1940, C. Berg), 19 , (VII-17-1941, C. Berg), 1 9 , ( I I -
2-1942, reared by C. Berg), 2 9 9 , (111-30-1942, C. Berg),
3 9 9, (V-3-1942, C. Berg), 5$ $, 23 9 9 , (VI-12-1942,
C. Berg), 2 9 9 , (VI-13-1942, C. Berg), 1 9 ; Whitmore
Lake, Washtenaw County, (III?5?1941, reared by C. Berg),
2$ $, (111-11-1941, reared by C. Berg), 1 9 .
Minnesota: Squaw Lake, Itasca State Park (VIII-14-
1963, D. L. Deonier), 19 ; Two Island, Itasca State Park
(VI-28-1963, D. L. Deonier), 1 3 , (VIII-8-1963, D. L.
Deonier), 7 9 9 , (VIII-18-1963, D. L. Deonier), 1 9 ;
1.3 miles south of main entrance, Itasca State Park (VI-
26-1963, D. L. Deonier), 79 9 .
New York: Ithaca (no dates or collector), \$.
Ontario: London (V-20-1958, W. Judd), 1 9 , (V-21-
1958, W. Judd), 1 9 , (VII-16-1958, W. Judd), 1$,
(VII-19-1958, W. Judd), 1 $ ; Midland (VIII-18-1955,
J. Chillcott), 3S $, 69 9 .
Quebec: Lac Bernard (VIII-7-1938, G. Shewell), I S ,
3 9 9 .
Washington: Orcas Island above Mountain Lake (VIII -
18-1925, A. Melander), 1 9 .
Wisconsin: Squaw Lake, Vilas County (VIII-19-1954,
J. Laffoon), 1 <$.
IMMATURE SPECIMENS EXAMINED: 67 from 4
localities:
Michigan: Cheboygan County (collected and reared by
NUMBER 68 93
C. Berg); Washtenaw County (collected and reared by C.
Berg).
Minnesota: Two Island Lake, Itasca State Park (col-
lected and reared by D. L. Deonier); 1.3 miles south of
main entrance Itasca State Park (collected and reared by
D. L. Deonier).
REMARKS.?The prominent face bordered by the
nonseriated primary facials together with its rela-
tively large size makes this species recognizable even
macroscopically in its habitat.
Adult. Habitats recorded: leaves of Potamogeton
amplifolius, P. gramineus, P. natans, Sagittaria
latijolia, and Nuphar advena.
Many observations of this conspicuous species
yielded little information on its behavior. In one in-
stance, a female caught a live chironomid and acted
very protective of her catch when a female H.
luctuosa darted in near it. On another occasion, a
female H. bergi attacked a female H. pulla. I col-
lected several adults by the lighted-receptacle method.
Berg found eggs in masses one layer deep con-
cealed in leaf folds, stipules, and broken mines of
Potamogeton species from June through August. I
observed a female return repeatedly over a 10-minute
period to an old aphid exoskeleton on a Sagittaria
leaf into which she inserted her ovipositor each time;
however, I found no eggs in it. In Berg's laboratory,
adults mated and oviposited within 48 hours after
emergence.
Egg and larva. According to Berg, the incubation
period ranged from 4 to 6 days, the first larval sta-
dium 4 to 8 days, and the second and third larval
stadia 6 to 10 and 5 to 15 days, respectively. In one
case, I found the third larval stadium to be 4 days.
All of these measurements were made in the
laboratory.
Puparium. The third-instar larva pupariates with
its spiracular peritremes inserted in the leaf midrib
or petiole. The puparial phase ranged from 10 to
15 days. Berg found several puparia in two bluegill
stomachs.
Host plants. I found larvae and puparia of this
species only in Potamogeton amplifolius. Berg found
them in P. amplifolius, P. gramineus, and P.
richardsonii.
Parasites. Berg (1950, p. 386) stated that "Appar-
ently H. pulla is much less susceptible to parasitism
than any other species of Hydrellia studied in this
investigation. Although H. pulla was often observed
closely associated with H. cruralis, the two braconid
parasites Chorebidea and Ademon, so frequently
reared from the latter, were never encountered in
puparia of the former." He reared only Trichopria
columbiana from H. pulla puparia. He found no
parasites in 80 H. pulla puparia collected along with
71 of H. cruralis from which 12 braconids emerged.
I reared Ademon niger and T. columbiana from
H. pulla puparia. Of 61 puparia collected or reared,
39 harbored hymenopterous parasites.
In one case of parasitism, a third-instar larva of
H. pulla fed actively from 8 July to 12 July, when it
pupariated. On 12 July, shortly after pupariation,
I observed a leechlike larva of Ademon niger moving
in the anterior part of the puparium. A subsequent
examination revealed that the anterior half of
the puparium was devoid of tissue. The adult
hymenopteron emerged 25 July. In another case, the
hymenopteron emerged 18 days after pupariation.
In four puparia, I found pharate adult hymenop-
terans with their ventral side toward the puparial
dorsum, and in another one, the hymenopteron's
head faced the posterior end of the puparium.
Several observations on the behavior of T. colum-
biana adults support previous statements about its
oviposition habit. Four adults lived submerged for
24 hours. In one instance, an adult emerged from
the puparium and surfaced immediately on 2
August. On 5 August, this adult submerged and
deposited three eggs near its former host puparium.
The hymenopteron held its wings in repose while
moving about submerged. She scraped a small bubble
loose from the wings and then tried to surface. The
next day she was alive on the surface.
Hydrellia rixator, new species
FIGURE 29
DIAGNOSIS.?Palpus moderate yellow; prementum
glossy brownish black; 6-8 aristal rays; vertex index
5.5-7.5; nearly all of frons velvety dark brown; wing
length 1.70-2.04 mm; body length:wing length 1.0-
1.2. Male length 1.50-1.70 mm; female 1.96-2.47
mm. Male terminalia as in Figure 29.
HEAD.?Face light yellowish brown; 4-6 pfa;
epistomal index 1.2-1.5; mesofacial index 2.0-2.3;
vertex index 5.5-7.5; A-index 1.3-1.7; ocular index
94 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
5.3-6.0. Palpus moderate yellow and triangular;
6-8 aristal rays; prementum glossy brownish black;
antenna dark brown; posterior half of fronto-orbital
area strong yellowish brown; remainder of frons
velvety dark brown; 12-16 postoculars.
THORAX.?Ppn and notopleuron yellowish gray;
remainder of mesonotum strong yellowish brown;
3-4 adc and 2 pdc; pleuron light gray; legs light-
gray pruinose except dark-brown tarsi. Wing length
1.70-2.04 mm; veins light yellowish brown; 6-7
setae on basal end of costa; 6-10 dorsal and 8-11
anterior interfractural costals; costal-section ratios:
11:1 1.6-2.3; 111:1V 2.7-3.2; V:IV 2.8-3.6; Mi * j
index 1.2-1.8.
ABDOMEN.?Terga light gray except medial parts
of 2-4 moderate brown. Male terminalia: median
third of posterior margin of sternum 5 concave and
congruent with distiphallus; median third of an-
terior margin of sternum 5 broadly notched; antero-
lateral margin of sternum 5 smoothly rounded, with
linear, bladelike process projecting laterad from
above; copulobus irregularly setose, with incurved
posterior arm bearing 5-6 fusiform macrochaetae
distally and a shorter posteriorly directed medial arm
hearing 3-4 macrochaetae distally. Postgonite bent
posteromediad and postgonite uncus anteromediad;
distiphallus short and nearly uniform in breadth to
ovoid apex. Anterior margin of fused surstyli with
3 deep clefts; B-index about 2.0; C-index 1.5-2.0.
TYPE.?Holotype male, USNM 70540.
TYPE-LOCALITY.?Samburg, Reelfoot Lake, Ten-
nessee, 36?22.9' N, 89?21.3' W (VIII-12-1962,
D. L. Deonier).
PARATYPES.?9 (4cf o*, 5 ? $ ) from 4 localities:
Kentucky: Fulton, Fulton County (no dates, W. Phillips),
1<$ (ISC).
Tennessee: Samburg, Reelfoot Lake (VIII-12-1962, D.
L. Deonier), 49 9 (USNM and ISC).
Texas: Kerrville (VH-no date-1954, L. Bottimer), 19
(USNM); Laguna Madre, 25 miles southeast of Harlington
(VII-7-1945, D. Hardy), 3$ g (USNM and ISC).
REMARKS.?Except for some inconspicuous color
differences and the ratio of body length to wing
length, this species could be readily confused physiog-
nomically with H. griseola or with the predominantly
Neotropical H. spinicornis.
Adult. Habitats recorded: leaves of Heteranthera
dubia and Sagittaria australis.
Hydrellia saltator, new species
FIGURE 56
DIAGNOSIS.?Palpus moderate yellow; 7-10 aristal
rays; ocular index 6.0-7.0; antenna velvety black;
2 basal coxals; male hind femur grooved ventrally
and hind tibia flanged, but not so distinctly as in
H. crassipes. Male length 2.72 mm; female length
3.06 mm. Male terminalia as in Figure 56.
HEAD.?Differing mainly from H. crassipes in the
following: mesofacial index 1.7-2.0; ocular index
6.0-7.0; antenna velvety black.
THORAX.?Differing mainly from H. crassipes in
the following: 2 basal coxals; wing length 2.47-2.98
mm; 7-10 dorsal and 10-12 anterior interfractural
costals.
ABDOMEN.?Terga semiglossy dark brown medi-
ally, light gray laterally and ventrally. Male termi-
nalia: median third of posterior margin of sternum
5 obtusely recessed; anterolateral margin of sternum
5 rounded through about 120? angle; copulobus
acutangular posteriorly (about 20?) and irregularly
setose except for medial marginal row. Postgonite
bent anterolaterad; postgonite uncus straight, trun-
cate, long, and directed laterad; distiphallus taper-
ing to mucronate apex (except small preapical
constriction) ; distiphallus carinate and finely lamel-
late ventrally. Anterior margin of fused surstyli
narrowly notched medially to about one-third the
length of structure, with about 6-8 setae on
anterolateral corners; B-index about 3.6; C-index
1.2.
TYPE.?Holotype male, CNC 11652.
TYPE-LOCALITY.?Grand Bend, Ontario, Canada
(VII-10-1939, G. E. Shewell).
PARATYPE.?1 $ with same data as holotype.
These are the only known specimens of this species.
REMARKS.?To distinguish this sibling of H.
crassipes, it may be necessary to examine the male
terminalia. One should not place too much reliance
on the basal-coxal differential at this time.
Hydrellia serena Cresson
FIGURE 35
Hydrellia serena Cresson, 1931, p. 104; 1934, p. 236; 1944b,
pp. 165, 174.?Wirth, 1965, p. 744 [catalog listing].
DIAGNOSIS.?Palpus moderate yellow; prementum
pale yellow; 7-9 aristal rays; ocular index 5.8-6.8;
NUMBER 6 8 95
most of parafrontale velvety black; thoracic pleuron
light gray except upper fifth dark brown and mesan-
episternum bronzed in posterolateral view; 2-3 basal
coxals; 7-10 anterior interfractural costals. Male
length 1.79-2.24 mm; female 2.18-2.40 mm. Male
terminalia as in Figure 35.
HEAD.?Face light gray or yellowish gray; 4-5
pfa; epistomal index 1.2-1.6; mesofacial index 1.9-
2.3; vertex index 6.0-9.0; A-index 1.4-2.2; ocular
index 5.8-6.8. Palpus moderate yellow; 7-9 aristal
rays; prementum pale yellow; antenna dark brown;
fronto-orbital area and frontal vitta strong yellowish
brown; most of parafrontale velvety black; 14-16
postoculars.
THORAX.?Ppn and notopleuron strong yellowish
brown; rest of mesonotum dark brown; 3 -4 adc
and 2-3 pdc; pleuron light gray except upper fifth
dark brown and mesanepisternum bronzed in
posterolateral view; legs light-gray pruinose except
dark-brown tarsi. Wing length 2.04-2.64 mm; wing
veins dark brown; 6-9 setae on basal end of costa;
6-9 dorsal and 7-10 anterior interfractural costals;
costal-section ratios: I I : I 2.0-2.5; I I I : I V 2.5-3.0;
V . I V 3.0-3.4; Mi ?
 2 index 1.3-1.7.
ABDOMEN.?Terga light brown dorsally and light
gray ventrally. Male terminalia: median third of
posterior margin of sternum 5 narrowly notched
(almost congruent with distiphallus); anterolateral
margins of sternum 5 acutangular ( 4 5 ? - 5 0 ? ) ;
anterior and lateral margins of sternum 5 distinctly
more heavily sclerotized; copulobus truncate pos-
teriorly and irregularly setose. Postgonite bent antero-
laterad and postgonite uncus slightly laterad; disti-
phallus apex lanceolate; basiphallus not nearly
covered laterally by surstyli. Anterior margin of fused
surstyli deeply and narrowly cleft medially, with 7-9
setae conspicuous on each corner; B-index about
5.0; C-index 0.3-0.8.
TYPE.?Holotype male, ANSP 6482.
TYPE-LOCALITY.?Ilwaco, Washington ( V H - n o
date-1917, A. L. Melander).
SPECIMENS EXAMINED.?181 ( 7 0 o * d \ 1 1 1 ? ? )
from 42 localities:
Alaska: Anchorage (VI-19-1921, J. Aldrich), 1 3 ; King
Salmon, Naknek River (VII-10-1952, W. Mason), 13 ,
69 9, (VII-23-1952, W. Mason), 19 , (VIII-1-1952,
W. Mason), 63 3 , 69 9 , (VIII-11-1952, W. Mason),
43 3 , 99 9 ; Savonoski, Naknek Lake (VII-26-1919, A.
Basinger), 13 , 1 9 ; Seward (VH-24-1921, J. Aldrich),
1 9 ; Sitka (VII-10-1907, W. Shaw), 19 .
Alberta: Manyberries (VI-4-1956, O. Peck), 18.
British Columbia: Atlin (VII-7-1955, H. Huckel), 73 3 ,
179 9 ; (VII-20-1955, H. Huckel), 39 9 , (VIII-4-1955,
B. Gibbard), 1 9 ; Huntingdon (VI-30-1953, W. Mason),
1 $ ; Masset, Queen Charlotte Islands (VI-3-1957, E.
MacDougall), 3 3 3 , 99 9 ; Milner (VII-12-1953, G.
Spencer), 29 9 ; Mission City (VI-6-1953, W. Mason),
1 3 , 69 9 , (VI-9-1953, W. Mason), 1 3 , 1 9 , (VI-18-
1953, E. Mason), 1 3 , (VI-20-1953, W. Mason), 19 ,
(VII-2-1953, E. Mason), 2 9 9 , (VII-5-1953, G.
Spencer), 19 , (VII-10-1953, W. Mason), 1 3 ; Moresby
Camp, Queen Charlotte Islands (V-31-1957, E. Mac-
Dougall), 1 3 ; peat bog at Pitt Meadows (VII-9-1953,
W. Mason), 1,5, 29 9 ; Vancouver (VI-20-1958, H. and
A. Howden), 1 9 ; Victoria (VII-10-1924, A. Melander),
1 3 , 2 9 9 .
Colorado: Bear Lake, Estes Park, 1,000 feet (VII-11-
1934, A. Melander), 1 9 ; Cameron Pass (VIII-19-22-1940,
C. Sabrosky), 1 3 ; Marshall Pass, 10,856 feet (VII-28-
1906, collector unknown), 13 , 1 9 ; Rocky Mountain
Biological Laboratory, Gunnison County, 9,500 feet (VIII-
26-1961, D. L. Deonier), 13 .
Idaho: Pottsville (VIII-23-1916, A. Melander), 13-
Manitoba: Churchill (VII-8-1952, J. Chillcott), 1 9 ,
(VII-30-1952, J. Chillcott), 1 3 ; Farnsworth Lake near
Churchill (VII-14-1952, J. Chillcott), 1 9 ; Fort Churchill
(VI-24-1952, J. Chillcott), 2 9 9 , (VII-4-1952, J. Chill-
cott), IS, (VII-10-1952, J. Chillcott), 1 9 ; Mile 505 of
Hudson Bay Railway (VI-29-1952, J. Chillcott), 1 9 ,
(VII-16-1952, J. Chillcott), 19 , (VII-26-1952, J. Chill-
cott), 19 .
Northwest Territories: Alakvik (VII-27-1932, Bryant),
IS.
Oregon: Marshfield (VI-27-no year, J. Aldrich), 39 9 ;
Newport (VI-9-1925, E. Van Dyke), IS.
Washington: Bellingham (VII-no date-1930, collector
unknown), 43 3 , 39 9 ; Blaine (VIII-7-1917, A.
Melander), IS; Everett (VI-19-1920, A. Melander),
IS, (VI-20-1920, A. Spuler), IS; Humptulips (IX-5-
1934, A. Melander), IS, 1 9 ; Ilwaco (VII-no date-1917,
A. Melander), 2 3 3 , 2 9 9 , (VIII-27-1917, A. Melander),
1 $ , (Vl-10-1918, A. Spuler), 2 3 3 , 2 9 9 , (VI-18-1918,
A. Spuler), 63 3 , 79 9, (VII-no date-1918, O. Miner),
2 9 9 , (VI-28-1925, A. Melander), 1 3 ; Longmire (VI-
27-1935, A. Melander), 1 3 ; Mount Constitution (VII-
31-1909, collector unknown), 1 3 , 1 9 ; Olga (V-17-1910,
collector unknown), 1 9 ; Olympia (VII-17-1922, A.
Melander), 1 3 ; Pluvius (VII-16-1922, A. Melander),
1 3 , 1 9 ; Point Gamble (VIII-16-1910, collector unknown),
1 3 ; Poulaho (VIII-17-1910, collector unknown), 1 3 ;
Roche Harbor (VII-3-1909, A. Melander), 1 3 ; Seaview
(VI-28-1925, A. Spuler), 33 3 , 3 9 9, (VI-30-1925, A.
Spuler), 1 3 ; Sequim (VIII-2-1951, collector unknown),
13.
REMARKS.?Adult. Habitats recorded: Equisetum
sp. at 9,500 feet, peat bog, and unspecified at 10,856
feet.
96 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Hydrellia spinicornis Cresson
FIGURES 40, 78, 93, 103, 105, 117, 124
Hydrellia spinicornis Cresson, 1918, p. 48 (pi. 3, fig. 5 ) ;
1947a, p. 38.?Wirth, 1968, p. 13 [catalog listing].
DIAGNOSIS .?Palpus moderate yellow; 6-8 aristal
rays; mesofacial index 2.6-3.4; body length:wing
length 0.9-1.0; thoracic pleuron light gray; distinct
black anteroventral spines on distal half of female
fore femur; costal section I I : I 3.0-3.7. Male length
1.53-1.75 mm; female length 1.70-1.87 mm. Male
terminalia as in Figure 40.
HEAD.?Face light gray; 5-7 pfa; epistomal index
1.5-1.9; mesofacial index 2.6-3.4; vertex index 8.0
or more; A-index 1.7-2.3; ocular index 5.8-7.6.
Palpus moderate yellow; 6-8 aristal rays; antenna
and most of parafrontale dark brown; fronto-orbital
area and frontal vitta strong yellowish brown; 13-16
postoculars.
