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 biological 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 apparatus 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. Additional 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.

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WASHINGTON : 1971

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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 California'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 recorded the pest status of H. griseola. He reported

the fairly widespread damage of barley, oats, and

timothy grass by that species in southern and southeastern 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 distributed 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 objectives: (1) to describe, redescribe, and construct keys

to the adults and as many immature stages as possible 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 presented 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 Hennig's (1943) specific key to larval Hydrellia including

Palaearctic species contributed most to the taxonomy

of Palaearctic species of Hydrellia. The recent revision 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 sections on Nearctic Hydrellia. Very little additional

taxonomic study of the genus was made until 1915,

when Cresson started describing new species. Johannsen (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) contributed 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) included a key to California species and a Nearctic

generic key; Grigarick (1959) redescribed the lifecycle 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 Galapagos 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 Oriental 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 discuss 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) described 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 representatives including H. griseola. With phylogenetic

interpretations as a primary objective, Hennig made

an important contribution here, especially in proposing 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 terminalia of 17 species of Hydrellia from the Stenhammar 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) illustrated parts of the male and female genitalia for

three new species of Hydrellia in eastern Siberia.

Sturtevant (1925, 1926) made the only contributions 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 morphology of these structures in Hydrellia.

Several authors contributed morphological data

on the immatures of Hydrellia: von Frauenfeld

(1866) described the gross aspects of the metamorphosis 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 investigation 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 physiological 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 morphological 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 illustrating most immature instars of six species. Seguy

(1950) referred briefly to some aspects of morphological 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 immature 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 contributions: 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; Balachowsky 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 Biological 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 Biological 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 Station, 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 Natural History (AMNH), California Academy of

Sciences (CAS), University of California at Davis

(UCD), Canadian National Collection (CNC), Carnegie Museum, Chicago Natural History Museum,

Cornell University (CU), Illinois State Natural

History Survey, Iowa State University (ISC), University of Kansas (UKC), Kansas State University,

University of Massachusetts (UMM), University of

Michigan, University of Minnesota (UMC), University 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 Agriculture (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. crassipes 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 accommodated dowels run vertically through them and

into the lake bottom to anchor the device. A detergent solution in the pan constituted the actual trap

mechanism. The detergent destroyed the effectiveness 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, Collembola, Hymenoptera, and several dipterous families in addition to those of Hydrellia.

I made most of my behavioral observations while

collecting. After observing the behavior of the individual, I collected it with a killing tube or livecapture tube. I tried to keep some captured adults

alive in the laboratory long enough to make additional observations or for inseminated females to

oviposit.

PREPARATION OF ADULTS.—I killed and preserved

in 80 percent ethanol specimens intended for anatomical and food-habits studies. I used a hydration

series on these specimens before dissecting or sectioning them. I hydrated some specimens just before

dissection and some just after fixation in Bouin's

solution or dioxane in preparation for serial sectioning.

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 terminalia, 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 stereomicroscope. 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 overwintering 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 examinations, I would occasionally find a first- or secondinstar 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 species 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 temperature 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 thermograph 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 specimens 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 moistened 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 microvials and placed them on the pin holding the corresponding 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 processes, 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 encountered in a taxonomic study must be named. Since

zoological terminology is already burdened with

much ambiguity, I constructed the following definitions 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 perpendicular 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 Mt + 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 Mt + 2 junction and the distal edge of the r-m

and M1 + 2 junction. (Figure 135.)

Mesanepimeron (aem^) : Part of the pteropleuron of other

terminology. (Figure 6.)

Mesanepisternum (aes2): The mesopleuron of other terminology. (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 terminology. (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 distiphallus. 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 postgonite which is often hooklike or clawlike and usually distinctly 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 projections 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 inserted 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 distance 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 distance 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 frontoclypeus at the level of the anterior edge of the cheliform

spot divided by the dorsoventral extent of the frontoclypeus 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 extent 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 distance 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 phragmatal ramus. (Figure 85.)
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Puparial length: The distance between the anterior prothoracic 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 homologous 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 becomes isolated by a membranization of the surrounding 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 membranization of the clypeus and the boundaries of the

frons in explanations of the evolution of the dipterous cibarium and proboscis.

Snodgrass never explained the fate of the epistomal (frontoclypeal) sulcus, but since, by the

accepted definition, the epistomal sulcus is the external cuticular furrow, or groove, between the anterior tentorial pits which delimits clypeus and frons,

I must assume that in the Snodgrass view the anterior 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 question 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 investigation 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 subcranial margins dorsad along the primary facial rows

as faint internal cuticular thickenings. These thickenings 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 supraantennal 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 opening are then supposed to be homologous with the

anterior arms of the tentorium, while the thickenings stretching from the occipital foramen downwards (pt.) to the mouth opening are supposed to

be homologous with the posterior arms of the tentorium." This concept of continuous sulci from

dorsal tentorial pits through anterior tentorial pits

to posterior tentorial pits can be visualized in illustrations by Bonhag (1951, figs. 1-4). For Bolwig's

interpretation to obtain, the anterior tentorial pits

need have shifted posteroventrad only a short distance. 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 attempted 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 extensively used in chaetotaxy. The area superior to this

epistoma between the indistinct arms of the epistomal 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 postgenal 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 theoretically 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 convenient 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 postoccipital 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 subcranial 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 extremities 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 conjunctiva basal to the labrum are the labral apodemes

and the trachea-like siphon connecting the labial

gutter with the cibarium. The distiproboscis consists essentially of the labella. Each labellum is partially supported above by a labellar sclerite and consists 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 antagonistic muscles as in Nematocera and many Brachycera

or by inflections of the lateral borders of the anteclypeus, 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 specimens 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 functional 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 compressors 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 macrochaetae 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 conspicuous. In most species, smaller secondary facials

occur, usually lateral to the primary facial row.

According to Hennig (1958), the vibrissa is present but indistinguishable in Ephydridae. Also,

according to Hennig, the postocellar has replaced

the postvertical at least in prominence and approximate location.

THORAX.—My presentation of the gross external

morphology of the thorax proper of Hydrellia is

essentially an interpretative synthesis of the contributions 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 prothoracic 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 posterior 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 termination 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 (Figure 6), the wings, and the mid legs are the remaining

main constituents of the mesothorax.

The dorsopleural (or notopleural) sulcus separates the notopleuron and the mesopleuron. At the

anterior end of this sulcus is a slightly distinguishable 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 conspicuous 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 occupies 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 secondary 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 probably 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 distinguished 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 mesokatepisternum 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 metapleuron consists of a metathoracic precoxale, apparently 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 metepisternum, 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 modified 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., postpronotals, propleural, mesanepisternals, mesokatepisternal(s), and basicoxal(s). On the mesonotum,

the dorsocentrals and acrostichals anterior to the

incomplete transverse sulcus are termed predorsocentrals 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 prescutellar is one large interalar (corrected from "intraalar"

by Sturtevant and Wheeler, 1954). Above the mesopleural wing process are two small supra-alar setae.

There are two notopleural macrochaetae and one lateral macrochaeta just anterior to the notopleural

apex. Inserted on the scutellar margin are the basal

scutellar, intermediate scutellar, and the apical scutellar.

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 interfractural 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 postabdomen. According to Steyskal's (1957) hypothetical 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 constitute 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 Figures 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 situation 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 taxonomically, 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 intromittent structure the phallus, partly because I was

uncertain how much of it represented the true

aedeagus and partly because phallus was a convenient suffix (basiphallus and distiphallus). I

needed these terms because of the taxonomic importance of the common division of the phallus into a

somewhat bulbous, heavily sclerotized basal part

(basiphallus) and a usually narrower, lightly sclerotized distal part (distiphallus). The basiphallus may

be the actual phallobase and the distiphallus—the

aedeagus of Snodgrass (1935, 1959). Phase microscopy 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 continuous 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 posterior termination. In many species, the pregonites

appear to have a more definite articulation with the

phallapodeme. I have called the more heavily sclerotized, 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 analyzed 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 depresses, 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 posterior end of the phallapodeme against the inferior

condyle of the basiphallic socket and by the phallic

depressors and associated muscles relaxing synchronously. 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 ventrolateral 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 understandable, 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 ventral 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 proctodeum. 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 circumversion. 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 composed 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 described the spermathecae and accessory glands of

Hydrellia in his comparative morphological study of

these structures in Aschiza and Schizophora Acalyptratae. 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 surmise 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 subfusiform, or cucumiform, with fairly symmetric contours 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 observation of sperm entry or sperm tracing.

The nature of the globose, lacunose, pluglike structure 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 apparatus 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 illustrated the morphological nomenclature of the feeding 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 mouthhooks 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 posteriorly into dorsal and ventral phragmatal rami

between which the cibarial dilator muscles can be

seen stretching from the dorsal rami to the epipharyngeal 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 unillustrated observations of feeding mechanics and some

dissections, I hypothesize that the fossae of the cheliform spots serve as insertions for protractor muscles

of the feeding apparatus, as does the prominent

clypeal arch, or dorsal angle of Allen (1957a), illustrated 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 phragmata below the cheliform spots.

The primary factor underlying the reduction of
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the clypeal arch and probably the entire streamlining 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 degrees 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 eliminate 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 metapneustic, but Keilin (1915, 1944) demonstrated a

vestigial, closed prothoracic spiracular atrium. According to Hennig (1943, 1952) there appears to

be a trend toward functional metapneustism in the

Ephydridae. Developmentally, the caudal tracheospiracular 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 conical with a terminal aciculous spine, the spiracular

atrium narrow, and the secondary atrial orifice medially located. The spiracular peritreme is probably

nonretractile. On abdominal segment 8 are supraspiracular 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 protuberances and conspicuous spinous processes occur

in the second-instar larvae, these are lost at the

second molt.

According to Hennig (1943, 1952), the thirdinstar 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 illustrated 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 prominence 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 setulose 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 segment . . . ." 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 innervation 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), locomotory spinules by Hinton (1955), tubercles by Allen

(1957b), and spiculi by Nye (1958) and Grigarick

(1959). Because the term is more descriptively flexible 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. According 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 considerable 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 translucency 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 morphological phase. The changes involved in the formation

of this uniquely economic "cocoon" are physiologically complex. Keilin (1944) used the term pupariogenesis to describe this formation. An equally satisfactory 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 operculum, 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 schwarzen, 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 puparium, except in aquatic larvae where lime deposition

is substituted for this endocuticular sclerotization.

Because of their endophytic environment and perhaps 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 inappropriate. The puparial cuticle is sclerotized, but darkening seems to vary ecologically and perhaps genetically. Hering (1950) observed that the prevalent

translucency of Hydrellia puparia makes them some

of the most suitable of all Acalyptratae for investigating 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 sufficiently 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 (including Hydrellia griseola), Psyllidae, early-instar

Ephemeroptera and Odonata, Collembola, Braconidae, several species of nematocerous Diptera, Musca

autumnalis, and Sarcophaga sp. Adults of some species prey actively on Collembola. Yeasts and similar

fungi, Cyanophyta, Chrysophyta, nectars, and leaf

epidermis are other dietary items of adults. Several species of lycosid and ctenid spiders, species of

Ochthera (Ephydridae), and species of Lispe and

Hydromyza (Muscidae) prey upon adults of Hydrellia. 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 influence the ecological distribution of Hydrellia species.