THORAX.?Ppn light gray; mesonotum strong yel-
lowish brown; 3-4 adc and 2 pdc; pleuron light
gray; legs light-gray pruinose except moderate-
yellow trochanters and dark-brown tarsi. Wing
length 1.53-1.87 mm; veins light yellowish brown;
6-8 setae on basal end of costa; 5-7 dorsal and
6-10 anterior interfractural costals; costal-section
ratios: I I : I 3.0-3.7; I I I : IV 2.6-3.5; V: IV 3.4-4.2;
Mi ?
 2 index 1.4-1.7.
ABDOMEN.?Terga moderate brown medially,
light gray laterally and ventrally. Male terminalia:
median third of posterior margin of sternum 5 con-
vex, with paired, posteriorly directed bispinate
processes; anterolateral margin of sternum 5
rounded; copulobus rounded or nearly truncate
posteriorly and irregularly setose. Postgonite directed
anteromediad, with short, medially incurved uncus;
distiphallus uniform in breadth (except preapical
constriction) and covered by sternum 5. Anterior
margin of fused surstyli deeply cleft medially to
central gibba; surstyli with acute anterolateral
processes; B-index about 3.3; C-index 8.0-10.0.
EGG.?Length 0.40-0.55 mm; maximum breadth
0.14?0.15 mm. Chorion (Figure 78), yellowish gray,
corrugate, with the few longitudinal ridges occa-
sionally anastomosing; spaces between ridges regu-
larly micropunctate. Micropylar protuberance in-
fundibulate, exposed in dorsal view.
FIRST-INSTAR LARVA.?Length 0.38-1.20 mm;
maximum breadth 0.06-0.20 mm. Clypeal arch dis-
tinctly angular (about 100?); feeding apparatus
black; frontoclypeal length 0.14-0.18 mm (Figure
105). Heavily sclerotized supraspiracular spinous
processes present.
SECOND-INSTAR LARVA.?Length 0.90-3.50 mm;
maximum breadth 0.20-0.80 mm. Clypeal arch not
so angular. Frontoclypeal length 0.24-0.28 mm
(Figure 93). Spiracular peritreme distinct as in Fig-
ure 79; 2 bispinate asteriform processes and 4-6
smaller spines just anterodorsal to each spiracular
peritreme; peritreme extraordinarily long and spicate.
Body translucent, yellowish gray.
THIRD-INSTAR LARVA.?Length 2.00-4.50 mm;
maximum breadth 0.50-1.00 mm. Frontoclypeal
length 0.36-0.42 mm (Figure 103). Ventral fronto-
clypeal index 3.4-3.6; phragmatal index 0.8-0.9;
bifurcation index 3.4-3.8; clypeal-arch index 1.4?
1.6. Clypeal arch sloping at 20?-30? in relation to
lower frontoclypeal margin. Mouth-hook beak and
base indistinct; mouth-hook light spot narrow, tri-
angular, and about 3.0 times longer than wide.
Supraspiracular bispinate processes absent or at least
indistinct; prothorax densely spinulose, abdominal
segment 8 without ventral, transverse row of setulae,
but with 4-5 annuli of spinules. Body opaque, with
yellow tinge.
PUPARIUM.?Length 3.00-4.00 mm; maximum
breadth 0.75-0.90 mm; fusiform (Figure 117).
Puparial length: minimum breadth 18.0-20.0; maxi-
mum breadth: minimum breadth 3.5-4.5; anal-plate
index 2.6-3.2. Prothoracic end semicircular in
ventral view; head-lobe scar triangular or obovoid;
maximum puparial breadth 1.5-1.7 times maximum
prothoracic breadth; anal plate reniform, with
anterior margin convex. Empty puparium trans-
lucent, light yellowish brown.
TYPE.?Holotype male, ANSP 6121.
TYPE-LOCALITY.?Alajuela, Costa Rica, sweeping
at 3,100 feet (IX-15-1909, P. A. Calvert).
ADULT SPECIMENS EXAMINED.?328 (190cfcf,
138$ $ ) from 5 localities:
Florida: Lacoochee (IX-18-1930, P. Oman), 49 9 .
Georgia: Rabun Bald, Rabun County (VIII-9-1957, W.
Richards), 1 3 .
Mississippi: Dickinson's Pond, sec. 3, T.4 north, R.14
west, Lamar County (VII-11-1962, D. L. Deonier), 79 3 3 ,
58 9 9, (VII-24-1962, D. L. Deonier), 70 3 3 , 4 7 9 9 ;
Lake Shady, Lamar County (VI-29-1962, D. L. Deonier),
23 3 , 49 9 , (VII-11-1962, D. L. Deonier), 383 3 ,
249 9 ; Vancleave Road, Jackson County (VI-28-1962,
D. L. Deonier), 19.
NUMBER 6 8 97
IMMATURE SPECIMENS EXAMINED.?26 from 2
localities:
Mississippi: Dickinson's Pond and Lake Shady, Lamar
County (collected and reared by D. L. Deonier).
REMARKS.?I have determined my material as Hy-
drellia spinicornis without access to the holotype; how-
ever, paratype comparison was made, and it seems un-
likely the material represents a new species. It is so
similar to Hydrellia griseola that a study of body
and wing lengths and male tenninalia was necessary
to detect its different status. Hydrellia spinicornis is
definitely a cryptic member of the H. griseola species
group.
Adult. Habitats recorded: leaves of Hydrochloa
caroliniensis, Nymphaea odorata, and Nelumbo
lutea.
Several observations yielded no information on
adult behavior.
Egg and larva. I collected a few eggs from leaves
of H. caroliniensis, but I did not obtain any data
on the incubation period. The larvae left evidence
that they initially enter the mesophyll of the leaf
sheath in most cases and then mine into the exceed-
ingly thin leaf blade. I had too few larvae to allow
them to develop sufficiently to measure the stadia.
Puparium. Most of the puparia I collected con-
tained late pupae. Mines in stems and stolons con-
taining puparia had an escape slit either lateral to
or slightly anterior to the operculum of the pupa-
rium, while those in leaves usually had none. The
larvae apparently made these slits before they
pupariated.
Host plants. I found larvae and puparia only in
Hydrochloa caroliniensis, a water grass growing in
vast mats in bay-tree swamps and small ponds
bordering lakes in southern Mississippi. Infestation
ranged from sparse to moderate in the three locali-
ties examined. Plants collected from isolated pools
during a late summer drought in September had a
lower percent infestation. The drought may have
indirectly decreased the local population of H.
spinicornis by causing numerous larvae to become
trapped in drying mats of water grass.
I examined samples of the following additional
plant species for immatures of H. spinicornis:
Juncus repens, Potamogeton diversifolius, Polygonum
hyperpiperoides, Xyris caroliniana, and Hypericum
punctatum.
Parasites. One undetermined hymenopteron
emerged through the posterior end of a puparium.
Hydrellia subnitens Cresson
FIGURE 45
Hydrellia subnitens Cresson, 1931, p. 106; 1936, p. 258;
1941, p. 37; 1944b, pp. 169, 174.?Wirth, 1965, p. 744
[catalog listing].
DIAGNOSIS.?Palpus moderate yellow; 8-9 aristal
rays; epistomal index 1.4-3.0; ocular index 9.0-
13.0; thoracic pleuron light gray; costal section I I : I
2.4-3.0. Male length 2.04-2.21 mm; female 2.04-
2.89 mm. Male terminalia as in Figure 45.
HEAD.?Face light gray or yellowish gray; 4-8
pfa; epistomal index 1.4-3.0; mesofacial index 2.0-
4.8; vertex index 4.4-6.3; A-index 2.1-2.6; ocular
index 9.0-13.0. Palpus moderate yellow; 8-9 aristal
rays; antennal segment 3 moderate yellow or dark
brown; antennal segments 2 and 3, fronto-orbital
area, and frontal vitta light brown; most of para-
frontale dark brown; 14-17 postoculars.
THORAX.?Ppn and mesonotum dark brown; 4 adc
and 2 pdc; pleuron light gray; 1-2 basal coxals
(1 macrochaetous); legs light-gray pruinose except
moderate-yellow tibial apices and tarsi. Wing length
2.47-3.23 mm; veins light yellowish brown; 6-8
setae on basal end of costa; 7-9 dorsal and 8-14
anterior interfractural costals; costal-section ratios:
I I : I 2.4-3.2; I I I : IV 2.5-3.5; V:IV 3.6-4.6; Mi ?
 2
index 1.3-1.6.
ABDOMEN.?Terga dark brown medially, light
gray laterally and ventrally. Male terminalia: median
third of posterior margin of sternum 5 concave;
anterolateral margin of sternum 5 prominent (30?-
40? angular; this prominence bare); copulobus long,
linguiform, curved slightly laterad, irregularly setose,
and verrucate anterolaterally. Postgonite bent antero-
laterad; postgonite uncus straight (directed antero-
laterad) ; distiphallus tapering to nearly truncate
apex; distiphallus lamellate ventromedially. Anterior
margin of fused surstyli broadly concave, with medial
cleft; B-index about 2.0; C-index 0.3-0.5.
TYPE.?Holotype male, ANSP 6484.
TYPE-LOCALITY.?Tacoma, Washington (VIII-
27-1911, A. L. Melander).
98 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
S P E C I M E N S EXAMINED.?4 (1 cT, 3 ? ? ) from 3
localities:
Oregon: Oregon City (IX-no other dates, collector
unknown), 19-
Washington: Tacoma (VIII-27-1911, A. Melander),
IS, 1 9 .
Wyoming: 12 miles northwest of Lusk (VH-no date-
1895, collector unknown), 19 .
Hydrellia surata, new species
FIGURE 25
DIAGNOSIS.?Palpus velvety black; 5-8 aristal
rays; vertex index 3.8-4.5; 4-7 sfa; antenna and
most of parafrontale velvety black; face centrally
protuberant, the lower third contrastingly light
gray; thoracic pleuron moderate olive-brown except
lower half of mesokatepistemum light gray; male
mid tibia expanded. Male length 1.39-1.78 mm;
female 1.19-1.67 mm. Male terminalia as in
Figure 25.
HEAD.?Face centrally protuberant, the lower third
contrastingly light gray; 3-4 pfa; 4-7 sfa; epistomal
index 1.0-1.4; mesofacial index 1.3-2.0; vertex in-
dex 3.8-4.5; A-index 2.0-2.6; ocular index 4.8-8.0.
Palpus velvety black; 5-8 aristal rays; antenna and
most of parafrontale velvety black; fronto-orbital
area and frontal vitta moderate olive-brown; 12-15
postoculars.
THORAX.?Ppn and mesonotum moderate olive-
brown; 4-6 adc and 2-3 pdc; pleuron moderate
olive-brown except lower half of mesokatepistemum
light gray; legs moderate olive-brown pruinose except
dark-brown tarsi; male mid tibia expanded. Wing
length 1.36-1.79 mm; veins dark brown; 6-7 setae
on basal end of costa; 5-6 dorsal and 6-8 anterior
interfractural costals; costal-section ratios: I I : I 1.7-
2.3; I I I : IV 2.5-3.0; V:IV 2.8-3.2: M, *
 2 index
1.4-1.8.
ABDOMEN.?Terga densely moderate olive-brown
pruinose over dark gray. Male terminalia: median
third of posterior margin of sternum 5 straight;
anterolateral margin of sternum 5 rounded; copu-
lobus short, ovoid posteriorly, with about 7-10
scattered setae. Postgonite large, bent anteromediad,
and not covered by copulobus or surstyli; postgonite
uncus very conspicuous and nearly straight; disti-
phallus transversely expanded at midlength and with
triangular apex. Anterior margin of fused surstyli
deeply and broadly concave; B-index about 0.6;
C-index 0.9-1.1.
TYPE.?Holotype male, U S N M 70542.
TYPE-LOCALITY.?Two Island Lake, Itasca State
Park, Becker County, Minnesota (VI -28-1963 ,
D. L. Deonier) .
P A R A T Y P E S . ? I l l (24o* d \ 8 7 ? 9 ) from 19 locali-
ties:
Connecticut: Avon Old Farms, Avon (VI-27-1929, C.
Curran), 29 9 (AMNH).
Florida: DeFuniak Springs, Walton County (VI I I -1 -
1962, D. L. Deonier), 29 9 (USNM and ISC).
Idaho: Echo Bay, Lake Coeur d' Alene (VIII-3-1934,
A. Melander), 19 (ANSP); Worley (VI-29-1919, A.
Melander), \$ (ANSP).
Manitoba: Whitewater Lake, 4 miles north of White-
water (VI-22-1958, C. Miller), 19 (ISC).
Massachusetts: Mashbee (VII-28-1950, collector un-
known), 29 9 (USNM and ISC); Woods Hole (VIII-
no date-1916, A. Sturtevant), 19 (USNM).
Minnesota: Squaw Lake, Itasca State Park (VIII-14-
1963, D. L. Deonier), 3?J,J, 29 9 (USNM, ISC and
DLD); Two Island Lake, Itasca State Park (VI-28-1963,
D. L. Deonier), 1<$, 5 9 9 , (VIII-8-1963, D. L. Deonier),
2$ $, 69 9 (USNM, ISC, and DLD); 0.5 mile south of
junction U.S. 71 and 113, Hubbard County (VII-5-1963,
D. L. Deonier), 1 5 , 2 9 9 (DLD).
Mississippi: Gravel pit at Hattiesburg (VI-30-1962,
D. L. Deonier), \$, (VI-30-1962, J. Ainsworth), 119 9
(USNM, ISC, and DLD); Lake Shady, Lamar County
(VI-29-1962, D. L. Deonier), 1$, 59 9 (USNM and
ISC); Lake Shelby State Park, Forrest County, 31 ?9' N,
89?14.6' W (VII-24-1962, D. L. Deonier), 1$, 129 9
(USNM and ISC).
New York: Vicinity of Jockeybush Outlet, Hamilton
County, 43?18'N, 74?34' W (VIII-5-1961, D. L. Deonier),
19 (USNM).
Ontario: Marmora (VIII-7-1952, J. McAlpine), \$,
29 9 , (VIII-9-1952, J. McAlpine), 66$, 129 9 , ( IX-
8-1952, E. Smith), 19 , (IX-9-1952, J. McAlpine), 5<$ $,
69 9 (CNC, USNM, and ISC); Midland (VII1-18-1955,
J. Chillcott), 119 9 (CNC, USNM, and ISC); Pembroke
(VII-3-1938, A. Melander), 1 ?$ (ANSP).
Pennsylvania: Ohiopyle, Fayette County (VHI-no date-
1907, H. Kahl), 19 (ISC).
Yukon: Reflection Lake, mile post 1160 of Alaska High-
way (VII-29-1952, C. Alexander), 19 (UMM).
REMARKS.?For taxonomic remarks see under
H. columbata.
Adult. Habitats recorded: leaves of Potamogeton
natans, Nymphaea tuberosa, Nelumbo lutea, Hyd.ro-
cotyle bonariensis, and Sparganium chlorocarpum.
NUMBER 6 8 99
Hydrellia suspecta Cresson
Hydrellia suspecta Cresson, 1936, pp. 258-259; 1944b, pp.
169, 174.?Deonier, 1964, p. 117; 1965, pp. 500, 506
[ecology].?Wirth, 1965, p. 744 [catalog listing].
DIAGNOSIS.?Palpus moderate yellow; 6-8 aristal
rays; ocular index 15.0 (measured in 1 female only);
costal section II: I 2.3; 6 pfa; antennal segment 3
moderate yellow or splotched with moderate yellow;
fused surstyli moderate yellow and much longer
than broad. Female length 2.47 mm (measured in
1 female only).
HEAD.?Face yellowish gray; 6 pfa; epistomal
index 1.7; mesofacial index 2.2; vertex index 5.2;
A-index 1.8; ocular index 15.0. Palpus moderate
yellow; 6-8 aristal rays; antennal segments 1 and 2
dark brown, segment 3 moderate yellow or splotched
moderate yellow and dark brown; fronto-orbital area
and frontal vitta strong yellowish brown; most of
parafrontale dark brown, but flecked with strong
yellowish brown pruinosity; 18 postoculars.
THORAX.?Ppn and mesonotum moderate brown;
3 adc and 2 pdc; pleuron light gray; legs light-gray
pruinose except tibial apices and tarsi moderate
yellow. Female wing length 2.60 mm; veins pale
yellow; 6 setae on basal end of costa; 8 dorsal and
12 anterior interfractural costals; costal-section
ratios: II:I 2.3; III:IV 2.7; V:IV 3.6; Mx + 2 index
1.4.
ABDOMEN.?Female terga semiglossy moderate
brown medially, with posterolateral light-gray
wedges; these terga light gray laterally and ven-
trally. Male terminalia: unavailable for study, but
Cresson (1936, p. 259) stated: "Abdomen of male
ovate; tergites II to V subequal in length; V of male
narrowly truncated apically. Genital segment surstyli
of male exerted, flattened much longer than broad."
TYPE.?Holotype male, ANSP 6529.
TYPE-LOCALITY.?New Mill Pond, Mount Desert
Island, Maine (VII-12-1935, W. Procter).
SPECIMENS EXAMINED.?3? $ from 2 localities:
Iowa: Springbrook State Park, Guthrie County (VII-
19-1960, D. L. Deonier), 2 9 9.
Maine: Mount Desert Island (VII-12-1935, W. Procter),
19.
REMARKS.?I have studied the holotype, but care-
ful measurements could be made only on the female
designated by Cresson as the paratype. The male
holotype was unavailable for dissection of the
terminalia.
Adult. Habitats recorded: leaves of Nuphar
advena; reed marsh.
Hydrellia tibialis Cresson
FIGURES 41, 87, 118
Hydrellia tibialis Cresson, 1917, p. 341; 1918, pp. 47, 48;
1931, p. 106; 1932, p. 14; 1941, p. 37; 1944b, pp. 164,
172; 1947, pp. 38, 39.?Johnson, 1925, p. 272.?Wirth
and Stone, 1956, p. 469.?Deonier, 1964, p. 116; 1965,
pp. 500, 506 [ecology].?Wirth, 1965, p. 744 [catalog
listing]; 1968, p. 13 [catalog listing].
DIAGNOSIS.?Palpus dark brown; 5-7 aristal
rays; body length:wing length 0.7-0.9; face light
gray, with small median carina; mesonotum and
abdominal dorsum glossy dark grayish green; male
mid tibia expanded. Male length 1.28-1.62 mm;
female 1.82-2.21 mm. Male terminalia as in Figure
41.
HEAD.?Face light gray; 4-5 pfa; 1 sfa; epistomal
index 1.0-1.3; mesofacial index 1.7-2.1; vertex
index 3.5-7.0; A-index 1.6-2.0; ocular index 7.5-
8.0. Palpus dark brown; 5-7 aristal rays; antenna,
fronto-orbital area, and frontal vitta dark brown;
most of parafrontale velvety dark brown; 12-18
postoculars.
THORAX.?Ppn semiglossy dark brown; mesonotum
glossy dark grayish green; 4-5 adc and 2 pdc;
pleuron densely strong yellowish-brown pruinose;
legs very sparsely light-gray pruinose except dark-
brown tarsi. Wing length 1.82-2.35 mm; veins light
brown; 5-7 setae on basal end of costa; 6-8 dorsal
and 8-10 anterior interfractural costals; costal-
section ratios: I I : I 2.0-2.3; III:IV 2.8-3.2; V: IV
3.2-3.8; Mi ?