Predominantly, these preferences center on species

of Potamogetonaceae and Gramineae. Other preferences include species of Alismataceae, Cyperaceae,

Hydrocharitaceae, Lemnaceae, and Liliaceae. Mines

of Hydrellia in dicotyledonous plants of Caryophyllaceae, Labiatae, Chenopodiaceae, Scrophulariaceae,

and Compositae apparently serve only for pupariation, 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. Nasturtium 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 competition with aquatic caterpillars, snails, phytophagous 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 ovipositing 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 intermedius 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 considerable 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 repeatedly 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 (Braconidae) 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 development of endoparasitic larvae, for in several observations 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 posteroventrally. 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 emerging adult hymenopteron for Ademon niger and
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Trichopria columbiana, at least, can remain submerged 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 laboratory 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 distribution 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 performed 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 zoogeographic consideration: (1) the known host preferences 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 tension of water and thus probably adversely affect

locomotion of adults and migrating larvae; (4) most

of the known species are Holarctic, the greatest number 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 operculum 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 species 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 periodically to clean their bodies and evert and then withdraw the ptilinum. The wings remain folded until

about 15-30 minutes after emergence, when they

rapidly expand. Often, they cannot fly until 45 minutes after emergence.

FEEDING.—Adults often search over the surface

film for entrapped insects and sometimes mistakenly

pounce on inanimate floating objects and manipulate 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 carnassially 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 peculiar behavior while feeding, e.g., H. biloxiae rhythmically 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 distinctive 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) dismounting, 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 Hydrellia, 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 mechanism. 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 predominant silvery or yellow facial pruinosity in

Hydrellia (and other schizophorans) reflect considerable 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 sometimes fail, as in cases where a male H. nobilis tried

to mount a male H. ischiaca after repeatedly scissoring his wings, or a male H. nobilis tried to mount a

female H. ischiaca, or a female H. amnicola attempted 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 several other conditions. A pair of H. ischiaca copulated

for 10 minutes and a pair of H. nobilis for 19 minutes. Hydrellia griseola have copulated from 1 to 50

minutes and at various times during daylight between 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 laboratory, 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 incubation, the pharate larvae often begin to make slight

movements several hours before eclosion. They slit

the micropylar end of the chorion with their mouthhook 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 crawling onto the abaxial surface. Larvae locomote on the

surface by anchoring their mouth-hook and contracting the body segments and by pushing with

their spiracular peritremes and creeping welts. Partial desiccation stimulates larvae, especially newly

eclosed ones, to twist and roll. This emergency reaction 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 perpendicularly 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 extremely thin mesophyll. By continuing this excavation 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 peristalsis. Also, even late third-instar larvae can turn

about within the mine. During endophyllous locomotion, 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 spiracular 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, because 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. Pupariation 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 puparium 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. Thirdinstar 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 spiracular peritremes embedded only in the epidermis, 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 Hydromyza 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 temperature 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 fecundity.
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. aurifacies 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 characters: 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, ventrally directed, secondary facial setula inserted above

primary facial row; two fronto-orbitals (anteriorly

and posteriorly inclined) ; apicodorsal antennals not

projecting noticeably; three scutellars (basal, intermediate, 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: metapneustic; spiracular peritreme spinous; mouth-hook

single.

Cresson (1944b) erected the tribe Hydrelliini

solely for Hydrellia and indicated its close relationship to the tribes Hydrinini and Ilytheini. As characters for Hydrelliini, he listed essentially the generic

characters. No author has disputed this classification.

However, some controversy has existed on the subfamilial classification. Loew (1860) placed Hydrellia

with Atissa, Philygria (as Hydrina), Hyadina, and

Axysta in Hydrellina, which corresponded to a subfamily. Becker (1926) grouped Hydrellia, Philygria,

and several parydrine genera in his subfamily "Hydrellinae." 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 immature instars of twelve species and redescribed

those of seven species previously known.

The phylogeny of this genus remains obscure concerning 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 distinguished 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 constituents 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 basiphallus) 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 character 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 identical in both groupings. Another computer group

correlated fairly well with two habitus groups—

morrisoni and platygastra. A computer program

integrating all quantitative and qualitative characters 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) ainsworthi.

crassipes group: (31) crassipes, (32) saltator, (33) procteri, (34) advenae.

nobilis group: (35) nobilis, (36) idolator, (37) atroglauca, (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) columbata, (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 terminalia 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 reference 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 II: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 II: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 II: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

III:IV 2.5 or less. (Male terminalia as in Figure 19.) H. caliginosa Cresson

Parafrontalia not velvety black; 6-8 aristal rays; costal section III: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 II: 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 II: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 II:I usually

2.4 or less 23

Prementum light gray; apicodorsal antennal prominent; costal section II:I 2.4 or more.

(Male unknown.) H. lata Cresson

23. Parafrontalia velvety black, contrasting with frontal vitta; 2 or 3 basal coxals; mesanepisternum 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 II: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 II: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; M1 + 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 mm; female cercus acute apically, rounded basally (Figure 71).

(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 rectangular 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 prothoracic 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; parafrontale 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 moderateyellow 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: II: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 concave; 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. Postgonite 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; mesonotum strong yellowish brown; thoracic pleuron light

gray except upper half of mesanepistemum semiglossy 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 olivebrown; 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: II: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; anterolateral margin of sternum 5 rounded; copulobus regularly 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 delimited 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. Vockeroth), 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; antecosta 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 frontoorbitals, 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 darkbrown 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; costalsection 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 medially, 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, ventrally 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; maximum 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 translucent, 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; maximum breadth: minimum breadth 4.0-4.5; anal-plate

index 2.3-2.6. Prothoracic end semicircular in ventral view; head-lobe scar subcircular or ovoid; maximum 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 localities :

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 (IV5-1953, 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), 13 , 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. Deonier); 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 caroliniana, Hypericum punctatum, Potamogeton diversifolius, 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 September, 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 frontoorbital 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 lightbrown 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 lightbrown 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: II:I 2.0-2.5; III: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 seriated 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),

299 .

Maryland: Chesapeake Beach (VIII-18-1922, J. Malloch), \$, (VII-3-1924, J. Malloch), 19 ; Kent Narrows

(VII-1954, M. Wheeler), 19 .

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 unknown), 19 ; Bellefontaine Point, Jackson County (VII10-1962, D. L. Deonier), 19 ; Dickinson's Pond, Lamar

County (VII-24-1962, D. L. Deonier), 19 ; Gulf Coast

Research Laboratory, Ocean Springs (VI-5-1962 , D. L.

Deonier), 1 <$ ; Vancleave Road, Jackson County, 30°25.1'

N, 88°46' W (VI-23-1962, D. L. Deonier), 13 .

New York: Cold Spring Harbor, Long Island (VH-dat e

and collector unknown), 19 ; Wilmington Notch, Adirondacks (VII-3-no year, J. Aldrich), 19 .

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; frontoorbital area moderate olive brown; 13—17 postoculars.

THORAX.—Ppn and mesonotum dark brown; 3-4

adc and 2 pdc; pleuron light gray except upper onesixth 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; costalsection 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 concave; 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 directed 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 collection 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 taxonomy 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 segment 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 sternum 5 with dorsal process projecting laterad (forming 90°-100° angle with copulobus); posterior margin of copulobus only slightly convex (partly or

wholly obscured by loose fascicula of 5-7 strong

macrochaetae) ; medial margin of copulobus distinctly 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 truncate 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; bifurcation index 4.2-4.4; clypeal-arch index 1.5-1.8.

Clypeal arch sloping at 10°-15°, with slight indentation 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 segment 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; maximum 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 subcircular, with maximum breadth about half of

prothorax length; anal plate subovoid, with distinctly

convex anterior margin. Empty puparium translucent, light yellowish brown. Early pupa with

greenish-yellow tinge.

TYPE.—Holotype male, ANSP 6621.

TYPE-LOCALITY.—Nigger Creek, Cheboygan County, Michigan (VIII-21-1941, C. O. Berg; reared

from Potamogeton tenuifolius).

ADULT SPECIMENS EXAMINED.—32 (HcfcT,

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), 19 ; 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), 19 , (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), 19 ;

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), 3tt ,

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 (collected and reared by C. Berg).

REMARKS.—As with many other species, especially

those of the H. bilobifera group, there is some possibility 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 medially, 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; epistomal 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: II:I 2.2-

2.5; III:IV 2.7-3.1; V:IV 3.0-3.8; Mi • 2 index

1.4-1.6.

ABDOMEN.—Terga semiglossy moderate brown medially and with conspicuous light blue-gray posterolateral 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 prementum, 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 immatures].—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: II:I 2.0-2.3; III: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; distiphallus long, asymmetrical, with apex curved to

right. Anterior margin of fused surstyli slightly convex; 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. General 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 thirdinstar 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 frontoclypeal 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; maximum 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 distinctly 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), 19 , (VII-18-1946, C. Berg), \$, (VIII-18 -

1946, C. Berg), 19 , (VII-21-1947 , C. Berg),

1 9 , (VIII-7-1947, C. Berg), 19 ; Douglas Lake,

Cheboygan County (VI1-5-1940, C. Berg), 13 , (VI16-1941, 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), 19 ,

(VIII-1-1941, C. Berg), 2 $ $ ; Third Sister Lake, Washtenaw County (VI-28-1942, C. Berg), 19 -

Minnesota: Eaglenest (IX-1-1958, W. Balduf), 19 ;

Iron Corner Lake, Itasca State Park (VI-18-1963, D. L.

Deonier), 19 ; Mississippi River at Sucker Creek, Clearwater County (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), 14 , 3 9 9 ;

Squaw Lake, Itasca State Park (VIII-14-1963, D. L. Deonier), 5$ $, 3 9 9 ; Rapid River Logging Camp, Hubbard

County (VII-2-1963, D. L. Deonier), 19 ; 1.3 miles south

of main entrance, Itasca State Park (VI-26-1963 , D. L.

Deonier) 9 3 4, 7 9 9 .

New York: Dryden Lake, Tompkins County (VIII-8 -

1961, D. L. Deonier), \1 $ &, 199 9 ; Franklinton (VI20-1923, M.Leonard), 2 9 9 .

Ontario: London (IX-12-1958, W. Judd), 19 ; Marmora (IX-9-1952, J. McAlpine), 19 ; Ottawa (VII-2 -

1947, G. Shewell), \$.

Quebec: Lac Bernard (VIII-7-1938, G. Shewell), 19 ;

Mt. Albert (VII-24-1954, J. Martin), 2 9 9 .

IMMATUR E SPECIMEN S EXAMINED. — 3 2 from 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 observation, 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 retreated 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 meeting. 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 covering 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 floating 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; apicodorsal 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, yellowish 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 antennal 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; anterolateral margin of sternum 5 rounded to a posteriorly

directed notch; posterior third of copulobus distinctly 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 terminalia: 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; maximum breadth 0.10-0.15 mm. Clypeal arch distinctly 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 dorsolateral seta (perhaps supraspiracular spinous seta of

first-instar larva). Body translucent, with light-green

tinge.

THIRD-INSTAR LARVA.—Length 2.17-5.85 mm; maximum 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. Clypeal arch gradually sloping at about 35° in relation 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; maximum breadth: minimum breadth 2.4-3.2; anal-plate

index 1.8-2.4. Prothoracic end semicircular in ventral 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), 13 , (X-21-1954, A.

Grigarick), 33 3 , 19, (VII-26-1955, W. Lange), 13 ,

(IX-10-1957, A. Grigarick), 2 9 9, (IX-17-1957, A. Grigarick), 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), 13 .