 2 index 1.7-2.1.
ABDOMEN.?Terga glossy dark grayish green
medially, light gray laterally and ventrally. Male
terminalia: median third of posterior margin of
sternum 5 concave and congruent with distiphallus;
anterolateral margin of sternum 5 smoothly rounded;
copulobus papilliform posteriorly (15? angular) and
irregularly setose. Postgonite bent anteriad; post-
gonite uncus directed anteriad or bent slightly
laterad; distiphallus short, tapering distally. Anterior
margin of fused surstyli truncate (indistinctly
mucronate) ; B-index about 1.6; C-index 0.8-1.2.
100 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
THIRD-INSTAR LARVA.?The frontoclypeus was lost
in the extraction of the only puparium. It was en-
tirely black like the mouth-hook (Figure 87). Mouth-
hook beak about 0.8 as long as base; maximum
mouth-hook base thickness about 1.6 times that of
beak.
PUPARIUM.?Length approximately 3.75 mm;
maximum breadth 0.95 mm; fusiform (Figure 118).
Puparial length: minimum breadth approximately
18.0; maximum breadth: minimum breadth 5.2; anal-
plate index 2.0. Prothorax and mesothorax dam-
aged, but apparently prothoracic end semicircular in
ventral view; abdominal segment 8 without ventral,
transverse row of setulae, but with 4-5 annuli of
spinules; anal plate subrectangular, with anterior
margin slightly convex; spiracular peritremes ter-
minal. Empty puparium translucent, light yellowish
brown.
TYPE.?Holotype male, ANSP 6141.
TYPE-LOCALITY.?Moscow, Idaho (X?9?1907, J.
M. Aldrich).
ADULT SPECIMENS EXAMINED.?840 (3180*0*,
522? $ ) from 148 localities:
Alabama: Unspecified locality, 12.
Alaska: King Salmon, Naknek River (VII-23-1952, W.
Mason), 23 3 , 82 2, (VII-31-1952, W. Mason), 22 2,
(VIII-1-1952, W. Mason), 2 2 2, (VIII-11-1952, W.
Mason), 1 2 ; Savonoski, Naknek Lake (VII-26-1919, A.
Basinger), 2 3 3 , 3 2 2.
Alberta: Grizzly Mountain, Slave Lake, 3,000 feet
(VIII-15-1924, O. Bryant), 12; McMurray (VII-14-
1953, G. Ball), 1 2 ; Medicine Hat (VI-16-1928, F. Carr),
It.
Arizona: Superior, Pinal County (IV-13-1935, A.
Melander), 12-
Arkansas: Calion, Union County (IX-3-1952, M.
Wheeler), 1 3 , 1 2 ; Rison, Cleveland County (IX-3-1952,
M. Wheeler), 12-
British Columbia: MacGillivray Creek Game Preserve
near Chilliwack (VII-14-1953, G. Spencer), 12 .
California: Big Pines (VIII-2-1944, A. Melander), 1 3 ;
Carpenteria (VIII-11-1950, A. Melander), 22 2 ; Cassel,
Shasta County (VII-15-1955, E. Schlinger), 22 2 ; Davis
(IV-17-1952, A. McClay), 1 3 , 12 , (VIII-13-1954, A.
Grigarick), 22 2 , (X-21-1954, A. Grigarick), 22 2 ,
(VII-27-1955, W. Lange), 12 ; El Capitan Reservation,
San Diego County (IX-25-1955, P. Arnaud), 42 2 ; Green
Valley (VII-26-1944, A. Melander), 1 2 ; Jenks Lake
(VII-14-1950, A. Melander), 22 2 , (VIII-18-1950, A.
Melander), 43 3 , 62 2 , (VIII-24-1950, A. Melander),
23 3 , 52 2 , (IX-7-1950, A. Melander), 202 2 ; Likeley,
Modoc County (X-l 1-1952, E. Schlinger), 1 3 , 62 2 ;
Manteca, San Joaquin County (IX-29-1902, J. Hesten),
1 2 ; Putah Canyon, Yolo County (XI-6-1954, W. Lange),
1 2 ; Rio Linda, Sacramento County (VIII-30-1957, A.
Grigarick), 1 3 ; San Diego (XI-18-1916, H. Dyar), 12 .
Colorado: Alamosa (VIII-1-no year, M. T. and G. M.
James), 1 2 ; Creede, 8,844 feet (VHI-no date-1914, S.
Hunter), 1 2 ; Creede (VII-1-1936, L. Tuthill), 12 ,
(VII-10-1939, L. Tuthill), 1 2 ; Deckers (VI11-25-1950,
collector unknown), 1 2 ; Electra Lake, about 37?33' N,
107 ?48' W, 8,400 feet (VI-28-VII-1-1919, collector un-
known), 23 3 ; Estes Park (VIII-11-1952, R. Dreisbach),
1 2 ; Grand Junction (IX-4-1938, D. Hardy), 1 3 , 32 2 ;
Tennessee Pass, 10,240 feet (VII-8-no year, J. Aldrich),
12 , (VII-24-1917, collector unknown), 1 2 ; Wray, about
40? N, 102?10' W (VIII-17-19-1919, collector unknown),
12 .
Delaware: Rehoboth (VI-25-1939, A. Melander), 1 2 ;
(VIII-4-1941, A. Melander), 13-
Florida: Baxter (IX-no date-1954, M. Wheeler), 1 2 ;
Fruitville (VIII-11-1930, R. Beamer), 12 .
Idaho: Moscow (X-9-1907, collector unknown), 1 2 ;
Soldier Creek, Priest Lake (VIII-22-1919, A. Melander),
1 3 , (VIII-22-1920, A. Melander), 1 2 ; Tule Bay, Priest
Lake (VIII-19-1919, A. Melander), 22 2 , (VIII-20-
1920, A. Melander), 12 , (VIII-22-1920, A. Melander),
22 2 .
Illinois: Champaign County (IV-26-1925, M. Shackle-
ford), 2 2 2 , (VII-10-1926, V. Smith), 1 2 ; Meredosia
(V-29-1917, collector unknown), 1 2 ; Peoria VIII-26-
1917, J. Aldrich), 1 2 ; Springfield (IX-7-1952, M.
Wheeler), 12 .
Indiana: Lafayette (X-22-1915, collector unknown), 13-
Iowa: Ames (IV-16-1927, collector unknown), 1 3 , 1 9 ,
(IV-24-1927, collector unknown), 12 , (V-12-1927, col-
lector unknown), 1 2 ; Ames, Skunk River (VII-17-1960,
D. L. Deonier), 1 2 ; Ames, Izaak Walton League Reserve,
Story County (IV-28-1960, D. L. Deonier), 1 2 , (VI-
17-1960, D. L. Deonier), 1 2 , (IV-23-1962, D. L.
Deonier), 1 3 , 22 2 ; Banner Mine Area, Warren County
(VIII-7-1960, D. L. Deonier), 62 3 3 , 29 2 2 , (V-23-
1963, D. L. Deonier), 1 3 , 1 2 ; Fraser Dam, Boone County
(IX-4-1960, D. L. Deonier), 1 2 , (VII-6-1961, D. L.
Deonier), 1 3 , 1 2 ; Lake Odessa, Louisa County, 41? 12.1'
N, 91?5.5' W (VIII-9-1960, D. L. Deonier), 123 3 ,
162 2 ; Ledges State Park, Boone County (V-7-1956, J.
Laffoon), 13 , (V-6-1958, J. Laffoon), 1 3 , (VI-14-1960,
J. Laffoon), 12 , (VIII-16-1960, D. L. Deonier), 1 2 ,
(VI11-18-1960, D. L. Deonier), 1 3 , 22 2 ; Ledges State
Park, Des Moines River (IX-12-1959, D. L. Deonier),
22 2 , (VIII-31-1960, D. L. Deonier), 3 3 3 , 52 2 ;
Ledges State Park, south pond (VII-10-1960, D. L.
Deonier), 1 2 ; Little Wall Lake, Hamilton County (VII-
13-1960, D. L. Deonier), 1 3 , 3 2 2 , (VIII-27-1960, D. L.
Deonier), 33 3 , 32 2 , (VI-18-1961, D. L. Deonier),
4 3 3 , 2 2 2 ; Mclntosh Woods State Park, Clear Lake
(IX-10-1961, D. L. Deonier), 4 3 3 , 32 2 ; Mississippi
River, 4 miles north of Oakville, Louisa County (VIII-10-
1960, D. L. Deonier), 1 2 ; near Missouri River, 7 miles
southwest of Whiting, Monona County (VI-6-1960, D. L.
Deonier), 1 3 ; Pilot Knob State Park, Hancock County
(VI-15-1960, D. L. Deonier), 1 2 ; Siewer's Springs State
NUMBER 6 8 101
Park, Decorah (IX-9-1961, D. L. Deonier), 2$ 3 , 7 9 9,
(IX-15-1962, D. L. Deonier), 4 3 3 , 39 9 ; Sioux City
(VI-20-1938, C. Ainslie), 1 2 ; Spring Lake, Greene County
(V-4-1962, D. L. Deonier), 6 3 3 , 1 4 9 9 , (IX-19-1962,
D. L. Deonier), 2 3 3 , 1 9 ; Springbrook State Park,
Guthrie County (VII-19-1960, D. L. Deonier), 1 3 , (VI-
23-1961, D. L. Deonier), 1 3 ; West Okoboji Lake, Dickin-
son County (IX-3-1949, J. Laffoon), 1 9 ; 4 miles east
of Gilbert, Story County (VI-25-1960, D. L. Deonier),
1 3 ; 3 miles east-southeast of Waterville, Allamakee County,
43?11' N, 91?14.1' W (VIII-2-1960, D. L. Deonier),
6 3 3 , 119 9 ; 3 miles southeast of Waterville, Allamakee
County, 43? 10.4' N, 91? 15.1' W (VIII-2-1960, D. L.
Deonier), 5 3 $, 99 9 , (VIII-2-1960, J. Laffoon), 2$ 3 ,
3 9 9 ; 4 miles northeast of Wapello, Louisa County, 41? 13.9'
N, 91?7.0' W (VIII-9-1960, D. L. Deonier), 63 3 , 8 9 9 ;
5 miles west of Yale, Guthrie County, 41?46.5' N, 94?27.6'
W (VII-19-1960, J. Laffoon), 13-
Kansas: Douglas County, 900 feet (no dates, R. Beamer),
13 ; Kansas University Natural History Reservation near
Lawrence, Douglas County (VI-7-1963, D. L. Deonier),
13 ; Leavenworth County Lake (VIII-27-1962, D. L.
Deonier), 123 3 , 199 9 ; Manhattan (X-18-1934, C.
Sabrosky), 1 3 ; Marais des Cygnes Wildlife Refuge, Linn
County (IV-9-1962, D. L. Deonier), 29 9 ; McPherson
County (VI-26-1923, W. Brown), 19 ; Meade County
(VIII-15?1915, R. Beamer), 1 9 ; Sappa Lake, Decatur
County (VIII-31-1961, D. L. Deonier), 4 3 3 , 109 9 ;
1.5 miles south of Bonner Springs (VI11-28-1962, D. L.
Deonier), 1 9 .
Louisiana: 15 miles east of Creole (VI-18-1948, B.
McDermott), 1 3-
Maine: Mount Desert Island (VII-25-1935, W. Procter),
19 .
Manitoba: Churchill (VIII-2-1937, D. Denning), 1 3 ,
(VIII-2-9-1937, D. Denning), 1 9 ; Herchmer ( V I I I - 1 -
1937, D. Denning), 1 3 , 29 9 ; Whitewater Lake, 4 miles
north of Whitewater (VII-17-1958, J. Chillcott), 1 9 .
Maryland: Plummer's Island, Potomac River ( IV-28-
1914, R. Shannon), 1 9 ; unspecified locality (no dates, H.
Loew), 1 3 .
Michigan: Baraga County (VIII-27-1952, R. Dreis-
bach), 1 9 ; Bay County (VIII-11-1951, R. Dreisbach),
1 9 ; Cass County (VIII-2-1953, R. Dreisbach), 1 3 , 1 9 ;
Cheboygan County (VII-26-1935, L. Penner), 1 9 ; Vine-
yard Lake, Jackson County (VII-26-1954, G. Steyskal),
2 3 3 , 99 9 , (VIII-1-1954, G. Steyskal), 13-
Minnesota: Bellingham (IX-11-1935, H. Telford), 1 9 ;
Chisago County (no dates, O. Oestlund), 29 9 ; Cook
County (VIII-21-1938, H. Milliron), 39 9 , (VII I -21-
1938, H. Peters), 1 9 ; Eaglenest (VIII-26-1959, W.
Balduf), 1 3 ; Freeborn County (IX-3-1937, R. Daggy),
1 3 , 1 9 ; Haydenville (IX-11-1935, A. Pritchard), 1 3 ,
2 9 9 ; Headwaters of LaSalle Creek, Itasca State Park
(VIII-6-1963, D. L. Deonier), 1 3 , 29 9 ; Itasca State
Park (VI-20-1938, H. Milliron), 1 9 ; Mille Lacs (VI-2-
1937, D. Denning), 1 9 ; St. Paul (V-25-1926, S.
Kepperley), 19 ; Washington County (V-7-1938, H. Mill-
iron), 1 9 ; west side across from Biological Station, Lake
Itasca (VII-28-1963, D. L. Deonier), 1 9 ; Yellow Medicine
County (IX-15-1938, C. Mickel), 29 9 ; 1 mile south of
main entrance, Itasca State Park (VI-26-1963, D. L.
Deonier), 1 3-
Mississippi: Bellefontaine Point, Jackson County (VII-
10-1962, D. L. Deonier), 43 3 , 99 9 ; Bluff Creek,
Vancleave, Jackson County (VI-18-1962, D. L. Deonier),
2 9 9 , (VI-21-1962, D. L. Deonier), 1 3 , 1 9 ; Dickinson's
Pond, Lamar County, sec. 3, T.4 north, R.I4 west (VII-
11-1962, D. L. Deonier), 5 3 3 , 6 9 9 ; near U.S. Highway
90, sec. 20, T.7 south, R.5 west, gravel pit (VI-9-1962,
D. L. Deonier), 1 3 , 69 9 , (VI-17-1962, D. L. Deonier),
2 3 3 , 49 9 ; Vancleave Road, Jackson County, 30?28' N,
88?43' W (VI-21-1962, D. L. Deonier), 2 9 9 , (VI-28-
1962, D. L. Deonier), 1 3 ; Vancleave Road, Jackson
County, 3O?25.1' N, 88?46' W (VI-23-1962, D. L.
Deonier), 1 9.
Missouri: Four miles northeast of LaRussell, Lawrence
County (IX-6-1961, D. L. Deonier), 9 3 3 , 159 9 ; 5
miles northeast of LaRussell, Lawrence County (IX-6-
1961, D. L. Deonier), 13-
Nebraska: Hastings (VIII-22-1950, collector unknown),
1 9 ; Lincoln, Lancaster County (IX-30-1952, W. Stevens),
19 , (VII-16-1960, W. Rapp), 1 3 ; Oakdale (VIII -21-
1950, collector unknown), 19 .
New Hampshire: Storm Lake, Mount Adams, 5,200 feet
(VIII-29-1943, M. Smith), 29 9 ; Summit of Mount
Washington, 6,100-6,280 feet (VIII-14-1958, J. Vocke-
roth), 2 9 9 .
New Jersey: Trenton (VII-11-1914, collector unknown),
19 .
New York: Dryden Lake, Tompkins County (VIII-8-
1961, D. L. Deonier), 3 3 3 , 2 9 9 .
North Carolina: Clingman's Dome, Great Smoky Moun-
tains National Park, 6,300-6,642 feet (V-28-1957, J.
Vockeroth), 13-
Oklahoma: Kingfisher (IX-13-1952, M. Wheeler), 13-
Ontario: Marmora (VIII-12-1952, J. McAlpine), 19 .
Oregon: Corvallis (VII-1-1925, collector unknown),
19 , (V-16-1941, J. Schuh), 6 3 3 , 29 9 ; North Powder
(VII-13-1931, R. Beamer), 1 9 .
Pennsylvania: Ohiopyle, Fayette County (VIII-7-1905,
H. Kahl), 1 3 , 29 9 , (VIII-11-1905, H. Kahl), 33 3 ,
19 ; Swarthmore (VI11-22-1909, E. Cresson), 1 9 .
Quebec: Gaspe (VIII-9-1937, C. Alexander), 1 9 ;
Gaspe Bay (VII-18-1931, J. Aldrich), 1 9 ; Great Valley,
Gaspe (VII-20-1931, J. Aldrich), 3 9 9 ; La Ferme (VII -
no date-1943, A. Roberts), 1 9 ; Notre Dame du Portage
(VIII-17-1957, W. Mason), 1 9 ; Rupert House (VII-25-
1949, D. Gray), 1 9 .
Tennessee: Dale Hollow Reservoir, 6 miles south of Byrds-
town, Pickett County (VIII-19-1962, D. L. Deonier),
443 3 , 679 9 ; Miller's Camp, Reelfoot Lake, 36?24.3'
N, 89?20' W (VIII-12-1962, D. L. Deonier), 1 9 ; Reel-
foot State Park, Reelfoot Lake (VIII-16-1962, D. L.
Deonier), 333 3 , 389 9 ; Samburg, Reelfoot Lake (VIII-
11-1962, D. L. Deonier), 2 3 3 , (VIII-12-1962, D. L.
Deonier), 4 9 9-
102 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Texas: Austin (III-2-1900, A. Melander), 2 9 9 , (X-
19-1950, M. Wheeler), 1 3 , (XI-19-1950, collector un-
known), 1 9 , (XII-1-1951, M. Wheeler), 1 9 ; Double
Lake (IV-11-1952, collector unknown), 1 $ ; Galveston
(IX-13-1953, M. Wheeler), 1 9 ; Goliad (IX-5-no year,
Townsend), 1 9 ; Guadalupe River, Gonzales (IV-22-1956,
W. Wirth), 2$ 2; Rio Hondo, Cameron County (VIII -
2-1950, M. Wheeler), 2$ $, 3 9 9 ; San Antonio (111-28-
1942, A. Melander), 29 9 ; Sinton (XII-25-1945, R. Bea-
mer), 1 $.
Virginia: Fairfax (VH-no date-1954, M. Wheeler), 1 $ ;
Norfolk (IX-4-1943, R. Beamer), 1 $ ; Warsaw, Richmond
County (VII-26-1952, W. Wirth), 1 9 .
Washington: Union Flat (IX-8-1918, collector un-
known), 1 S ?
Wyoming: Lander (VIII-16-1950, M. Wheeler), 1 9 ;
Yellowstone National Park, Biscuit Basin (VIII-2-1934, A.
Melander), 49 9 ; Yellowstone National Park, northwest
entrance (VII-27-1923, A. Melander), 1 9 ; Yellowstone
National Park, Yellowstone Lake (VII-18-1923, A. Me-
lander), 2 9 9 .
IMMATURE SPECIMENS EXAMINED.?1 from Banner
Mine Area, Warren County, Iowa (collected and
reared by D. L. Deonier).
REMARKS.?This species is relatively readily recog-
nized by the light-gray, slightly carinate face and
metallic appearance of the mesonotum and abdomen;
however, female H. tibialis may be confused with
females of H. americana, H. definita, and H. prudens.