Iowa: Banner Mine Area, Warren County, 41°26.4' N,

93°33.8' W (VIII-7-1960, D. L. Deonier), 13 ; 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, Hamilton County (VIII-27-1960, D. L. Deonier), 2 3 3 , 19 ;

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), 13 , (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), 13 , 29 9 ; Leavenworth 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), 13 ; 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) 19 , (XI-26-1946, C. Berg), 23 3 , (VIII-11-1947,

C. Berg), 13 .

Minnesota: Biological Station, Itasca State Park (VI26-1963), D. L. Deonier), 13 ; 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), 13 ; 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), 13 , (VIII-8-1963, D. L. Deonier), 13 ;

6.5 miles east of Waubun (VIII-18-1963, D. L. Deonier),

599 .
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 ; Miller's Camp, Reelfoot Lake, 36°24.3' N, 89°20' W (VIII12-1962, D. L. Deonier), 15 5 5, 152 9 ; Samburg, Reelfoot 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, (IX13-1953, M. Wheeler), 15 ; Kerrville (111-17-1955, W.

Wirth), 15.

IMMATURE SPECIMENS EXAMINED.—296 from 7

localities:

Iowa: Siewer's Springs State Park near Decorah (collected 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 misdetermined 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 Reservation, 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 consisted 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 collection. 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 Zannichellia palustris in Davis, California, during his research 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, Jussiaea repens, Nasturtium officinale, Eleocharis palustris, 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 commonly 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. (probably 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 34,81 , 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 moderate 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; costalsection ratios: 11:1 1.4-2.2; III: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 irregularly setose. Postgonite bent anteriad and postgonite 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 frontoclypeal 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; maximum breadth: minimum breadth 3.6-4.2; anal-plate

index 1.8-3.2. Prothoracic end semicircular in ventral 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 ventral, transverse row of setulae, but with 3-4 annuli

of spinules; anal plate subovoid, with anterior margin 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 localities:

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, 49 $ (USNM and DLD); Lake

Shady, 7 miles west of Hattiesburg, Lamar County (VII11-1962, 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 Nymphaea 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 specimen pushed its body forward and upward rhythmically. 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 discovery 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 remained 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 setaceum, 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 (infrequently partly moderate orange-yellow); 7-8 aristal

rays; apicodorsal antennal prominent; antenna and

frontal vitta brown; most of parafrontale olivebrown; 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; costalsection 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 Philadelphia 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. Dreisbach), 1$.

Nebraska: Crete (VII-3-1960, W. Rapp), 1$.

Ontario: Ottawa (VI-1-1951, J. McAlpine), 1 $, (VII14-1952, G. Shewell), 1$, (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 III: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 parafrontale 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 interfractural costals; costal-section ratios: II:I 2.2-2.6;

III: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 rightangled (90°-100°); posterolateral angle of

copulobus roundly 45° from the lateral corner; the

medial margin broadly notched; copulobus irregularly 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 examined. Frontoclypeal length 0.46-0.59 mm (Figure

98). Ventral frontoclypeal index 5.0-5.5; phragmatal 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 tapering 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. Melander), 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), 13 ;

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 unknown), 16, 19 ; 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 species 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 index 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 olivebrown; 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:

II:I 2.0-2.5; III: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 acutangular 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 holotype 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 moderate 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 Figure 59.

HEAD.—Face light gray or light yellowish brown;

3-6 pfa; 0-5 sfa; epistomal index 1.0-1.4; mesofacial 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 mesomeron light gray; 1-2 basal coxals (1 macrochaetous); 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; III: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 concave; 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 VI4-1967, D. L. Deonier), \$, 19 (DLD); Rapid River

Logging Camp, Hubbard County (VII-2-1963, D. L. Deonier), 2$ (DLD); Sucker Lake at dam (reared adult V27-1967, 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 submerged 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 thirdinstar, 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 temperatures 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. zosteriformis, 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 olivebrown; 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 except 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 distinctly 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 localities:

Maine: Mount Desert Island (VIII-12-1935, W. Procter)^? 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 immatures of this species I examined Potamogeton

amplifolius, P. natans, P. gramineus, P. robbinsii,

Zannichellia palustris, Eleocharis palustris, Sparganium 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 Figure 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: II:I 2.2-

2.6; III:IV 2.8-3.2; V:IV 3.5-4.0; M, • 2 index

1.2-1.8.

ABDOMEN.—Terga semiglossy dark brown medially, light gray laterally and ventrally. Male terminalia : median third of posterior margin of sternum

5 deeply concave; anterolateral margin of sternum 5

rounded through angle of 100°; copulobus acutangular (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, 879? )

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), 15 , (VIII-13-1933, G. Steyskal), 19.

Minnesota: Basswood Lake, Lake County (VIII-16-1950,

R. Namba), 1 9 ; Chambers' Creek, Itasca State Park (VII18-1963, D. L. Deonier), 453,299 ; Douglas Lodge Bay,

Itasca State Park (VI-20-1963, D. L. Deonier), 1 9 ; Mississippi 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), 15 , 19-

New Jersey: Manahawkin (VIII—1—1912, collector unknown), 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, collector unknown), 85 5 , 59 9 ; (VII-14-1903, collector

unknown), 2 5 5,2 9 9.

Ontario: Bark Lake (VII-22-1939, D. Bullock), 19 ;

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), 15 , 69 9, (VIII-25 and IX-8-1952, E.

Smith), 39 9.

Oregon: Oregon City (IX-no date and year, collector unknown), 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 aggregation 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 aggregation. 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 captivity 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 information 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 localities 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 taxonomy 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; mesofacial 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 parafrontale velvety dark brown; 12-16 postoculars.

THORAX.—Ppn light gray; mesonotum strong yellowish 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:

II:I 1.8-2.2; III: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 squarenotched and congruent with distiphallus; anterolateral 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 undulate than in H. ischiaca. Micropylar protuberance

infundibulate and visible in dorsal view. This description 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 conspicuous. 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 conspicuous. 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; bifurcation index 2.4-2.7; clypeal-arch index 1.8-2.3.

Clypeal arch smoothly sloping at about 20° in relation to lower frontoclypeal margin. Mouth-hook beak

distinctly longer than base; maximum mouth-hook

base thickness about 1.5 times that of beak; mouthhook light spot large and ovoid. Prothorax and mesothorax 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 ventral 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 unknown), 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), 19 ; Plummer's

Island, Potomac River (IX-1-1962, K. Krombein), 10 3 3 ,

799 .

Massachusetts: Fall River (VII-3-1930, A. Melander),

1 9.

Michigan: Black River, Cheboygan County (VI-20-

1941, C. Berg), 833 , 899 ; Carp River, Cheboygan

County (VII-1-1941, C. Berg), 59 9 ; Cheboygan Pool,

Cheboygan County (VIII-6-1941, C. Berg), 13 , 19 ;

Cheboygan County (VI-12-1940, C. Berg), 13 , (VIII14-1940, C. Berg), 13 , (VIII-6-1941, C. Berg), 13 , (VI26-1946, C. Berg), 3 9 9, (VII-5-1946, C. Berg), 2 3 3 ,

1 9 , (VII-16-1946, C. Berg), 13 , (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), 13 , 19, (VII3-17-1941, C. Berg), 39 9, (VI-26-1946, C. Berg), 19 ,

Grosse Isle, Wayne County (IX-12-1948, G. Steyskal),

1 9 ; Hamburg, Livingston County (VIII-13-1933, G. Steyskal), 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), 19 , (VI-6-1941, C. Berg), 13 , (VI-29-

1941, C. Berg), 33 3 , 109 9 ; Midland County (IX-5-

1937, G. Steyskal), 19 ; Monroe (VII-2-1939, G. Steyskal), 19 ; Ocqueoc Lake, Presque Isle County (VII-13-

1941, C. Berg), 2 3 3,299 , (VII-15-1941, C. Berg), 13 ;

Third Sister Lake, Washtenaw County (all reared by C.

Berg: 111-15-30-1940, V-l 1-20-1940, VI-29-1940, VII2-1940, 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 ; Whitmore 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 (VIII6-1963, 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), 19 ; Squaw Lake,

Itasca State Park, (VIII-14-1963, D. L. Deonier), 4 3 3 ,

49 9.

New Jersey: Trenton (VII-21-1909, collector unknown),

299 .

Ohio: Cedar Point, Sandusky River (VIII-5-1902, collector unknown), 1 3 •

Ontario: Arnprior (VIII-19-22-1930, C. Curran), 13 ,

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, VII8-30-1958, VIII-14-31-1958, IX-2-27-1958, and X-9-

1958), 1013 3 , 1869 9 ; Marmora (VII-21-1952, J. McAlpine), 23 3 , 19, (VIII-4-1952, J. McAlpine), 19,

(VIII-7-1952, J. McAlpine), 19 , (VIII-11-1952, J. McAlpine), 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), 13 ,

(IV-27-1942, A. Melander), 19 ; Garner State Park,

Uvalde County (IV-3-1955, W. Wirth), 13 ; Pedernales

River, Gillespies County (111-19-1955, W. Wirth), 49 9.

Virginia: Alexandria (VIII-5-1922, J. Aldrich), 13 .

West Virginia: Marlington (VIII-15-1930, G. Auxier),

1 3 , 49 9.

IMMATURE SPECIMENS EXAMINED: 60 from 6 localities :

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 (collected and reared by D. L. Deonier) ; Long Lake, Clearwater 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 coordination 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 specimens approached each other while holding their

wings about 45° from their abdomina. They suddenly 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 singlelayered 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, respectively, 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 overwintering 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 thirdinstar 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 dorsal 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 mesonotum 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 lightgray 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; costalsection ratios: II:I 2.2-2.6; III:IV 2.8; V:IV 3.2-

3.6; Mi + 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), 19 ; 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: II:I 2.2-

2.5; III: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 concave; anterolateral margin of sternum 5 acutangular

in form of blunt hook; copulobus acuminate posteriorly and irregularly setose. Postgonite and postgonite uncus bent slightly laterad; distiphallus ventrally 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 (IX20-1957, 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 terminalia to confirm determinations.

Host plants. Grigarick reared a few specimens

from Sagittaria sp. collected near Sacramento, California, 20 and 21 September, 1957 during his research 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 mesokatepisternum, 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; III: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 (VIII31-1961, 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 (VIII18-1963, 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 unknown), 19 ; 12 miles northwest of Lusk (VH-no date1895, 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, Tompkins 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 distinct bare space posteriad from indistinct central

verruca. Postgonite bent laterad then mediad; posgonite uncus spicate, directed anteromediad; distiphallus slightly constricted in middle third and

indistinctly carinate and lamellate ventrally to

mucronate apex; most of basiphallus exposed ventrally. 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 longitudinal 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 protuberance 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 frontoclypeal 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. Mouthhook 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; maximum breadth: minimum breadth 4.2-5.0; anal-plate

index 3.5-4.0. Prothoracic end semicircular or subtriangular 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 translucent, light yellowish brown. Pupal color as in H.

bilobifera.

TYPE.—Holotype male, USNM 70532.

TYPE-LOCALITY.—Samburg, Reelfoot Lake, Tennessee, 36°22.9' N, 89°21.3' W (VIII-11-1962, D.