Adult. Habitats recorded: stems of Eleocharis
calva and Eleocharis sp.; leaves of Sagittaria lati-
folia, Potamogeton nodosus, Nasturtium officinale,
Sporobolus indicus, Leersia oryzoides, Hydrochloa
caroliniensis, Heteranthera dubia, Glyceria sp.,
Juncus repens, Nuphar advena, Nymphaea tuberosa,
Zizania aquatica, Oryza sativa, Pontederia cordata,
and Polygonum scabrum; mats of Eragrostis hyp-
noides; flowers of Ranunculus aquatilis; limnic
wrack, sandbars, rocky shore, riffle rocks, and aquatic
liverwort (Riccia fluitans).
I found this species to be nearly as eurytopic and
abundant as H. griseola. It was one of the most
characteristic dipterous species in my samplings of
the sedge meadow community. At Banner Mine Area,
Warren County, Iowa, 50 net sweeps yielded 212
adults from a clump of Eleocharis calva covering
about a square meter. Also, I found adults very
abundant in mats of Eragrostis hypnoides in central
Iowa. My surveys revealed a greater preference for
emergent vegetation than for floating leaves as indi-
cated by data from Grigarick floating traps set among
floating leaves of P. gramineus and Jussiaea repens
on Reelfoot Lake. Four traps set for a total of 5 days
yielded only 14 adults.
Adults collected on flowers of R. aquatilis were
evidently feeding on nectar or pollen. I examined
the gut contents of 75 adults and found greenish-
yellow, granular material with some diatoms.
Host plants. I reared two adults from Eleocharis
obtusa collected at Banner Mine Area, Warren
County, Iowa, in April. Either the larvae had over-
wintered in the plants or had recently hatched, for
the adults did not emerge until 28 days after collec-
tion of the plants. The larvae pupariated in the
basal sheaths of the culms. Judd (1962) trapped
four adults in June and July in an emergence trap
floating where the depth ranged from 0.6 to 1.4
meters.
Hydrellia trichaeta Cresson
FIGURES 37, 68, 88, 115
Hydrellia trichaeta Cresson, 1944a, p. 7; 1944b [in part],
pp. 170, 175.?Deonier, 1964, pp. 115, 125, fig. 4.
Hydrellia coniformis.?Johnson [in part], 1925, p. 271
[lapsus calami, fide Cresson, 1944b, p. 135].
Hydrellia conformis.?Cresson, 1936, p. 261.
DIAGNOSIS.?Palpus moderate yellow; 8-9 aristal
rays; face planate; vertex index 5.4-6.8; ocular index
6.1-8.6; male abdomen rounded posteriorly; female
cercus mucronate apically and truncate basally;
antennal segment 3 partly moderate yellow; apico-
dorsal antennal prominent. Male length 1.82-2.13
mm; female 1.71-2.21 mm. Male terminalia as in
Figure 37; female as in Figure 68.
HEAD.?Differing mainly from H. bilobifera in the
following: ocular index 6.1-8.6; 8-9 aristal rays.
THORAX.?Differing mainly from H. bilobifera in
the following: 3-4 adc and 2 pdc; wing length
1.67-2.04 mm; veins dark brown; 6-8 dorsal and
8-10 anterior interfractural costals.
ABDOMEN.?Terga semiglossy dark gray medially,
light gray laterally and ventrally (at least on pos-
terior half of each segment). Male terminalia:
median third of posterior margin of sternum 5 deeply,
rectangularly recessed and congruent with disti-
phallus; anterolateral margin of sternum 5 with
inconspicuous dorsal process projecting laterad;
copulobus slightly convex posteriorly, without macro-
chaetae, the medial margin with convexity just
anteriad of postgonite uncus having seriated setae,
NUMBER 6 8 103
and with lateral triangular lobe. Postgonite bent
mediad; acuminate postgonite uncus curved anteriad;
distiphallus constricted proximally at midlength and
preapically, with ventrally lamellate apex, and
bicarinate ventrally on proximal two-thirds (carinae
connected preapically) ; basiphallus covered. Anterior
margin of fused surstyli concave medially with mod-
erately prominent lateral convexities; B-index about
2.0; C-index 0.6-0.8. Syntergum 9+10 rounded pos-
teriorly. Female terminalia: length of tergum 8 and
sternum 8 subequal; cercus irregularly setose dorsally
and laterally, mucronate apically, truncate basally
and less than 1.5 times as long as wide (lateral view).
Segment 7 with group of 3-4 seriated setae laterally
on posterior margin; median spermatheca similar to
that of H. discursa (Figure 70).
THIRD-INSTAR LARVA.?Body of larva not examined.
Frontoclypeal length 0.50 mm (Figure 88). Ventral
frontoclypeal index 2.7; phragmatal index 1.0;
bifurcation index 3.6; clypeal-arch index 2.0. Clypeal
arch slightly concave from noticeable convexity at
the level of the cheliform spot (angled 25? in relation
to lower frontoclypeal margin). Mouth-hook beak
length only 0.7-0.8 of base length; maximum mouth-
hook base thickness about 2.2 times that of beak;
mouth-hook light spot touching margin of mouth-
hook base.
PUPARIUM.?Length 3.70 mm; maximum breadth
0.76 mm; fusiform (Figure 115). Puparial length:
minimum breadth 20.0; maximum breadth: minimum
breadth 4.2; anal-plate index 2.6. Prothoracic end
describing more than half a circle in ventral view;
head-lobe scar ovoid to obovoid, nearly as long as
prothorax; prothorax and mesothorax sparsely spinu-
lose ventrally and laterally; abdominal segment 8
with ventral, transverse row of 6 setulae and 3-4
annuli of spinules; anal plate subelliptical, with
anterior margin straight to slightly convex. Empty
puparium translucent, light yellowish brown.
TYPE.?Holotype male, ANSP 816.
TYPE-LOCALITY.?Redding, Connecticut (VIII-
3-1931, A. L. Melander).
ADULT SPECIMENS EXAMINED.?98 (44cfcf,
54 ? $ ) from 45 localities:
Iowa: Goose Lake, Hamilton County (VII-2-1961, D. L.
Deonier), 49 9 : Mclntosh Woods State Park, Clear Lake
(IX-10-1961, D. L. Deonier), 3 3 3 , 49 9 ; Spring Lake,
Greene County (VII-14-1961, D. L. Deonier), 4 3 3 , 7 9 9 ,
(IX-19-1961, D. L. Deonier), 63 4 , 39 9 , (V-4-1962,
D. L. Deonier) 1 9 ; Springbrook State Park, Guthrie
County (VI-23-1961, D. L. Deonier), 1 4 , 19.
Kansas: Kansas University Natural History Reservation
near Lawrence, Douglas County (VI-7-1963, D. L.
Deonier), 13-
Massachusetts: Horseneck Beach (VIII-10-1896,
Hough), 19 .
Minnesota: Biological Station, Lake Itasca (VI-26-
1963, D. L. Deonier), 1 9 ; Bohall Lake, Itasca State Park
(VI-25-1963, D. L. Deonier), 63 3 , 39 9 , (VIII-17-
1963, D. L. Deonier), 10 3 3 , 49 9 ; Squaw Lake, Itasca
State Park (VIII-14-1963, D. L. Deonier), 19 ; 1.3 miles
south of main entrance, Itasca State Park (VI-26-1963,
D. L. Deonier), 3 3 5 , 1 9 .
New York: Dryden Lake, Tompkins County (VIII-8-
1961, D.L. Deonier), 19.
North Carolina: Highlands Biological Station, High-
lands, Macon County (VI-15-26-1968, D. L. Deonier),
83 3 , 2 0 9 9-
Ontario: Marmora (VII-25-1952, E. Smith), 13-
Tennessee: Miller's Camp, Reelfoot Lake, 36?24.3' N,
89?20' W (VIII-12-1962, D. L. Deonier), 13 , 19.
Virginia: White Oak Canyon, south slope of Stony Man,
3,500 foot, Madison County (VII-22-1961, D. L. Deonier),
19 .
IMMATURE SPECIMENS EXAMINED.?23 from 3
localities:
Iowa: Spring Lake, Greene County (collected and reared
by D. L. Deonier).
Minnesota: Bohall Lake, Itasca State Park (collected and
reared by D. L. Deonier).
North Carolina: Highlands Biological Station, Highlands,
Macon County (collected and reared by D. L. Deonier).
REMARKS.?Adult. Habitats recorded: leaves of
Potamogeton natans, P. nodosus, P. gramineus, and
Nuphar advena.
One male and two females reared in the laboratory
and caged on 18 July copulated. The females de-
posited 11 eggs on floating leaves of P. nodosus on
20 July. The specimens died on 21 July.
Egg and larva. The incubation period for three
eggs lasted 4 days. The first larval stadium ranged
from 4?7 days for two larvae. The first and second
larval stadia combined amounted to 13 days for two
larvae. The second and third larval stadia ranged
from 5 to 11 and 4 to 14 days respectively for five spe-
cimens. I found larvae of Dytiscidae to be efficient
predators on H. trichaeta larvae in the laboratory.
I made the following observations on larvae from
P. nodosus collected from ice at Spring Lake, Greene
County, Iowa, on 25 January. One third-instar larva
migrated from its first host plant to a leaf of P.
berchtoldii placed near it and pupariated in it. On
104 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
24 February, I found a late first-instar larva mining
in a leaf of P. nodosus and placed a fresh leaf of
P. berchtoldii near it. The larva molted on 26 Feb-
ruary and crawled around the bottom of the dish
until I placed it on the fresh leaf. It started exca-
vating a mine at 2:10 p.m. and had burrowed in
entirely by 3:35 p.m. During this period, the larva
did not pierce the water surface with its spiracular
peritremes. After it had burrowed in entirely, a small
bubble formed at the mine entrance. This larva
mined out most of the mesophyll of two leaves of
P. berchtoldii and part of that of a submerged leaf
of P. amplifolius before pupariating in the apex of a
second submerged leaf of P. amplifolius. As a third
instar, the larva burrowed completely into a leaf in
1 hour. Another larva starting as a first instar mined
throughout a submerged leaf of P. berchtoldii and
partially in one of P. amplifolius before pupariating.
As a second instar, this larva required 2 hours to
burrow into a leaf. In one observation of it as a
third instar, it defecated explosively, with the green
feces so particulate that it diffused rapidly in water
within the old mine. Shortly after this occasion, the
larva pierced the water surface film with its
spiracular peritremes three times in about 3 minutes.
Puparium. The puparial phase ranged from 8 to
10 days for six puparia in the laboratory. All of these
six pupariated in leaf axils or on the abaxial side of
floating leaves.
Host plants. I found larvae and puparia only in
Potamogeton gramineus, P. epihydrus, P. nodosus,
and Elodea = Anacharis) canadensis.
Hydrellia valida Loew
FIGURE 22
Hydrellia valida Loew, 1862, p. 153.?Osten Sacken, 1878,
p. 202.?Becker, 1896, p. 269 [index listing].?Johnson,
1904, p. 163; 1925, pp. 271-272.?Aldrich, 1905, p. 627
[catalog listing].?Jones, 1906, p. 185 [catalog listing].?
Cresson, 1944a, p. 7; 1944b, pp. 167, 173.?Wirth, 1965,
p. 744 [catalog listing].
DIAGNOSIS.?Palpus moderate yellow; 5-7 aristal
rays; vertex index 6.0-7.5; 12-15 anterior inter-
fractural costals; mesonotum light gray; thoracic
pleuron light gray; abdomen light gray. Male length
2.21-2.50 mm; female 2.38-2.81 mm. Male termi-
nalia as in Figure 22.
HEAD.?Face light yellowish brown; 4-6 pfa; 1-2
sfa; epistomal index 1.0-1.2; mesofacial index 2.0-
2.2; vertex index 6.0-7.5; A-index 1.5-2.0; ocular
index 6.3-6.7. Palpus moderate yellow; 5-7 aristal
rays; antenna and most of parafrontale dark brown;
frontal vitta and fronto-orbital area strong yellowish
brown; 12-15 postoculars.
THORAX.?Ppn light gray; mesonotum light gray
(infrequently with sparse light yellowish-brown
pruinosity) ; 3-4 adc and 2-3 pdc; pleuron light
gray except upper edge of mesanepisternum in-
frequently light yellowish brown; legs light-gray
pruinose except dark-brown tarsi. Wing length 2.47-
3.01 mm; veins light yellowish brown; 6-8 setae on
basal end of costa; 8-12 dorsal and 12-15 anterior
interfractural costals; costal-section ratios: I I : I 2.2-
2.8; I I I : IV 2.3-2.5; V:IV 3.4-4.4; M, ?
 2 index
1.1-1.5.
ABDOMEN.?Terga and sterna light gray. Male
terminalia: median third of posterior margin of
sternum 5 concave; anterolateral margin of sternum
5 obtusely rounded; copulobus nearly truncate
(occasionally slightly convex) posteriorly and ir-
regularly setose except on posterior margin.
Postgonite bent anterolaterad; postgonite uncus
short, bent laterad; distiphallus uniform in breadth;
basiphallus with black, spinous lateral process.
Anterior margin of fused surstyli broadly emarginate;
B-index about 2.7; C-index 8.0-10.0.
TYPE.?Holotype female, MCZ 11156.
TYPE-LOCALITY.?"Middle States." (Loew, 1862,
stated that Osten Sacken was the collector.)
SPECIMENS EXAMINED.?86 (22o*cf, 6 4 $ ? )
from 24 localities:
Connecticut: Goose Island (VII-21-1913, collector un-
known), 1 $ ; Greenwich (VI-21-1916, E. Kalmbach), 1 $ ;
New Haven, west shore of harbor (VII-29-1961, D. L.
Deonier), 3$$, 1 8 $ $ , (VII-29-1961, J. Laffoon), \$;
Stony Creek (VII-27-1904, H. Viereck), 1 $ ; Westport
(VII-13-1932, A. Melander), 4$ $.
Delaware: Rehoboth (VI-25-1939, A. Melander), 19 .
Florida: Unspecified locality (no data), 1,5.
Maine: Machias (VII-17-no year, collector unknown),
IS, 1 $ ; Orrs Island (VII-26-no year, collector unknown),
2 $ $ ; Trenton (VIII-1-1930, A. Melander), 1 $ .
Massachusetts: Brewster (VIII-24-1936, A. Melander),
1 $ ; Fall River (VII-3-1930, A. Melander), 2 $ $ ; Horse-
neck Beach (VIII-6-1896, Hough), 1 $ ; Ipswich (VII-22-
1961, W. Wirth), 1<$, 1 $ ; New Bedford (VIII-20-1896,
Hough), 2$ $ ; Woods Hole (VII-14-1899, W. Wheeler),
6 $ $ , (VI-20-30-1923, collector unknown), 1$, (VII-
21-1954, M. Wheeler), 5$ $, 8 $ $ ; 3 specified localities
(no data), 2$ <$, 1 $ .
NUMBER 6 8 105
Mississippi: Camp Graveline, Jackson County, 30?22.3'
N. 88?42.3' W (VI-12-1962 , D. L. Deonier), 1 $ ; Gulf
Coast Research Laboratory, Ocean Springs (VI-5 -1962 ,
D. L. Deonier), 1 $ .
New Jersey: Waterwitch (VI-5-1937 , A. Melander),
3 $ 9 .
New York: Cold Spring Harbor, Long Island ( V U - n o
other data), 4SS, 2 9 9 , (VII-6-1927 , A. Melander),
1$, 19 (VII-26-1932, C. Curran), 1 9 .
Nova Scotia: Lockeport (VII-18-1958, J. Vockeroth),
1 9 ; Smith's Cove (VI-24-1955, G. Steyskal), 1 9 .
Rhode Island: Watch Hill (VIII -5-1939 , A. Melander),
1 9 ; unspecified locality (no dates, Osten Sacken), 1$.
Texas: Galveston ( IX-13-1953 , M. Wheeler), 1 9 .
REMARKS.?Hydrellia valida is a sibling species of
H. griseola, but it has some differential characters
other than the male terminalia.
Adult. Habitats recorded: tidal marsh and moist
intertidal sand.
I examined the following plant species for imma-
tures of H. valida: Cyperus sp., Scirpus olneyi,
Juncus sp., Spartina patens, Spartina sp., and
Sporobolus indicus. All were negative.
Hydrellia ivilburi Cresson
FIGURE 38
Hydrellia wilburi Cresson, 1944b, p. 168, 172.?Wirth,
1965, p. 744 [catalog listing].
DIAGNOSIS.?Palpus dark brown; 8-11 aristal rays;
mid and hind tarsi moderate orange-yellow; face
white; thoracic pleuron moderate brown; 9-11 setae
on basal end of costa; male abdomen compressed.
Male length 2.30-2.42 mm; female 2.38-2.72 mm.
Male terminalia as in Figure 38.
HEAD.?Face white; 3-5 pfa; 4-11 sfa; epistomal
index 1.0-1.4; mesofacial index 1.6-2.2; vertex
index 4.2-7.0; A-index 1.7-2.0; ocular index 4.5?
6.0. Palpus dark brown; 8-11 aristal rays; apico-
dorsal antennal prominent; antennal segments 2 and
3 and frons (except light-brown ocellar triangle
and dark-brown fronto-orbital area) velvety black;
antennal segment 1 dark brown; 13-16 postoculars.
THORAX.?Ppn and mesonotum semiglossy dark
brown; 4-5 adc and 2-3 pdc; pleuron semiglossy
moderate brown; legs dark brown except dark-yellow
trochanters and moderate orange-yellow mid and
hind tarsi. Wing length 2.70-3.32 mm; veins light
yellowish brown; 9-11 setae on basal end of costa;
6-8 dorsal and 9-11 anterior interfractural costals;
costal-section ratios: I I : I 2.2-2.5; I I I : IV 2.5-3.0;
V: IV 3.0-3.8; Mi ?
 2 index 1.3-1.8.
ABDOMEN.?Terga and sterna dark brown. Male
terminalia: see the description for H. platygastra.
TYPE.?Holotype male, ANSP 6657.
TYPE-LOCALITY.?Bear Lake, 8,500 feet, Routt
County, Colorado (VIII-19-1935, D. A. Wilbur).
SPECIMENS EXAMINED.?71 (35o* cf, 36$ $ ) from
9 localities:
British Columbia: Shuswap Lake (VII-22-1926, J. Mc-
Dunnough), 1<5, 1 9 .
California: Glacier Point Road, Yosemite National Park
(VII-6-1947, A. Melander), 1 9 ; Sequoia National Park
(VIII-6-1940, D. Hardy) , 1 4 , 2 9 9 .
Colorado: Bear Lake, Routt County, 8,500 feet ( V I I I -
19-1935, D. Wilbur), 1 $ ; Lake Brennan, 10,000 feet ( V I I -
15-1934, C. Alexander), 31 $ <}, 2 7 9 9 ; Nederland, Boul-
der County (VII-10-1937, H. Peters), 19 ; Tennessee Pass
(VII-25-1917, J. Aldrich), 1 9 .
Wyoming: Kemmerer (VIII-14-1950, M. Wheeler), 1$,
3 9 9 .
REMARKS.?The male terminalia of H. wilburi
and H. platygastra appear to be identical. The two
species may be conspecific, but since they are sym-
patric and no evidence of intergradation in the facial
color is apparent I am treating them as separate
species. See additional remarks under H. platygastra.