L. Deonier).

PARATYPES.—112 (48c" d1, 64? ? ) from 24 localities :

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), 13 , 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 (VI26-1963, 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 identified the others from other Potamogeton species as a

variety of H. ascita." Since I discovered several specimens 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 aquatica, 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, Sagittaria 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 undetermined 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; costalsection ratios: II:I 2.5-3.2; III:IV 2.6-3.0; V:IV

3.5-3.9; M1 + 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, Colorado (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 mountain 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 olivebrown; 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 olivebrown; coxae dark gray and tarsi dark brown; remainder 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: II:I 2.0-2.4; III: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; distiphallus lamellate ventrally and ventrally bicarinate

preapically. Anterior margin of fused surstyli undulate (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 localities :

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 distinguished 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 moderate 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; remainder of pleuron velvety brownish black; legs

light-gray pruinose except moderate-yellow trochanters, tarsi, and at least distal third of mid and hind

tibiae. Wing length 1.36-1.96 mm; veins light yellowish brown; 4-7 setae on basal end of costa; 5-7

dorsal and 5-9 anterior interfractural costals; costalsection ratios: II:I 2.4-2.7; III:IV 3.4-4.2; V:IV

3.4-4.0; Mi . 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 sternum 5 roundly obtusangular; copulobus acutangular

posteriorly (rounded 75°) ; posterior half of copulobus 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), 13 , (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), 19 ; Washington (VIII-17-1913, J. Aldrich),

5 9 9, (VIII-16-1952, J. Laffoon), IS.

Florida: Baxter (IX-no date-1954, M. Wheeler), 19 ;

DeFuniak Springs, Walton County (VIII-1-1962, D. L.

Deonier), 19 ; 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), 19 ,

(VII-4-1916, J. Aldrich), 2$$, 29 9, (VI-25-1916, J.

Aldrich), 23 3, 29 9, (VI-29-1916, J. Aldrich), 19 ,

(VII-7-1916, J. Aldrich), 19 , (VIII-31-no year, J.

Aldrich), 19 ; Lake County (VII-25-1950, W. Kwolek),

1 9 , (IX-15-1950, W. Kwolek), 19 : 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-southeast 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 ; Chesapeake Beach (VII-3-1924, J. Malloch), 19 ; (VIH-no

date-1926, J. Aldrich), 19 ; 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), 19 , (IX-1-1913, W.

McAtee), 19 ; near Plummer's Island, Potomac River

(V-17-1914, R. Shannon), 19 , (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), 13 ; Woods Hole

(VII-22-no year, A. Sturtevant), 19.

Michigan: Detroit (VI-19-1938, G. Steyskal), 19 ,

(IX-6-1942, G. Steyskal), 15 , (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 (VII10-1962, D. L. Deonier), 19 ; Bluff Creek, Vancleave,

Jackson County (VI-18-1962, D. L. Deonier), 19 ;

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), 19 ; Lake Shady, Lamar County (VII11-1962, D. L. Deonier), 5 5 3 , 13 9 9, (VI-29-1962,

D. L. Deonier), 3 3 5,399 ; 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), 19 ,

(IX-27-1915, C. Adams), 19 , (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 , 19 ; Cold Spring Harbor, Long Island (VII—no dates.

A. Melander), 23 3 ; Ithaca, campus (IX-24-1925, P

Babing), 19 ; Ithaca (VI—no dates, collector unknown)

1 9 ; Lancaster (VII-25-1946, L. Beamer), 13 , (VII25-1946, 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), 13 ,

1 9 , (V-28-1957, J. Vockeroth), 29 9 ; Highlands, 3,800

feet (V-8-1957, J. Vockeroth), 19, (V-10-1957, J.

Vockeroth), 13 , (V-13-1957, J. Vockeroth), 19, (VII23-1957, J. Vockeroth), 19 ; Highlands, Macon County,

35°8.2' N, 83° 11.3' W, 3,850 feet (VI-28-1958, J.

Laffoon), 19 .

Nova Scotia: Truro (no dates-1913, R. Matheson), 13 .

Ohio: Rome (VII-19-1946, R. Beamer), 13 .

Ontario: Ottawa (VIII-31-1908, J. Fletcher), 19,

(VI-3-1958, J. Vockeroth), 13 , 5 9 9 , (VIII-26-1959,

J. Vockeroth), 13 .

Pennsylvania: Alleghany County (VII-10-no year, H.

Smith), 2 3 3 ; Harmarville, Alleghany County (VI1-2 3-

1916, H. Kahl), 13 , 29 9 ; Hartstown, Crawford County

(VII-4-1921, H. Kahl), 19 ; Hills Station (VIII-30-1905,

H. Kahl), 19 ; Lansdalc (VII-3-1910, E. Cresson, Jr.),

2 3 3 , 19 ; Ohiopyle, Fayette County (VII-26-1905, H.

Kahl), 13 , (VII-28-1905, H. Kahl), 13 , 49 9, (VIII7-1905, 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 , 3399 ;

Pittsburgh (VIII-29-1908, H. Kahl), 2 3 5, (VI-28-1909,

H. Kahl), 19 ; Point Pleasant (V-30-1914, E. Cresson,

Jr.), 15 ; Swarthmore (V-13-1906, E. Cresson, Jr.), 19 ,

(VII-1-1908, E. Cresson, Jr.), 15 , 69 9 , (VI-20, VII-18,

VIII-22-1909, E. Cresson, Jr.), 8 9 9, (V-l, VII-31-1910,

E. Cresson, Jr.), 15 , 39 9 , (V-16, VI-27-1920, E. Cresson, Jr.), 2 3 3 , 29 9 ; Westmoreland County (no dates,

H. Kahl), 13 , 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), 19 ;

Knoxville (VIII-18-1939, D. Hardy), 19, (IV-16-no

year, collector unknown), 19 ; Reelfoot State Park, Reelfoot 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), 13 , 49 9 ;

Palmetto State Park, near Ottine (VII-10-1950, M.