106 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 1.?Checklist of known host plants of Hydrellia.
Species of
Hydrellia
albiceps
albifrons
albilabris
ascita
bergi
bilobifera
biloxiae
butomi
Host
plant
Alismataceae
Alisma
plantago-aquatica
Cruciferae
Nasturtium
officinale
Alismataceae
Alisma
plantago-aquatica
Lemnaceae
Lemna
minor
Spirodela
Potamogetonaceae
Potamogeton
alpinus
amplifolius
epihydrus
foliosus
illinoensis
oakesianus
richardsonii
zosteriformis
Potamogetonaceae
Potamogeton
natans
richardsonii
zosteriformis
Potamogetonaceae
Potamogeton
nodosus*
berchtoldii*
gramineus*
epihydrus*
Najadaceae
Zannichellia
palustris^
Juncaceae
Juncus
repens*
Butomaceae
Butomus
Gramineae
Zoogeographic Major
region
Palaearctic
Palae arctic
Palaearctic
Palaearctic
Nearctic
Nearctic
Nearctic
Nearctic
Nearctic
Palaearctic
Palaearctic
references
Hendel (1926), Hering (1935, 1957)
Seguy (1950)
Marchal (1903), Hering (1925,
1926), Thieneman (1912)
von Frauenfeld (1866), Linnaniemi
(1913), Hendel (1926), Hering
(1925, 1926, 1937, 1957), Seguy
(1950)
Berg (1949, 1950)
Berg (1949, 1950)
Hendel (1926), Hering (1937, 1951,
1957)
Hering (1951)
NOTE: ? Reared from host for the first time during this study, t Recorded first in this
study. % Reared from host during this study.
NUMBER 6 8 107
TABLE 1.?Checklist of known host plants of Hydrellia.?Continued
Species of
Hydrellia
caliginosa
chrysostoma
cochleariae
concolor
cruralis
deceptor
discursa
fascitibia
flaveola
Host
plant
Potamogetonaceae
Potamogeton
praelongus
Alismataceae
Alisma
plantago-aquatica
Potamogetonaceae
Potamogeton
lucens
Callitrichaceae
Callitriche
Hydrocharitaceae
Hydrocharis
morsus-ranae
Potamogetonaceae
Potamogeton
crispus
perfoliatus
Alismataceae
Alisma
plantago-aquatica
Hydrocharitaceae
Stratiotes
aloides
Potamogetonaceae
Potamogeton
alpinus
amplifoliust
epihydrus
foliosus
gramineus
illinoensis
natans
nodosus
praelongusX
richardsoniit
zosteriformis
Alismataceae
S a git t aria
sp.t
Potamogetonaceae
Potamogeton
gramineus*
Potamogetonaceae
Potamogeton
Geraniaceae
Tropaeolum
minus
majus
Zoogeographic
region
Nearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Nearctic
Nearctic
Nearctic
Palaearctic
Palaearctic
Major
references
Berg (1949, 1950)
Hendel (1926), Grunberg (1910),
Hering (1937, 1957)
Hering (1925, 1937, 1957)
Hering (1957)
Seguy (1950)
Hering (1950, 1951, 1957), Seguy
(1950)
Hering (1924, 1925, 1926, 1937,
1957)
Hering (1924, 1925, 1957), Wahlgren
(1947)
Berg (1949, 1950)
Hering (1937, 1950, 1951)
Frost (19?4) [as Notiphila flaveola],
Marchal (1903)
108 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 1.?Checklist of known host plants of Hydrellia.?Continued
Species of
Hydrellia
Host
plant
Zoogeographic
region
Major
references
flaveola
(cont'd)
flavicornis
fulviceps
gladiator
glycerine
griseola
Papaveraceae
Papaver
sp.
Alismataceae
Alisma
plantago-aquatica
Alismataceae
Alisma
plantago-aquatica
Hydrocharitaceae
Vallisneria
americana\
Gramineae
Glycerin
aquatica
Gramineae
Agropyron
Agrostis
Alopecurus
Anthoxanthum
odoratum
Apera
Avena
sativa
Briza
Bromus
Brachypodium
Calamagrostis
Catabrosa
Cynodon
Dactylis
glomerata
Digitaria
Echinochloa
crusgalli
Eleusine
Eragrostis
Festuca
pratensis
Gaudinia
Glyceria
aquatica
Hierchloe
Holcus
lanatus
Palaearctic Frost (1924) [as Notiphila flaveola]
Palaearctic Hering (1925, 1926, 1937, 1957)
Palaearctic Gercke (1882)
Nearctic
Palaearctic Hendel (1926), Hering (1937, 1957)
Palaearctic Grigarick (1959), Hering (1957)
Palaearctic Grigarick (1959), Hering (1957)
Palaearctic Grigarick (1959), Hering (1937,
1957), Linnaniemi (1913)
Palaearctic Grigarick (1959), Wilke (1924)
Palaearctic Hering (1957)
Palaearctic Grigarick (1959), Balachowsky and
Nearctic Mesnil (1935), Hering (1937),
Hendel (1926), von Frauenfeld
(1869), Sorauer and Reh (1913),
Kirchner (1923)
Palaearctic Hering (1957)
Palaearctic Grigarick (1959), Hering (1937),
Nearctic Hendel (1926)
Palaearctic Hering (1957)
Palaearctic Grigarick (1959), Hering (195 7)
Palaearctic Grigarick (1959), Hering (1957)
Palaearctic Grigarick (1959)
Palaearctic Grigarick (1959), Hering (1957),
Wilke (1924)
Palaearctic Grigarick (1959), Hering (195 7)
Nearctic Grigarick (1959)
Palaearctic Hering (1957)
Palaearctic Grigarick (1959)
Palaearctic Grigarick (1959), Hering (1957),
Wilke (1924)
Palaearctic Hering (1957)
Palaearctic Grigarick (1959), Hering (1922)
Palaearctic Grigarick (1959)
Palaearctic Grigarick (1959), Hering (1951,
1957), Wilke (1924)
NUMBER 6 8 109
TABLE 1.?Checklist of known host plants of Hydrellia.?Continued
Species of
Hydrellia
griseola
(cont'd)
Host
plant
Hordeum
vulgare
Lagurus
Lolium
perenne
Muhlenbergia
Oryza
Panicum
P asp alum
Phalaris
arundinacea
Phleum
pratense
Phragmites
Poa
annua
pratensis
trivialis
Polypogon
Scleropoa
Secale
cereale
Setaria
Triticum
sativum
Zea
Zizania
aquatica*
Alismataceae
Alisma
plantago-aquatica
Sagittaria
Damasonium
Caryophyllaceae
Lychnis
sp.
flos-cuculi
Stellaria
media
Chenopodiaceae
Kochia
Compositae
Bellis
Zoogeographic
region
Palaearctic
Nearctic
Palaearctic
Palaearctic
Nearctic
Palaearctic
Palaearctic
Palaearctic
Nearctic
Nearctic
Palaearctic
Nearctic
Palaearctic
Palaearctic
Palaearctic
Nearctic
Palaearctic
Palaearctic
Nearctic
Palaearctic
Palaearctic
Nearctic
Palaearctic
Palaearctic
Nearctic
Palaearctic
Nearctic
Nearctic
Nearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Major
references
Kirchner (1923), Grigarick (1959),
Frost (1924), Balachowsky and
Mesnil (1935), von Frauenfeld
(1869), Hendel (1926), Brischke
(1883), Seguy (1951)
Hering (1957)
Sorauer and Reh (1913), Wilke
(1924), Hering (1925, 1937,
1957), Grigarick (1959), Frost
(1924), Hendel (1926)
Grigarick (1959)
Grigarick (1959), Lange, et al.
(1953), DeOng (1922) [as H.
scapularis], Darby (1962), Hendel
(1926)
Grigarick (1959), Hering (1957),
Malloch (1915) [as H. scapularis]
Grigarick (1959)
Grigarick (1959), Hendel (1926),
Hering (1937, 1957)
Grigarick (1959), Hendel (1926),
Hering (1937), Linnaniemi
(1913), Wilke (1924)
Grigarick (1959), Hering (1957)
Grigarick (1959), Frost (1924),
Hendel (1926), Hering (1937),
Kirchner (1923)
Grigarick (1959)
Hering (1957)
Grigarick (1959), Hendel (1926),
Hering (1957)
Grigarick (1959), Hering (1957)
Grigarick (1959), Hendel (1926),
Hering (1957)
Grigarick (1959)
Grigarick (1959)
Grigarick (1959), Grimshaw (1925)
Grigarick (1959), Lange et al. (1953)
Grigarick (1959)
Grigarick (1959), Balachowsky and
Mesnil (1935), Hendel (1926),
Hering (1951, 1957)
Grigarick (1959), Frost (1924),
Hendel (1926), Hering (1957)
Grigarick (1959), Hcring (1937,
1957)
Grigarick (1959), Hering (1937,
1957)
110 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 1.?Checklist of known host plants of Hydrellia.?Continued
Species of Host
Hydrellia plant
griseola Cruciferae
(cont'd) Nasturtium
officinale*
Cyperaceae
Car ex
Cyperus
Scirpus
Hydrocharitaceac
Hydrocharis
morsus-ranae
Stratiotes
aloides
Labiatae
Lamium
purpureum
Leguminosae
Trifolium
Lcmnaceac
Lemna
minor
Liliaceae
A Ilium
cepa
porrum
Polygonaceae
Polygonum
Scrophulariaceae
Veronica
Typhaceae
Typha
latifolia
hydrocharitis Hydrocharitaceac
Hydrocharis
morsus-ranae
ischiaca Gramineae
Zizania
aquatica*
Glyceria
grandis*
obtusa*
Potamogetonaceae
Potamogeton
natans*
itascae Potamogetonaceae
Potamogeton
zosteriformis*
leucostoma Chenopodiaceae
Chenopodium
album
Zoogeographic Major
region
Nearctic
Palaearctic
Nearctic
Palaearctic
Nearctic
Nearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Nearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Nearctic
Palaearctic
Nearctic
Nearctic
Nearctic
Palaearctic
references
Grigarick (1959), Hendel (1926),
Hering (1937, 1957), Seguy
(1950)
Grigarick (1959), Hendel (1926)
Grigarick (1959), Lange et al. (1953)
Grigarick (1959), Hendel (1926),
Grimshaw (1925), de Meijere (1902)
Grigarick (1959), Hendel (1926),
Ruschka and Thienemann (1913),
Seguy (1950), Wahlgren (1947)
Grigarick (1959), Frost (1924),
Hendel (1926), Hering (1937,
1957)
Grigarick (1959), Hendel (1926),
Hering (1937, 1951, 1957)
Grigarick (1959), Bertrand (1954),
Grimshaw (1925)
Grigarick (1959), Balachowsky and
Mesnil (1935), Hering (1957)
Grigarick (1959), Hendel (1926)
Grigarick (1959), Hering (1937,
1957)
Grigarick (1959)
Hendel (1926), Hering (1925, 1926,
1931, 1937, 1951, 1957)
Frost (1924)
NUMBER 68
TABLE
Species of
Hydrellia
luctuosa
maura
modest a
mutata
nasturtii
nigricans
nigripes
potamogeti
propinqua
1.?Checklist of
Host
plant
Potamoge tonaceae
Potamogeton
alpinus
amplifolius
richardsonii
natans
zosteriformist
Cruciferae
Nasturtium
officinale
Potamoge tonaceae
Potamogeton
perfoliatus
Hydrocharitaceae
Hydrocharis
Potamoge tonaceae
Potamogeton
natans
sp.
Alismataceae
Alisma
plantago
Hydrocharitaceae
Stratiotes
aloides
Hydrocharis
morsus-ranae
Lemnaceae
Lemna
Cruciferae
Nasturtium
officinale
Juncaginaceae
Juncus
bufonius
articulatus
Potamogetonaceae
Potamogeton
Potamogetonaceae
Potamogeton
natans
drucei
Hydrocharitaceae
Stratiotes
aloides
known host plants of Hydrellia.?Continued
Zoogeographic Major
region
Nearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
references
Berg (1949, 1950)
Seguy (1950), Hering (1951)
Hering (1957), Bertrand (1954),
Hering (1950)
Keilin (1915)
Keilin (1915), Hering (1937),
Hendel (1926), Thienemann
(1912)
Hendel (1926), Hering (1937, 1957),
Griinberg (1910)
S6guy (1950), Wahlgren (1947),
Hering (1959)
Hendel (1926), Linnaniemi (1913),
Gercke (1878), Hering (1925,
1926), Grunberg (1910)
Linnaniemi (1913)
S?guy (1950), Hering (1937), Taylor
(1928), Vayssiere (1933), Collin
(1928), Hering (1957)
Hering (1957)
Hering (1937, 1957)
Hering (1937, 1957)
Wahlgren (1947), Hering (1957)
111
112 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 1.?Checklist of known host plants of Hydrellia.?Continued
Species of
Hydrellia
pulla
ranunculi
spinicornis
stratiotae
stratiotetla
tibialis
thoracica
trichaeta
viridescens
williamsi
xenophaga
spp.
Host
plant
Potamogetonaceae
Potamogeton
amplifoliust
gramineus
richardsonii
Cruciferae
Nasturtium
officinale
Gramineae
Hydrochloa
caroliniensis*
Hydrocharitaceae
Stratiotes
aloides
Hydrocharitaceae
Stratiotes
aloides
Cyperaceae
Eleocharis
obtusa*
Gramineae
Glyceria
aquatica
Hydrocharitaceae
Elodea
canadensis*
Potamogetonaceae
Potamogeton
epihydrus*
gramineus*
nodosus*
Alismataceae
?A. llSTYlG,
Potamogetonaceae
Potamogeton
perfoliatus
crispus
sp.
Lemnaceae
Lemna
Potamogetonaceae
Potamogeton
Alismataceae
Alisma
Sagittaria
latifoliat
Zoogeographic
region
Nearctic
Palaearctic
Nearctic
Palaearctic
Palaearctic
Nearctic
Palaearctic
Palaearctic
Nearctic
Nearctic
Palaearctic
Palaearctic
Australian
Palaearctic
Palaearctic
Nearctic
Major
references
Berg (1949, 1950)
Thienemann (1912), Hendel (1926),
Hering (1925), Taylor (1928)
Hering (1926, 1937, 1957), Seguy
(1950), Hendel (1926)
Hering (1957), Wahlgren (1947)
Hering (1926)
Hering (1925)
Hering (1957)
Hering (1950, 1951, 1957)
Hardy (1960)
Hering (1957)
Hering (1957)
NUMBER 6 8 113
TABLE 1.?Checklist of known host plants of Hydrellia.?Continued
Species of
Hydrellia
Host
plant
Zoogeographic
region
Major
references
Ranunculaceae
Haloragaceae
Myriophyllum
Potamogetonaceae
Potamogeton
Potamogetonaceae
Potamogeton
amplifolius
Polygonaceae
Polygonum
amphibium
persicaria
Palaearctic Hering (1937)
Palaearctic Schiner (1864)
Palaearctic Schiner (1864), Seguy (1930)
Nearctic Frost (1924)
Palaearctic Frost (1924)
TABLE 2.?Checklist of known parasitic wasps of Hydrellia.
Species of
Hydrellia
Parasitic
wasp
Zoogeographic
region
Major
references
albilabris
ascita
bergi
Braconidae
Chaenusa
conjungens
Chorebus
uliginosus
Dacnusa
liminicola
Braconidae
Chorebidella
sp.
Dacnusa
sp.
Diapriidae
Trichopria
columbiana
Braconidae
Ademon
niger
Chorebidea
sp.2
Dacnusa
sp. 1
Diapriidae
Trichopria
columbiana
Palaearctic Fulmek (1962)
Nearctic Berg (1950)
Nearctic Berg (1950)
Nearctic Berg (1950)
Nearctic Berg (1950)
NOTE: ? Reared from host for the first time during this study, t Reared from host
during this study.
114 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 2.?Checklist of known parasitic wasps of Hydrellia.?Continued
Species of
Hydrellia
Parasitic
wasp
Zoogeographic
region
Major
references
bilobifera
butomi
cochleariae
cruralis
fascitibia
griseola
Braconidae
Aphanta
sp.?
Chorebidella
bergi*
Opius*
sp.
Braconidae
Ademon
decrescens
Pteromalidae
genus undet.
(hyperparasite)
Braconidae
Ademon
decrescens
Chaenusa
conjungens
Chorebus
uliginosus
Opius
caesius
Pachysema
discolor
Chalcidae
Gonatocerus
uliginosus
Braconidae
Ademon
niger
Chorebidea
sp.
Chorebidella
bergi
Diapriidae
Trichopria
columbiana
Pteromalidae
genus undet.
Braconidae
Ademon
decrescens
Braconidae
Ademon
decrescens
sp.
Chaenusa
conjungens
sp.
Nearctic
Palaearctic Hering (1925)
Palaearctic Hering (1925)
Palaearctic Fulmek (1962), Thienemann (1916)
Palaearctic Fulmek (1962)
Nearctic Berg (1950)
Nearctic Berg (1950)
Nearctic Berg (1950)
Palaearctic Fulmek (1962)
Palaearctic Burghele (1959a, 1959b), Grigarick
(1959), Seguy (1934), Fulmek
(1962)
Palaearctic Burghele (1959a, 1959b), Grigarick
Nearctic (1959), Fulmek (1962)
NUMBER 6 8 115
TABLE 2.?Checklist of known parasitic wasps of Hydrellia.?Continued
Species of
Hydrellia
griseola
(cont'd)
Parasitic
wasp
Chorebus
uliginosus
nixoni
dense punctatus
orghidani
aquaticusi
sp.
Coelinius
hydrelliae
Merites(?)
Opius
hydrelliae
punctiventris
sp.
sp.
Pachysema
temula
Diapriidae
Trichopria
columbiana
sp.
Eucoilidae
Kleidotoma
striaticollis
Eulophidae
Achrysocharis
sp.
Asecodes
sp.
Chrysocharis
sp.
Derostemus
sp.
Elachertus(?)
sp.
Mestocharis(?)
sp.
Neochrysocharis
sp.
Paracrias
sp.
Pnigalio
sp.
Rhopalotus
sp.
Solenotus
sp.
intermedius
Sympiesis
sp.
Zoogeo graphic
region
Palaearctic
Palaearctic
Nearctic
Nearctic
Palaearctic
Palaearctic
Palaearctic
Nearctic
Palaearctic
Palaearctic
Neotropical
Palaearctic
Nearctic
Palaearctic
Neotropical
Palaearctic
Palaearctic
Nearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Nearctic
Palaearctic
Palaearctic
Nearctic
Nearctic
Major
references
Burghele (1959a, 1959b), Grigarick
(1959), Fulmek (1962),
Kuwayama (1955)
Burghele (1959a, 1959b), Grigarick
(1959)
Grigarick (1959), Malloch (1915)
[as Gyrocampa]
Kawall (1867), Grigarick (1959),
Fulmek (1962)
Kuwayama (1955), Grigarick (1959)
Burghele (1959a, 1959b), Fulmek
(1962), Grigarick (1959)
Grigarick (1959), Thompson (1943)
Kuwayama (1955)
Parker, et al. (1952)
Fulmek (1962)
Grigarick (1959)
Burghele (1959a, 1959b), Fulmek
(1962)
Parker, et al. (1952)
Kuwayama (1955)
Grigarick (1959)
Grigarick (1959)
Kuwayama (1955)
Kuwayama (1955)
Kuwayama (1955)
Kuwayama (1955)
Grigarick (1959)
Kuwayama (1955)
Kuwayama (1955)
Grigarick (1959)
Grigarick (1959)
116 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 2.?Checklist of known parasitic wasps of Hydrellia.?Continued
Species of
Hydrellia
griseola
(cont'd)
ischiaca
itascae
luctuosa
maura
mutata
Parasitic
wasp
Ichneumonidae
Hemiteles
sp.
Horogenes
fenestralis
Proctotrupidae
Ismarus
sp.
Pteromalidae
Eupteromalus
americanus
Halticoptera
sp.
Merismus
sp.
Polycystis
clavicornis
Pteromalus
sp.
Trichomalus
sp.
Braconidae
Chorebus
aquaticus*
Chorebidella
bergi*
Chaenusa
sp.*
Opius
sp.*
Diapriidae
Trichopria
columbiana*
Braconidae
Aphanta
sp.*
Chorebidella
bergi*
Braconidae
DacnusQ.
sp.
Diapriidae
Trichopria
columbiana
Braconidae
Pachysema
discolor
Braconidae
Chaenusa
conjungens
Zoogeographic
region
Palaearctic
Palaearctic
Palaearctic
Nearctic
Nearctic
Palaearctic
Palaearctic
Neotropical
Palaearctic
Nearctic
Nearctic
Nearctic
Nearctic
Nearctic
Palaearctic
Palaearctic
Major
references
Kuwayama (1955)
Fulmek (1962)
Kuwayama (1955)
Grigarick (1959)
Grigarick (1959)
Kuwayama (1955)
Burghele (1959a, 1959b), Fulmek
(1962)
Parker, et al. (1952)
Kuwayama (1955)
Berg (1950)
Berg (1950)
Fulmek (1962)
Fulmek (1962), Rimsky-Korsakov
(1917)
NUMBER 68 117
TABLE 2.?Checklist of known parasitic wasps of Hydrellia.?Continued
Species of
Hydrellia
nasturtii
pulla
stratiotae
spp.
viridescens
Parasitic
wasp
Braconidae
Ademon
decrescens
Braconidae
Ademon
niger*
Diapriidae
Trichopria
columbianai
Braconidae
Ademon
decrescens
Chorebus
uliginosus
Braconidae
Dacnusa
obscuripes
[listed as probable
parasite]
Gyrocampa
sp.
Ademon
mutator
Chorebidea
motasi
Chorebus
striola
Ademon
decrescens
mutator
Chaenusa
najadum
natator
Dacnusa
obscuripes
Gyrocampa
stagnalis
Opius
hydrelliae
Pachysema
discolor
Diapriidae
Ceratopria
lacustris
Braconidae
Ademon
decrescens
Chorebus
najadum
natator
Zoogeographic
region
Palaearctic
Nearctic
Nearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Palaearctic
Major
references
Thompson (1943, 1953), Fulmek
(1962), Vayssiere (1933)
Berg (1950)
Fulmek (1962)
Thienemann (1916), Burghele
(1959b)
Thienemann (1916)
Thompson (1953)
Burghele (1959a)
Burghele (1959b)
Fulmek (1962)
Fulmek (1962), Thienemann (1916)
[as H. chrysostoma]
118 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Literature Cited
Aldrich, J. M.
1905. A Catalogue of North American Diptera (or Two-
winged Flies). Smithsonian Miscellaneous Collec-
tions, volume 46, number 1444.
1912. The Biology of some Western Species of the Dip-
terous Genus Ephydra. Journal of the New York
Entomological Society, 20:77-103.
Allen, Pamela
1957a. The Larval Morphology of Agromyzidae (Dip-
tera). Proceedings of the Royal Entomological
Society of London, series A, 32:59-66.
1957b. The Larval Morphology of some Species of Phy-
tomyza Fallen ( Diptera: Agromyzidae). Pro-
ceedings of the Royal Entomological Society of
London, series A, 32:171-181.
Balachowsky, A., and L. Mesnil
1935. Les Insectes Nuisibles aux Plantes Cultivees. Vol-
ume I. Paris: Busson.
Becker, Theodor
1896. Dipterologische Studien IV. Ephydridae. Berliner
Entomologische Zeitschrift, 41:91-276, plates 4-7.
1903. Aegyptische Dipteren Gesammelt und Beschrieben.
Mitteilungen aus der Zoologischen Sammlung des
Museums fur Naturkunde in Berlin, 2 ( 1 ) : 67?195,
4 plates.
1919. Dipteres Brachypteres. In: Arc de Meridien Equa-
torial en Amerique du Sud, 1899-1906, volume 10
(Ent.-Bot.), fascicle 2, pages 163-215, plates 14-
17.
1926. Ephydridae [Family 56]. In: Erwin Lindner, edi-
tor, Die Fliegen der Palaearktischen Region,
volume 6, part 1. Stuttgart: E. Schweizerbart.
Berg, C. O.
1949. Limnological Relations of Insects to Plants of the
Genus Potamogeton. Transactions of the American
Microscopical Society, 68:279-291.
1950. Hydrellia (Ephydridae) and some Other Acalyp-
trate Diptera Reared from Potamogeton. Annals
of the Entomological Society of America, 43:374?
398.
Bertrand, Henri
1954. Les Insectes Aquatiques d'Europe (Genres: Larves,
Nymphes, Imagos). Volume 1. In: P. Lechevalier,
editor, Encyclopedie Entomologique, series A,
volume 30. Paris: Lechevalier et fils.
Bezzi, Mario
1894. I Ditteri del Trentino. Saggio di un Elenco Delle
Specie de Ditteri Finora Osservate nel Trentino.
Atti della Societa Veneto-Trentino di Scienze
Naturali, series 2, 1: 275-353.
1921. Ditteri di Cirenaica Raccolti dal Prof. Alessandro
Ghigi Durante l'Escursione Organizata dal Tour-
ing Club Italiano nel Mese d'Aprile 1920. Atti
della Societa Italiano di Scienze Naturali, 60:1-12.
Bolwig, Niels
1940. The Reproductive Organs of Scatophila unicornis
(Diptera). Proceedings of the Royal Entomological
Society of London, series A, 15:97-102.
1941. The Head of Scatophila unicornis Czerny (Dip-
tera). Proceedings of the Royal Entomological
Society of London, series A, 16:1-10.
Bonhag, P. F.
1949. The Thoracic Mechanism of the Adult Horsefly
(Diptera: Tabanidae). Memoir of the New York
Cornell University) Agricultural Experiment
Station, 285:3-39.
1951. The Skeleto-muscular Mechanism of the Head
and Abdomen of the Adult Horsefly (Diptera:
Tabanidae). Transactions of the American En-
tomological Society, 77:131-202.
Brischke, C. G. A.
1883. Breschreibung der Forst-, Garten- und Landwirt-
schaftlichen Feinde und Freunde unter den In-
sekten. Schriften der Naturforschenden Gesellschaft
in Danzig, new series, 5:97-125.
Brocher, Frank
1910. Observations Biologiques sur quelques Dipteres et
Hymenopteres dits "Aquatiques." Annales de
Biologie Lacustre, 3:170-176.
Burghele, Anca
1959a. New Rumanian Species of Dacnusini (Hym.,
Braconidae) and some Ecological Observations
upon Them. Entomologist's Monthly Magazine,
95:121-126.
1959b. Contributions a la Connaissance des Hymenopteres
Parasitant les Jeunes Stades d'Insectes Aquatiques.
[In Rumanian, with Russian and French sum-
maries]. Bucharest, Universitatea "C. I. Parhon,"
Analele, Seria Stuntelor naturii, 22:143-169.
Collin, J. E.
1928. A New Species of Hydrellia (Diptera, Ephydridae)
Mining the Stems of Watercress. Entomologist's
Monthly Magazine, 64:128-129.
Comstock, J. H.
1940. An Introduction to Entomology. 9th edition, re-
vised. Ithaca, New York: Comstock.
Coquillett, D. W.
1900. Papers from the Harriman Alaska Expedition. IX.
Entomological Results (3) : Diptera. Proceedings
of the Washington Academy of Science, 3:389-
464.
1903. A New Ephydrid from Australia. Entomological
News, 14:324.
1904. New North American Diptera. Proceedings of the
Entomological Society of Washington, 6:75.
1910a. New Genera and Species of North American
Diptera. Proceedings of the Entomological Society
of Washington, 12:124-131.
1910b. The Type-Species of the North American Genera
of Diptera. Proceedings of the U. S. National
Museum, 37:499-647.
NUMBER 6 8 119
Crampton, G. C.
1942. The External Morphology of the Diptera. In:
Guide to the Insects of Connecticut. Part VI. The
Diptera or True Flies of Connecticut. Fascicle 1.
Connecticut State Geological and Natural History
Survey Bulletin, 64:10-165.
1944. A Comparative Study of the Terminalia of Male
Calypterate Cyclorrhaphous Diptera and Their
Acalypterate Relatives. Bulletin of the Brooklyn
Entomological Society, 39:1-31.
Cresson, E. T., Jr.
1915. Descriptions of New Genera and Species of the
Dipterous Family Ephydridae. II. Entomological
News, 26:68-72.
1917. Descriptions of New Genera and Species of the
Dipterous Family Ephydridae. IV. Entomological
News, 28:340-341.
1918. Costa Rican Diptera Collected by Philip P. Cal-
vert, Ph.D., 1909-1910. Paper 3. A Report on
the Ephydridae. Transactions of the American
Entomological Society, 44:39-68, plate 3.
1924. Descriptions of New Genera and Species of the
Dipterous Family Ephydridae. VI. Entomological
News, 35:159-164.
1930. Studies in the Dipterous Family Ephydridae. III.
Transactions of the American Entomological So-
ciety, 56:93-131.
1931. Descriptions of New Genera and Species of the
Dipterous Family Ephydridae. IX. Entomological
News, 42:104-108.
1932. Studies in the Dipterous Family Ephydridae,
Paper IV. Transactions of the American Entomo-
logical Society, 58:1-34.
1934. Two New Species of the Genus Hydrellia from
Mount Desert, Maine (Diptera: Ephydridae).
Entomological News, 45:234-236.
1936. Descriptions and Notes on Genera and Species of
the Dipterous Family Ephydridae. II. Transac-
tions of the American Entomological Society, 62:
257-270.
1938. Notes on and Descriptions of some Neotropical
Ephydridae (Dipt.). Revista de Entomologia,
8:24-40.
1941. New Genera and Species of North American Eph-
ydridae (Diptera). Entomological News, 52:35-38.
1942. Descriptions of Two New Nearctic Species of the
Genus Hydrellia Reared from Pond-weed (Dip-
tera: Ephydridae). Entomological News, 53:
78-79.
1944a. Descriptions of Genera and Species of the Dip-
terous Family Ephydridae. Paper XIV. Notulae
Naturae, 135:1-9.
1944b. Synopses of North American Ephydridae (Dip-
tera). la. Supplement to Part I on the Subfamily
Psilopinae. II. The Tribes of Hydrelliini, Hydri-
nini, and Ilytheini of the Subfamily Notiphilinae,
wth Descriptions of New Species. Transactions of
the American Entomological Society, 70:159-180.
1947a. A Systematic Annotated Arrangement of the Gen-
era and Species of the Neotropical Ephydridae
(Diptera). II. The Subfamily Notiphilinae. Trans-
actions of the American Entomological Society,
73:35-61.
1947b. A Systematic Annotated Arrangement of the
Genera and Species of the Ethiopian Ephydridae
(Diptera). II. The Subfamily Notiphilinae. Trans-
actions of the American Entomological Society,
73:105-124.
1948. A Systematic Annotated Arrangement of the
Genera and Species of the Indo-Australian Ephy-
dridae (Diptera). II. The Subfamily Notiphilinae
and Supplement to Part I on the Subfamily Psilo-
pinae. Transactions of the American Entomological
Society, 74:1-28.
Curran, C. H.
1930. Report of the Diptera Collected at the Station
for the Study of Insects, Harriman Interstate Park,
New York. Bulletin of the American Museum of
Natural History, 61:21-115.
1934. The Families and Genera of North America Dip-
tera. New York: Author.
Dahl, R. G.
1959. Studies on Scandinavian Ephydridae (Diptera
Brachycera). Opuscula Entomologica. Supplemen-
tum, 15:1-224, figure 84.
1964. Revision av Slaktet Hydrellia R. D. (Dipt. Ephy-
dridae) i Coll. Stenhammar och Zetterstedt. Opus-
cula Entomologica, 29:179-187.
1967. Ephydridae. In: J. lilies, editor, Limnofauna Euro-
paea. Stuttgart: G. Fisher.
1968. Zur Kenntnis der Ephydriden (Diptera, Brachy-
cera) von Ostsibirien. Notulae Entomologicae, 48:
35-39.
Darby, R. E.
1962. Midges Associated with California Rice Fields,
with Special Reference to Their Ecology (Diptera:
Chironomidae). Hilgardia, 32:1-206.
DeOng, E. R.
1922. A Rice Leaf-miner. Journal of Economic En-
tomology, 15:432.
Deonier, D. L.
1964. Keys to the Shore Flies of Iowa (Diptera, Ephy-
dridae). Iowa State Journal of Science, 39:103?
126.
1965. Ecological Observations on Iowa Shore Flies (Dip-
tera, Ephydridae). Proceedings at the Iowa Acad-
emy of Science, 71:496-510.
Downes, W. L.
1955. Notes on the Morphology and Classification of the
Sarcophagidae and other Calypterates (Diptera).
Proceedings of the Iowa Academy of Science,
62:514-538.
1958. The Nearctic Miltograminae (Diptera, Sarcopha-
gidae) and Certain Allies. Unpublished Ph.D.
thesis. Library, Iowa State University of Science
and Technology, Ames, Iowa.
120 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Emden, F. van, and W. Hennig
1956. Diptera. In: S. L. Tuxen, editor, Taxonomist's
Glossary of Genitalia in Insects, p a g e s 1 1 1 - 1 2 1 .
Copenhagen: Ejner Munksgaard.
Fallen, C. F.
1813. Beskrifning Ofver nagra i Sverige funna Vattenflu-
gor (Hydromyzides). Kongliga Vetenskaps Aca-
demien Handlingar; 34:240-257.
Fassett, N. C.
1960. A Manual of Aquatic Plants. With Revision Ap-
pendix by E. C. Ogden. 2d edition. Madison:
University of Wisconsin Press.
Frauenfeld, G. R. von
1866. Uber die metamorphosis von Hydrellia albilabris
Meigen. Verhandlungen der Zoologisch-Botani-
schen Gesellschaft in Wien, 16:685.
1869. Ueber einige Pflanzenverwiister des Jahres 1869.
Verhandlungen der Zoologisch-Botanischen Gesell-
schaft in Wien, 19:601-604.
Frey, Richard
1921. Studien uber den Bau des Mundes der niederen
Diptera Schizophora nebst Bemerkungen iiber die
Systematik dieser Dipterengruppe. Ada Societas
pro Fauna et Flora Fennica, 48(3): 1-245.
1933. En Ammarkningsvard Insektstandort vid Viisjoki
(Ik). Notulae Entomologicae, 13:81-87.
Frick, K. E.
1952. A Generic Revision of the Family Agromyzidae
(Diptera) with a Catalogue of New World Spe-
cies. University of California Publications in En-
tomology, 8:330-452.
Frost, S. W.
1924. A Study of the Leafmining Diptera of North
America. Memoir of the New York (Cornell Uni-
versity) Agricultural Experiment Station, 78:1-
228.
Fulmek, L.
1962. Parasitinsekten der Blattminierer Europas. Haag,
Holland: W. Junk.
Gercke, G.
1879. Ueber die Metamorphose Nacktfliigeliger Cera-
topogon-Artcn sowie iiber die von Tanypus nigro-
punctatus Steg. und von Hydrellia mutata Meig.
Verhandlungen Vereins fur Naturwissenschaft-
liche Unterhaltung zu Hamburg, 4:522-228, 1
plate.
1882. Ueber die Metamorphose einiger Dipteren. Ver-
handlungen Vereins fur Naturwissenschaftliche
Unterhaltung zu Hamburg, 5:68-80, plates 1, 2.
1889. Dipterologische Miscellanen. (II. Serie). Wiener
Entomologische Zeitung, 8:219-222.
Glick, P. A.
1960. Collecting Insects by Airplane, with Special Ref-
erence to Dispersal of the Potato Leaf-hopper.
U. S. Department of Agriculture Technical Bul-
letin, number 1222.
Gobert, E.
1887. Catalogue des Dipteres de France. Revue d'En-
tomologie Publiee par la Sociiti Francaise d'En-
tomologie (Caen), 6(3): 1-87.
Goetghbuer, M.
1942. Faunule Dipterologique des Brise-Lames. Bulletin
de Musee Royale d'Histoire Naturelle de Belgique,
18(24):1-10.
Grensted, L. W.
1944. Records of Diperta and other Insects from Dinas
Head, Pembrokeshire. Entomologist's Monthly
Magazine, 80:201-203.
Grigarick, A. A.
1959. Bionomics of the Rice Leaf Miner, Hydrellia
griseola (Fallen), in California (Diptera: Ephy-
dridae). Hilgardia, 29:1-80.
Grimshaw, P. H.
1925. The Study of Flies (Diptera). Naturalist, 816:
5-20.
Griinberg, K.
1910. Diptera, Zweifliigler. In: A Brauer, editor, Die
Susswasserfauna Deutschlands. Eine Exkursions-
fauna. Volume 2A, part 1. Jena, Germany: G.
Fischer.
Hardy, D. E.
1960. Diptera: Nematocera-Brachycera (except Dolicho-
podidae). Volume 10. In: E. C. Zimmerman, edi-
tor, Insects of Hawaii. Honolulu: University of
Hawaii Press.
Harrison, R. A.
1959. Acalypterate Diptera of New Zealand. Bulletin
of the New Zealand Department of Scientific and
Industrial Research, 128:1-382.
Hendel, Friedrich
1926. Blattminenkunde Europas I. Die Dipterenminen.
Vienna: Fritz Wagner.
1933. Neue Acalyptrate Musciden aus der Palaarktischen
Region (Dipt.). Deutsche Entomologische Zeit-
schrift, 1933:39-56.
Hennig, Willi
1943. Ubersicht iiber Bisher Bekannten Metamorphoses-
tadien der Ephydriden, mit Neubeschreibungen
nach dem Material der Deutschen Limnologischen
Sunda-Expedition. Arbeiten iiber Morphologische
und Taxonomische Entomologie aus Berlin-
Dahlem, 10:105-138.
1952. Die Larvenformen der Dipteren. Part 3. Berlin:
Akademie-Verlag.
1958. Die Familien der Diptera Schizophora und
Ihre Phylogenetischen Verwandtschaftsbeziehun-
gen. Beitrage zur Entomologie (Berlin), 8:506-
688.
Hering, E. M.
1922. Monatliche Anweisungen: Blattminen und Blatt-
minierer. Entomologisches Jahrbuch, 31:36.
1924. Minenstudien IV. Zeitschrift fur Wissenschaftliche
Biologie, 2:217-250.
1925. Minenstudien VI. Zeitschrift fur Morphologie und
Okologie der Tiere, 4:502-539.
1926. Die Okologie der Blattminierenden Insektenlarven.
Zoologische Bausteine, 1:1-253, plates 1, 2.
NUMBER 6 8 121
1937.
1950.
Die Blattminen Mittel-und Nord-Europas ein-
schliesslich Englands?Bestimmungstabellen alter
von Insektenlarven der verschiedenen Ordnungen
erzeugten Minen. Neubrandenburg, Germany: G.
Feller.
Beitrag zur Kenntnis der Gattung Hydrellia (Dip-
tera: Ephydridae). Archiv fur Hydrobiologie, 43:
234-240.
1951. Biology of the Leaf Miners. 's-Gravenhage, Hol-
land: W. Junk.
1957. Bestimmungstabellen der Blattminen von Europa.
Volumes 1?3. 's-Gravenhage, Holland: W. Junk.
Hinton, H. E.
1955. On the Structure, Function, and Distribution of
the Prolegs of the Panorpoidea, with a Criticism
of the Berlese-Imms Theory. Transactions of the
Royal Entomological Society of London, 106:455?
556, plate 1.
Hollande, A.; J. Cachon; and F. Vaillant
1951. Recherches sur quelques Larves d'Insectes Termi-
tophiles (Muscidae, Calliphoridae, Oestridae,
Tineidae, Melandryidae). Annales des Sciences
Naturelles, Zoologie, series 11, 13:25-396.
Howard, L. O.
1900. A Contribution to the Study of the Insect Fauna
of Human Excrement. Proceedings of the Wash-
ington Academy of Science, 2:541-604, plates
30, 31.
Imms, A. D.
1957. A General Textbook of Entomology, Including
Anatomy, Physiology, Development and Classifi-
cation of Insects. 9th edition, revised by O. W.
Richards and R. G. Davies. London: Methuen.
International Commission on Zoological Nomenclature
1961. International Code of Zoological Nomenclature
Adopted by the XV International Congress of
Zoology. London: The International Trust for
Zoological Nomenclature.
Johannsen, O. A.
1935. Aquatic Diptera. Part II. Orthorrhapha-Brachy-
cera and Cylorrhapha. Memoir of the New York
(Cornell University) Agricultural Experiment
Station, 177:1-62, plates 1-12.
Johnson, C. W.
1904. A Supplementary List of the Diptera of New
Jersey. Entomological News, 15:157?163.
1925. Fauna of New England. 15. List of the Diptera or
Two-winged Flies. Occasional Papers of the Boston
Society of Natural History, 7:1-326.
Jones, B. J.
1906. Catalogue of the Ephydridae with Bibliography
and Description of New Species. University of
California Publications in Entomology, 1: 153-198.
Judd, W. W.
1964. A Study of the Population of Insects Emerging as
Adults from Saunders Pond at London, Ontario.
American Midland Naturalist, 71:402-414.
Kahl, Hugo
1917. Notes upon the Genus Leucophenga Mik (Dip-
tera) with Descriptions of New Species from
South America, West Africa, and the Philippine
Islands. Annals of the Carnegie Museum, 11:364-
393.
Kato, S.
1955. [Morphology of the Smaller Rice Leaf-miner]. In:
S. Kuwayama, editor [Investigations on Hydrellia
griseola Fallen, the Smaller Rice Leaf-miner. Spe-
cial Report Number 3 of the Society for Plant
Protection of North Japan]. Author, title, and
journal name translated from Japanese.
Kawall, J. H.
1867. Miscellanea Entomologica. Stettiner Entomolo-
gische Zeitung, 28:120-121.
Keilin, D.
1915. Recherches sur les Larves de Dipteres Cyclor-
haphes. Bulletin de Science des France et Belgique,
series 7, 49:15-198, plates 1-16.
1944. Respiratory Systems and Respiratory Adaptations
in Larvae and Pupae of Diptera. Parasitology, 36:
1-66, plates 1,2.
Kelly, K. L., and D. B. Judd
1955. The ISCC-NBS Method of Designating Colors
and a Dictionary of Color Names. National Bureau
of Standards, U. S. Department of Commerce,
Circular 553.
Kim, K. C , and E. F. Cook
1966. A Comparative External Morphology of Adult
Sphaeroceridae (Diptera). Miscellaneous Publica-
tions of the Entomological Society of America,
5:78-89.
Kirchner, O.
1923. Die Krankheiten und Beschddigungen Unserer
Landwirtschaftlichen Kulturpflanzen. 3d Edition.
Stuttgart: Eujen Ulmer.
Kloet, G. S., and W. D. Hincks
1945. A Check List of British Insects. Stockport, Eng-
land: Authors.
Kowarz, Ferdinand
1894. Verzeichnis der Insekten Bohmen's. II. Fliegen.
Catalogus Insectorum Faunae Bohemicae. Prague:
Physiokratisch Gesellschaft.
Kreuter, E. A.
1927. [Contribution to the Biology of the Barley Fly.
Report of the Bureau of Applied Entomology, 3:
92-98]. Author, title, and journal name translated
from Russian.
Kuwayama, S., editor
1955. [Investigations on Hydrellia griseola Fallen, the
Smaller Rice Leaf-Miner. Special Report Number
3 of the Society for Plant Protection of North
Japan]. Editor, title, and journal name translated
from Japanese.
Lameere, A.
1907. Manuel de la Faune de Belgique. Volume 3.
Brussels: H. Lambertin.
122 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Lampa, Sven
1906. Berattelse till Kungl. Landtbruks-styrelsen An-
gande Verksamheten vid Statens Entomologiska
Anstalt under Ar 1905. Entomologisk Tidskrift,
27:17-64.
Lange, W. H., Jr.; K. H. Ingebretsen; and L. L. Davis
1953. Rice Leaf Miner. California Agriculture, 7:8-9.
Laurence, B. R.
1952. Observations on Hydrellia (Hydropota) griseola
(Fallen) (Dipt., Ephydridae). Entomologist's
Monthly Magazine, 88:31-33.
Le Berre, J. R.; H. Chevin; and J. P. Moreau
1961. Longevite, Fecondite et Nutrition de Deux Dip-
teres: "Oscinella frit" L., "Hydrellia griseola"
Fall. Revue de Zoologie Agricole (Talence), 60:
151-161.
Lilljeborg, W.
1861. En Flugas Harjningar a Kornfalten i Ostra
Skane, Blekinge och Sddra Delarna af Kalmar Ian
under Sommaren 1860. Tidskrift for Landtmanna
och Kommunalekonomien (Upsala), 1861:205-
215.
Linnaniemi, W. M.
1913. Zur Kenntnis der Blattminierer, Speziell derjeni-
gen Finnlands. I. Ada Societas pro Fauna et Flora
Fennica, 37:1-138.
Loew, Hermann
1860. Die Europaeischen Ephydrinidae und die Bisher
in Schlesien Beobachteten Arten Derselben. Neue
Beitrage zur Kenntnis der Dipteren. Part 7. Berlin:
Mittler und Sohn.
1861. Diptera Americae Septentrionalis Indigena. Gen-
turia prima. Berliner Entomologische Zeitschrift,
5:307-359.
1862. Monographs of the Diptera of North America.
Part 1. Smithsonian Miscellaneous Collections,
volume 6, number 141.
1864. Diptera Americae Septentrionalis Indigena. Cen-
turia quinta. Berliner Entomologische Zeitschrift,
8:47-104.
1870. Ueber die Bisher auf der Galizischen Seite des
Tatragebirges Beobachteten Dipteren. Jahrbuche
der k. k. Gelehrten Gesellschaft in Krakau, 41:
1-19.
1872. Diptera Americae Septentrionalis Indigena. Cen-
turia decima. Berliner Entomologische Zeitschrift,
16:49-115.
Macquart, Jean
1835. Histoire Naturelle des Insectes Dipteres. Volume 2.
Paris: Roret.
Malloch, J. R.
1915. Some Additional Records of Chironomidae for
Illinois and Notes on other Illinois Diptera. Bul-
letin of the Illinois State Laboratory of Natural
History, 11:305-364.
Marchal, Paul
1903. Sur la Biologie des Hydrellia (Dipt.) Degats Ex-
erces sur le Cresson par VHydrellia ranunculi Hal.
Bulletin de la Societe Entomologique de France,
1903:236-237.
Meijere, J. C. H. de
1902. Ueber die Prothorakalstigmen der Dipterenpuppen.
Zoologische Jahrbucher. Abteilungen fur Anatomie
und Ontogenie, 15:623-692.
1939. Naamlijst van Nederlandsche Diptera. Tijdschrift
voor Entomologie, 82:137-174.
Miller, Albert
1950. Internal Anatomy and Histology of Imago. In:
M. Demerec, editor, Biology of Drosophila. Pages
420-534. New York: John Wiley and Sons.
Milne, L. J., and Margery Milne
1959. What Do Animals See? American Scholar, 28:
39-48.
Moragues y de Manzanos, F.
1894. Insectos de Mallorca. Anales de la Sociedad Es-
pafiola de Historia Natural, 23:73-87.
Morley, Claude, and E. A. Atmore
1915. The Diptera of Norfolk and Suffolk. Transactions
of the Norfolk and Norwich Naturalists' Society,
10:162.
Nemenz, H.
1960. Beitrage zur Kenntnis der Biologie von Ephydra
cinerea Jones 1906. (Diptera, Ephydridae). Zoolo-
gischer Anzeiger, 165:218-226.
Nye, I. W. B.
1958. The External Morphology of some of the Dipterous
Larvae Living in the Gramineae of Britain. Trans-
actions of the Royal Entomological Society of
London, 110:411-487.
Osten Sacken, C. R.
1878. Catalogue of the Described Diptera of North
America. 2d edition. Smithsonian Miscellaneous
Collections, volume 16, number 270.
1881. An Essay of Comparative Chaetotaxy, or the Ar-
rangement of Characteristic Bristles of Diptera.
Mitteilungen Munchener Entomologische Verein,
5:121-140.
Parker, H. L.; P. A. Berry; and A. Silveira Guido
1952. Host-parasite and Parasite-host Lists of Insects
Reared in the South American Parasite Labora-
tory During the Period 1940-1946. Revista de la
Asociacion de Ingenieros Agronomos, 92:15-112.
Ping, Chih
1921. The Biology of Ephydra subopaca Loew. Memoir
of the New York (Cornell University) Agricul-
tural Experiment Station, 49:557-616.
Procter, William
1938. Biological Survey of the Mount Desert Region
Founded and Directed by William Procter. Part
VI. The Insect Fauna. Philadelphia: Wistar
Institute of Anatomy and Biology.
Rimsky-Korsakov, M.
1917. Observations Biologiques sur les Hymenopteres
Aquatiques. Revue Russe d'Entomologie (for
1916),16:209-225.
NUMBER 6 8 123
Robineau-Desvoidy, A. J. B.
1830. Essai sur les Myodaires. Memoires da I'Academic
Royale des Sciences de I'Institut de France, 2:1?
813.
Ruschka, F., and August Thienemann
1913. Zur Kenntnis der Wasserhymenopteren. Zeitschrift
fur Wissenschaftliche Insektenbiologie, 9:82?87.
Schiner, J. R.
1864. Fauna Austriaca. Die Fliegen (Diptera). Volume
2, part 1. Vienna: Carl Gerold's Sohn.
Schoyen, T. H.
1930. Beretning om Skadeinsektenes optreden i Land-og
Havebruket in Arene 1928 og 1929. Landbrucks-
direkt. Arsberet, 1930:1-36.
Schiitte, L.
1921. Das Tonnchen der Musciden. Zoologischer Anzei-
ger, 53:49-51.
Scotland, Minnie B.
1940. Review and Summary of Studies on Insects Asso-
ciated with Lemna minor. Journal of the New
York Entomological Society, 48:319-333.
Se'guy, E.
1930. Contribution a l'Etude des Dipteres du Maroc.
Memoires de la Societi des Sciences Naturelle du
Maroc, 24:1-206.
1934. Dipteres (Brachyceres) (Muscidae Acalypterae et
Scatophagidae). Faune de France. Volume 28.
Paris: Lechevalier et fils.
1950. La Biologie des Dipteres. In: P. Lechevalier, edi-
tor, Encyclopedic Entomologique, series A, volume
26. Paris: Lechevalier et fils.
1951. Atlas des Dipteres de France.?Belgique?Suisse.
Paris: N. Boubee.
Slosson, Annie T.
1902a. Additional List of Insects Taken in Alpine Region
of Mt. Washington. Entomological News, 13:319-
321.
Snodgrass, R. E.
1935. Principles of Insect Morphology. New York: Mc-
Graw-Hill.
1944. The Feeding Apparatus of Biting and Sucking
Insects Affecting Man and Animals. Smithsonian
Miscellaneous Collections, volume 104, number
3773.
1953. The Metamorphosis of a Fly's Head. Smithsonian
Miscellaneous Collections, volume 122, number
4133.
1957. A Revised Interpretation of the External Repro-
ductive Organs of Male Insects. Smithsonian
Miscellaneous Collections, volume 135, number
4309.
1959. The Anatomical Life of the Mosquito. Smithsonian
Miscellaneous Collections, volume 139, number
4388.
Sorauer, P., and L. Reh
1913. Handbuch der Pflanzenkrankheiten. Die Tierischen
Feinde. Volume 3. Berlin: Paul Parey.
Stein, Friedrich
1867. Eine der Gerste Schadliche Fliege. Berliner En-
tomologische Zeitschrift, 11:395-397, plate 3,
figures 7-10.
Steyskal, G. C.
1957. The Postabdomen of Male Acalyptrate Diptera.
Annals of the Entomological Society of America,
50:66-73.
St8rmer, K., and R. Kleine
1911. Die Getreidefliegen, mit Besonder Beriicksichti-
gung Ihrer Wirtschaftlichen Bedeutung und der
Abhangigkeit Ihres Augtretens von Wittersungs-
verhaltnissen. Frohlings Landwirtschaftliche Zei-
tung, 60:682-703.
Strobl, P. G.
1900. Spanische Dipteren. VIII. Wiener Entomologische
Zeitung, 19:1-10.
1904. Neue Beitrage zur Dipterenfauna der Balkanhal-
binsel. Wissenschaftliche Mitteilungen aus Bos-
nien und der Herzegowina, 9:519?581. (Heraus-
gegeben vom Bosnisch-Herzegowinischen Landes-
museum in Sarajevo).
Sturtevant, A. H.
1925. The Seminal Receptacles and Accessory Glands of
the Diptera, with Special Reference to the Acalyp-
teratae. Journal of the New York Entomological
Society, 33:195-215.
1926. The Seminal Receptacles and Accessory Glands of
the Diptera, with Special Reference to Acalyp-
teratae. Journal of the New York Entomological
Society, 34:1-21, plates 1-3.
Sturtevant, A. H., and M. R. Wheeler
1954. Synopses of Nearctic Ephydridae (Diptera).
Transactions of the American Entomological So-
ciety, 79:151-257.
Taylor, T. H.
1928. The Watercress Stem-miner. Entomologist's
Monthly Magazine, 64:126-128.
Thienemann, August
1912. Bemerkungen zum Erstein Dipterenheft der
"Siisswasserfauna Deutschlands." Entomologische
Mitteilungen, 1:275-279.
1916. Ueber Wasserhymenopteren. Zeitschrift fiir Wissen-
schaftliche Insektenbiologie, 12:49-54.
Thompson, W. R.
1943. A Catalogue of Parasites and Predators of Insect
Pests. Section 1, part 2. Bellville, Ontario: Im-
perial Parasite Service.
1953. A Catalogue of Parasites and Predators of Insect
Pests. Section 2, part 2. Ottawa, Ontario: Com-
monwealth Bureau of Biological Control.
Timon-David, Jean
1961. Contribution a l'Etude de l'Ecologie et du Peuple-
ment des lies de la Region Marseillaise. Colloques
Internationaux du Centre National de la Recher-
ches Scientifique. XCIV. Le Peuplement des lies
Mediterraneennes et le Probleme de L'Insularite.
II. Liste des Dipteres Captures a Ratonneau-
Pomegues. Pages 228-230.
124 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Tonnoir, A. L., and J. R. Malloch
1926. New Zealand Muscidae Acalyptratae. Records of
the Canterbury Museum, 3:1-26.
Tragardh, I.
1903. Beitrage zur Kenntnis der Dipterenlarven. 1. Zur
Anatomie und Entwicklungsgeschichte der Larve
von Ephydra riparia Fall. Arkiv fiir Zoologie,
1:1-42, plates 1-4.
Tsacas, L.
1959. Contribution a la Connaissance des Dipteres de
Grece. 1. Bulletin de la Societi Entomologique de
France, 64:123-130.
1960. Sur Quelques Dipteres de l'lle Majorque. Eos,
36:237-244.
Tuxen, S. L.
1956. Taxonomist's Glossary of Genitalia in Insects.
Copenhagen: Ejner Munksgaard.
Vayssiere, P.
1933. La Mouche du Cresson (Hydrellia nasturtii Collin
[Dipt. Ephydridae]) et son Parasite (Ademon
decrescens Nees [Hym. Braconidae]). Bulletin de
la Societi Entomologique de France, 38:86-87.
Wahlgren, Einar
1947. Hydrellia-?Arten als Blattminierer in Stratiotes
aloides L. (Dipt.). Opuscula Entomologica, 12:
78-79.
Walker, Francis
1856. Insecta Brittanica. Diptera. Volume 3. London:
John Edward Taylor.
Washburn, F. L.
1906. Additional Minnesota Diptera. Eleventh Annual
Report of the State Entomologist of Minnesota,
1906:79-82.
Westwood, J. O.
1840. An Introduction to the Modern Classification of
Insects. Synopsis of the Genera of British Insects.
London: Ornie, Brown, Green, and Longmans.
Wigglesworth, V. B.
1956. Insect Physiology. 5th edition, revised. London:
Methuen.
Wilke, S.
1924. Die Graue Gerstenminierfliege, Hydrellia griseola
Fall. (Diptera: Ephydridae). Deutsche Ento-
mologische Zeitschrift, 6:172-179.
Wirth, W. W.
1948. A Taxonomic Study of Hawaiian Ephydridae
(Diptera) Related to Scatella Robineau-Desvoidy.
Proceedings of the Hawaiian Entomological So-
ciety, 13:277-304.
1956. The Ephydridae (Diptera) of the Bahama Islands.
American Museum Novitates, 1817:1-20.
1965. Ephydridae. In: Alan Stone; C. W. Sabrosky; W.
W. Wirth; R. H. Foote; and J. R. Coulson. A
Catalog of the Diptera of America North of Mex-
ico. Pages 734-759. Washington, D. C : U. S.
Government Printing Office.
1968. Family Ephydridae. In: P. Vanzolini, editor, A
Catalogue of the Diptera of the Americas South of
the United States. Sao Paulo: Departmento de
Zoologia, Secretaria da Agricultura.
1969. New Species and Records of Galapagos Diptera.
Proceedings of the California Academy of Sciences,
series 4, 36(20): 571-594.
Wirth, W. W., and Alan Stone
1956. Aquatic Diptera. In: R. L. Usinger, editor. Aqua-
tic Insects of California. Pages 372-482. Berkeley,
California: University of California Press.
Wulp, F. M. van der, and J. C. H. de Meijere
1898. Nieuwe Naamlijst van Nederlandsche Diptera.
Tijdschrift voor Entomologie, 41:1?143.
Zetterstedt, J. W.
1846. Diptera Scandinaviae Disposita et Descripta.
Volume 5. Lund, Sweden: Author.
NUMBER 6 8 125
Localities of Specimens Illustrated
ALASKA
King Salmon, Naknek River; Figure 35.
Matanuska Valley; Figures 19, 111.
BRITISH COLUMBIA
Qualicum; Figure 66.
CALIFORNIA
Davis; Figures 84, 95, 116.
Maxwell, Colusa County; Figure 138.
Monterey County; Figure 57.
Sacramento; Figures 42, 94.
Sardine Creek, Mono County; Figure 51.
COLORADO
Walden Lake; Figure 62.
CONNECTICUT
Canaan; Figure 52.
New Haven Harbor, west shore; Figure 22.
FLORIDA
Biscayne Bay; Figure 54.
Lacoochee; Figure 31.
Port St. Joe Beach, Gulf County; Figure 58.
(Unspecified) Figures 24, 71.
ILLINOIS
Havana; Figure 67.
IOWA
Banner Mine Area, Warren County; Figures 41, 87, 118.
Little Wall Lake, Hamilton County; Figures 1, 3, 4, 5, 6,
10,11,13,130,133-137,139-142.
Springbrook State Park, Guthrie County; Figures 9, 64.
Spring Lake, Greene County; Figures 20, 37, 73, 75, 88,
115.
Three miles east-southeast of Waterville, Allamakee
County; Figure 8.
KANSAS
Kansas University Natural History Reservation, Douglas
County; Figures 2, 12, 14, 85, 91, 109.
MAINE
Bar Harbor, Mount Desert Island; Figure 39.
MARYLAND
Plummer's Island, Potomac River; Figure 46.
MASSACHUSETTS
Woods Hole; Figure 18.
MICHIGAN
Bessey Creek, Cheboygan County; Figure 92.
Douglas Lake, Cheboygan County; Figures 89, 96, 107,
114, 132.
Nigger Creek, Cheboygan County; Figures 86, 112.
Third Sister Lake, Washtenaw County; Figures 61, 106,
108.
(Unspecified) Figure 123.
MINNESOTA
Biological Station, Itasca State Park; Figure 28.
Bohall Trail Bog, Itasca State Park; Figure 36.
Bohall Lake, Itasca State Park; Figure 68.
Chambers' Creek, Itasca State Park; Figures 7, 65.
Douglas Lodge Bay, Lake Itasca; Figures 44, 76, 102, 110.
Lake Itasca, Itasca State Park; Figures 82, 126, 129.
Long Lake, Clearwater County; Figure 128.
Mississippi River at Sucker Creek, Clearwater County;
Figures 100, 122.
Mississippi River, sec. 34, T. 145 north, R. 36 west, Clear-
water County; Figure 59.
One and three-tenths miles south of main entrance to
Itasca State Park; Figures 33, 74.
Rapid River Logging Camp, Hubbard County; Figures
98, 131.
Sucker Creek, 100 yards north of State Highway 31, Clear-
water County; Figure 47.
Squaw Lake, Itasca State Park; Figures 55, 63.
Two Island Lake, Itasca State Park; Figure 25.
MISSISSIPPI
Dickinson's Pond, Lamar County; Figures 43, 99, 103,
117, 120.
Gravel pit near U.S. Highway 90, sec. 20, T. 7 south,
R. 5 west, Jackson County; Figures 34, 81, 101, 119.
Gulf Coast Research Laboratory, Ocean Springs; Figure
27.
Lake Shady, Lamar County; Figures 32, 40, 78, 79, 83,
90, 93, 105, 124.
NEW YORK
Dryden Lake, Tompkins County; Figures 17, 60.
Inlet Valley, Ithaca; Figures 104, 121.
ONTARIO
Grand Bend; Figure 56.
Midland; Figure 23.
Ottawa; Figures 21, 53.
PENNSYLVANIA
Ohiopyle; Figure 26.
QUEBEC
Perkins Mills; Figures 15, 69.
LaTrappe; Figure 48.
TENNESSEE
Brewer's Bar Ditch, Reelfoot Lake; Figures 70, 72, 97, 113.
Dale Hollow Reservoir, six miles south of Byrdstown,
Pickett County; Figure 50.
Miller's Camp, Reelfoot Lake; Figures 30, 77, 125.
Samburg, Reelfoot Lake; Figures 29, 80, 127.
WASHINGTON
Ilwaco; Figure 38.
O'Sullivan Dam, Grant County; Figure 49.
Tacoma; Figure 45.
Three miles west of Kettle Falls; Figure 16.
Abbreviations Used on Illustrations
AAC, preacrostichal
AC, anteclypeus
ACB, anteclypeal breadth
ACW, anterior cibarial wall
ADC, predorsocentral
ADT, accessory-gland duct
AEM2, mesanepimeron
AES2, mesanepisternum
AFR, anterior fronto-orbital
AG, accessory gland
AM, adhesive material
AN, anus
ANT, antenna
AOD, anteocellar distance
AP, anal plate
APR, supraspiracular asteriform process
ARR, aristal rays
AS, annuli of spinules
ASC, apical scutellar
A3AP, apicodorsal antennals
BC, basicoxal
BEC, basal end of costa
BF, basal follicle
BP, basiproboscis
BPH, basiphallus
BS, basiphallic socket
BSC, basal scutellar
CA, cardia
CAN, canaliculi
CAP, halter capitellum
CAR, clypeal arch
CB, cibarium
CE, compound eye
CER, cercus
CF, distal costal fracture ("break")
CH, chorion
CL, copulobus
CM, crop meatus
CO, common oviduct
CR, crop proper
CS, cervical sclerite 2
CSN, cibarial sensillum
CSP, cheliform spot
CTH, canalicular teeth
CW, creeping welt
CXI, fore coxa
CX2, mid coxa
CX3, hind coxa
CY, calyx
DC, cibarial dilators
DCW, dorsal cibarial wall
DPH, distiphallus
DPR, dorsal phragmatal ramus
EDT, ejaculatory duct
EE, escape exit made by parasite
EO, dehydrated embryo
EP, epistoma
ES, epithelial sheath
ESP, esophagus
FCS, epistomal sulcus
FE, femur
FPL, frontoclypeal plate
GA, gonal arch
GC, genital chamber
GE, genal
h, humeral crossvein
HLS, head-lobe scar
IA, interalar
IFC, interfractural costals
ISC, intermediate scutellar
IV, inner vertical
KES2, mesokatepisternum
LA, labellum
LAB, labium (LA in Figure 81)
LAP, labral apodeme
LDT, labial-glad duct
LF, parafrontale
LG, labial gland
LGD, labial-gland duct opening
LM, labrum
LO, lateral ocellus
LR, longitudinal ridges
LS, labial sclerite
LT, laterotergite
LU, frontal lunule
M, mesomeron
m, medial crossvein
Mx + 2, combined first and second
medial vein
M3 + Cuj, combined third medial and
first cubital vein
MDA, mouth-hook depressor apodeme
ME, meconium
MF, frontal vitta
MFH, mesofacial height
MG, midgut
MH, mouth-hook
MO, median ocellus
MOT, mediotergite
MPS, maxillary palpus
MP, metapleuron
MPP, micropylar protuberance (MP in
Figure 125)
MS, mesopleural sulcus
MT, Malpighian tubules
NP, notopleuron
OC, ocellar
OL, lateral oviduct
OO, oocyte
OR, ovary
OV, outer vertical
PA, postalar
PAC, postacrostichal
PAP, phallapodeme
PBS, probasisternum
PCP, paraclypeal phragma
PCW, posterior cibarial wall
PD, halter pedicel
PDC, postdorsocentral
PDO, ovariole pedicel
PF, parafaciale
PFA, primary facials
PFR, posterior fronto-orbital
POG, postgonite
PH2, thoracic phragma 2
PHD, phallic depressors
PHL, phallic levators
PLP, postlabial pad
PM, prementum
PO, postocellar
PP, propleuron
PPH, paraclypeal phragma (Figures 84,
85).
PPN, postpronotum
PR, prestomum
PRG. pregonite
PRS, prescutellar
PSP, peritremal spinous seta
PT, spiracular peritreme
PTF, ptilinal fissure
PTL, ptilinum
PTO, primary tracheal orifice
PU, postgonite uncus
Rj, first radial vein
R-2 ? 3> combined second and third ra-
dial vein
R4 + 5, combined fourth and fifth ra-
dial vein
RG, rectal gland
r-m, radiomedial crossvein
RT, rectum
RV, rectal valve
SI?5, sterna 1?5
S8, sternum 8
SA, subalare
SAO, secondary atrial orifice
SAT, spiracular atrium
SB, halter scabellum
SBP, subgenital plate
SCB, subcranial breadth
SI, food-neatus siphon
SL, supra-alar
NUMBER 6 8 127
Abbreviations Used on Illustrations?Continued
SO, spermatogonia
SOH, subocular height
SP, lateral spermatheca
SP1, prothoracic spiracle
SP3, metathoracic spiracle
SPP, supraspiracular protuberance
SSP, supraspiracular spinous seta
SPT, subspiracular protuberance
SS, fused surstyli
ST, spermatic tubule
SV, stomodeal valve
SVS, seminal vesicle
SZ, spermatozoa
T4, tergum 4
T5, tergum 5
T9 ? io, syntergum 9-(-10
TB, tibia
TF, terminal filament
TS, testis
VB, vertex breadth
VD, vas deferens
VOD, vertical ocular height
VPR, ventral phragmatal ramus
VR, median spermatheca
WA, wing area
WL, wing length
WVR, wall of median spermatheca
128 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
LO MO PO IV OV
CM SV
0.10MM
FIGURES 1-3.? 1, Hydrellia griseola (Fallen), male: head, cleared, frontal view. 2, H. bilobifera
Cresson, female: internal genitalia, ventral view. 3, H. griseola (Fallen), female: gut, dorsal
view.
NUMBER 6 8 129
PDC
FIGURES 4-6.?Hydrellia gnseola (Fallen): 4, abdomen, male, ventral view; 5, posterior half
of abdomen, male, lateral view; 6, thorax proper, male, lateral view.
130 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
13
FIGURES 7-14.?7, Hydrellia crassipes Cresson, male: left hind femur and tibia, anteroventral
view. 8, H. morrisoni Cresson, male: left hind tibia, anterior view. 9, H. harti Cresson, male:
head, lateral view. 10, 11, H. griseola (Fallen): 10, male, terminalia, lateral view; 11, male,
head, cleared, frontal view. 12, H. bilobifera Cresson, male: musculature of gonal arch and
phallapodeme, dorsal view. 13, H. griseola (Fallen) : male terminalia, dorsal view. 14. H.
bilobifera Cresson: male internal genitalia, ventral view.
NUMBER 6 8 131
22
FIGURES 15?22.?Male terminalia of adult Hydrellia species (ventral view of left half unless
otherwise specified): 15, H. harti Cresson; 16, H. morrisoni; 17, whole structure for H. bergi
Cresson; 18, H. prudens Curran; 19, H. caliginosa Cresson; 20, H. procteri Cresson; 21,
H. idolator, new species; 22, H. valida Loew.
132 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
27 28
FIGURES 23?30.?Male terminalia of adult Hydrellia species (ventral view of left half unless
otherwise specified): 23, H. ascita Cresson; 24, H. gladiator, new species; 25, H. surata, new
species; 26, H. griseola (Fallen); 27, H. americana Cresson; 28, H. itascae, new species; 29,
H. rixator, new species; 30, H. discursa, new species.
NUMBER 6 8 133
'l '? * * * ? ? '
tot.
35
1
36 37
FIGURES 31?38.?Male terminalia of adult Hydrellia species (ventral view of left half unless
otherwise specified): 31, H. floridana, new species; 32, H. ainsworthi, new species; 33, H.
crassipes Cresson; 34, H. biloxiae, new species; 35, H. serena Cresson; 36, H. nobilis (Loew) ;
37, H. trichaeta Cresson; 38, H. platygastra Cresson.
134 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
fV'/l
43 ?.??
FIGURES 39?46.?Male terminalia of adult Hydrellia species (ventral view of left half unless
otherwise specified) : 39, H. advenae Cresson; 40, H. spinicornis Cresson; 41, H. tibialis
Cresson; 42, H. deceptor, new species; 43, H. formosa Loew; 44, H. ischiaca Loew; 45, H.
subnitens Cresson; 46, H. insulata, new species.
NUMBER 6 8 135
vr.J
-i
53
FIGURES 47?54.?Male terminalia of adult Hydrellia species (ventral view of left half unless
otherwise specified): 47, H. amnicola, new species; 48, H. manitobae, new species; 49, H.
personata, new species; 50, H. bilobifera Cresson; 51, H. melanderi, new species; 52, H. notata,
new species; 53, H. borealis Cresson; 54, H. cavator, new species.
136 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
59 62
FIGURES 55-62.?Male terminalia of adult Hydrellia species (ventral view of left half unless
otherwise specified): 55, H. columbata, new species; 56, H. saltator, new species; 57, H.
proclinata Cresson; 58, H. agitator, new species; 59, H. cessator, new species; 60, H. definita
Cresson; 61, H. pulla Cresson; 62, H. flavicoxalis Cresson.
NUMBER 6 8 137
72
FIGURES 63?72.?63?66, Male terminalia of adult Hydrellia species (ventral view of left
half unless otherwise specified): 63, H. cruralis Coquillett; 64, H. luctuosa Cresson; 65,
H. notiphiloides Cresson; 66, H. penicilli Cresson. 67-72, Female terminalia of adult Hydrellia
species (lateral view of left side): 67, H. ascita Cresson; 68, H. trichaeta Cresson; 69, H.
harti Cresson; 70, H. discursa, new species; 71, H. gladiator, new species; 72, H. bilobifera
Cresson.
138 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
83
FIGURES 73?83.?73, Hydrellia bilobifera Cresson: egg chorion, parasagittal section. 74, H.
bergi Cresson: egg, lateral view. 75, H. bilobifera Cresson: egg, dorsal view. 76, H. ischiaca
Loew: egg, ventral view. 77, H. discursa, new species: egg, dorsal view. 78, H. spinicornis
Cresson: egg, dorsal view, shading indicates numerous punctulae or pits. 79, H. spinicornis
Cresson: posterior part of abdominal segment 8 and tracheospiracular siphon of second-instar
larva, dorsal view. 80 H. discursa, new species: abdominal segment 8 and spiracular peritreme
of first-instar larva, lateral view of left side. 81, H. biloxiae, new species: anterior part of
third-instar larva, lateral view of left side. 82, H. ischiaca Loew: posterior part of abdominal
segment 8 and tracheospiracular siphon of second-instar larva, dorsal view. 83, H. ainsworthi,
new species: left spiracular peritreme and ramus of third-instar larva, lateral view.
NUMBER 6 8 139
84
87
89
FIGURES 84-95.?Feeding apparatus of larval Hydrellia (third-instar and lateral view unless
otherwise specified): 84, H. notiphiloides Cresson: dorsal view; 85, H. bilobifera Cresson;
86, H. ascita Cresson; 87, H. tibialis Cresson: mouth-hook, lateral view; 88, H. trichaeta
Cresson; 89, H. bergi; 90, H. ainsworthi, new species: second instar; 91, H. bilobifera Cresson;
92, H. luctuosa Cresson; 93, H. spinicornis Cresson: second instar; 94, H. deceptor, new species;
95, H. notiphiloides Cresson.
140 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
96
101
103 010MM
104 105
FIGURES 96-106.?Feeding apparatus of larval Hydrellia (third instar and lateral view unless
otherwise specified) : 96, H. cruralis Coquillett; 97, H. discursa, new species; 98, H. caliginosa
Cresson; 99, H. ainsworthi, new species; 100, H. ischiaca Loew; 101, H. biloxiae, new species;
102, H. itascae, new species; 103, H. spinicornis Cresson; 104, H. morrisoni Cresson; 105, H.
spinicornis Cresson: first instar; 106, H. pulla Cresson.
NUMBER 6 8 141
- - H I S
107
109
0.5MM
110
108
O.SMM
112
O.SMM O.SMM 114
O.SMM
FIGURES 107-114.?Puparia of Hydrellia (ventral view; setae and creeping welts on middle
part omitted): 107, H. luctuosa Cresson; 108, H. cruralis Coquillett; 109, H. bilobifera
Cresson; 110, H. itascae, new species; 111, H. caliginosa Cresson; 112, H. ascita Cresson;
113, H. discursa, new species; 114, H. pv.Ua Cresson.
142 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
115
0.5MM
O.SMM
119
117 118
120 121 122
O.SMM
123 O.SMM
FIGURES 115-123.?Puparia of Hydrellia (ventral view; setae and creeping welts on middle
part omitted): 115, H. trichaeta Cresson; 116 H. notiphiloides Cresson; 117, H. spinicornis
Cresson; 118, H. tibialis Cresson; 119, H. biloxiae, new species; 120, H. ainsworthi, new species;
121, H. morrisoni Cresson; 122, H. ischiaca Loew; 123, H. bergi Cresson.
NUMBER 6 8 143
FIGURES 124?129.?124, Hydrellia spinicornis Cresson: late second-instar larva with opiine
Hymenoptera parasite, lateral view ( X 16). 125, H. discursa, new species: egg, lateral view
^X 48). 126, H. griseola (Fallen) and H. ischiaca Loew: left puparium with early pupa of
H. griseola; right one with late pupa of H. ischiaca, lateral view ( x 4). 127, H. discursa,
new species: posterior third of third-instar larva, ventrolateral view ( x 17). 128, H. cruralis
Coquillett: puparia on left void, two on right with hymenopterous parasites, dorsal and ventral
views ( x 4.5). 129, H. itascae, new species: puparium in situ in submergent Potamogeton
leaf, dorsal view ( X 5.2).
144 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
132
FIGURES 130-132.?130, Hydrellia griseola (Fallen): male with mite parasite, right lateral
view ( x 8.2). 131, H. ischiaca Loew: puparia, left one with braconid Hymenoptera parasite,
right one with pharate adult fly, dorsal view ( x 9.5). 132, H. cruralis Coquillett: puparium,
ventral view, and emerged adult hymcnopterous parasite, dorsal view ( X 6.2; photograph from
slide borrowed from C. O. Berg).
NUMBER 6 8 145
OR-
M3+Cu,
135
FIGURES 133-135.?133, Hydrellia griseola (Fallen): female internal genitalia, left view of
parasagittal section ( x 17). 134, H. griseola (Fallen), male: proboscis, uncleared, left lateral
view ( x 54). 135, H. griseola (Fallen), female: left wing, dorsal view ( x 18.7).
146 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
138
FIGURES 136-138.?136, Hydrellia griseola (Fallen), male: abdomen, ventral view ( x 34).
137, H. griseola (Fallen), male: abdomen, left lateral view (X 34). 138, H. bilobifera
Cresson, male: abdomen, left lateral view ( x 36.5), phallus partly depressed.
NUMBER 6 8 147
FIGURES 139-142.?139, Hydrellia griseola (Fal len) , male: head, front view ( x 32 ) . 140,
H. griseola (Fal len) , male: cibarium of probosis, front view ( X 32) . 141, H. griseola (Fal len) ,
male: cibarium, posterior view ( X 80) . 142, H. griseola (Fal len) , male: cibarium, right lateral
view ( X 80) .

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