Wheeler), 19-

Virginia: Alexandria (VI-14-1952, W. Wirth), 19 ;

Falls Church (VII-4-1950, W. Wirth), 19 , (VIII-8-

1950, W. Wirth), 19 , (VI-30-1951, W. Wirth), 19 ;

(VII-22-1951, W. Wirth), 19, (VI-9-1954, W. Wirth),

19, (VII-15-1954, W. Wirth), 15 , 19 , (VH-no date1954, M. Wheeler), 19 , (VI-30-1960, W. Wirth), 19,

(VII-24-1960, W. Wirth), 19 , (VIII-6-1960, W. Wirth),

19, (VIII-8-1960, W. Wirth), 15 ; Great Falls (VII21-1962, G. Steyskal), 19 ; Holmes Run, Falls Church

(VI-20-1960, W. Wirth), 13 , (VII-1-1960, W. Wirth),

19, (VII-3-1960, W. Wirth), 19 , (VII-31-1960, W.

Wirth), 19 ; Maywood, Alexandria County (VII-9-

1922, M. McAtee), 19 ; 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), 13 .

REMARKS.—This species is the most readily distinguishable 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 species 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 posteriorly 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 index 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; anterolateral 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 digitiform lobe projecting posteriad medial to postgonite)

and serially setulose; remainder of copulobus irregularly 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 margin 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 semicircular 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

II: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 olivebrown; 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 darkbrown 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: II:I 2.6-2.8; III: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; copulobus 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 supraspiracular asteriform process with 4-7 spinous projections (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 relation 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 asteriform 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 margin straight or slightly concave. Empty puparium

translucent, light yellowish brown.

TYPES.—Notiphila griseola Fallen: Fallen apparently 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). TYPELOCALITY: 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: RobineauDesvoidy very probably used only syntypes for this

binomen. Most of them are probably in the Paris

Museum (Natural History). TYPE-LOCALITY: probably 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 collected 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, Clearwater 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 species 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 determined by Mik, Schiner, Loew, and Becker. The variations 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, strawberries, 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 altitude 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 probably 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 composition 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 supplied with food, 50 percent died after 34 days.

Considering these and other factors, the mean ecological 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 species of Braconidae (Hymenoptera), and the following 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 specimens were greenish yellow and granular. Examination of sections of 125 adults revealed granular contents, 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 defensive 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 important predators of adults. He witnessed several captures by these spiders. In June and July, he caught

more lycosid spiders than H. griseola adults in floating 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 attempted to mate with the female, first grasping her

head, then her wing, and finally grasping the copulating male. When the third one intruded, the copulating 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 laboratory, 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 aggregations 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 microenvironmental 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 inseminated 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 stadium 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 figures 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 exposed 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, especially 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 following 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 thirdinstar larvae often sought a new mine for pupariation. 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 gramineous genera reported as containing host plants for

this species. Additionally, various authors have reported 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 infestations 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 following four conditions that increase susceptibility to

infestation of rice with H. griseola: (1) low soil fertility 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 species of Polypogon, Avena, Hordeum, and other wild

grasses in rain pools and irrigated rice fields and

three generations in May-June on rice and associated 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 infested rice plants only rarely after the middle of July.

Parasites. Forty-three species of Hymenoptera reported 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 highest 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 prominent. 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 lightgray 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, rectangularly 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 constricted proximally and preapically and bicarinate

ventrally; carinae partly lamellate; most of basiphallus 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 inconspicuously bilobate posteriorly. Female terminalia:

tergum 8 much longer than sternum 8; cercus irregularly 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 margin; 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), 13 .

Iowa: Spring Lake, Greene County (IX-19-1961, D. L.

Deonier), 13 , 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), 13 ;

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 prominent; 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: II:I 2.0-

2.5; III:IV 2.8-3.2; V:IV 3.3-3.8; M, *2 index

1.3-1.7.

ABDOMEN.—Terga semiglossy dark brown medially, 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 distiphallus; 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 (VII2-1947, G.E. Shewell).

PARATYPES.—4 (1 d\ 3 ? $ ) from 4 localities:

Mississippi: Bellefontaine Road, Jackson County (VI12-1962, 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, conspecificity 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 orangeyellow; prementum glossy brownish black; 8-10

aristal rays; antenna velvety dark brown; frontoorbital area and frontal vitta moderate olive-brown;

most of parafrontale dark brown; 13-17 postoculars.

THORAX.—Ppn and postalar bridge densely moderate 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 interfractural 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, shallow 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 listing]—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 except 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 mesonotum, and notopleuron light gray; remainder of

mesonotum moderate brown; 3-4 adc and 2 pdc;

pleuron light gray except upper fourth of mesanepistemum 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 olivebrown; 6-8 setae on basal end of costa; 6-8 dorsal

and 7-12 anterior interfractural costals; costalsection ratios: II:I 2.7-3.0; III:IV 2.1-2.6; V:IV

3.0-3.9; Mi . 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 posteromedial and medial margins, otherwise bare. Postgonite 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, corrugate, with longitudinal ridges occasionally anastomosing 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 distinctly 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; bifurcation 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 thickness about 2.0 times that of beak. Supraspiracular

asteriform process absent; prothorax and mesothorax

densely spinulose; abdominal segment 8 with ventral, 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; maximum breadth: minimum breadth 4.0-6.0; 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 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, 19 ; 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), 19 , (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), (VII10-1960, 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 Waterville, 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, 19 ;

Beltsville (V-21-1922, J. Malloch), 19 ; Glen Echo (VII20-1924, J. Malloch), 19 ; Plummer's Island, Potomac
NUMBER 6 8 73

River (VIII-5-1913, R. Shannon), 19 , (IX-5-1915, W.

McAtee), 19 ; near Plummer's Island, Potomac River

(V-22-1915, R. Shannon), 19 , (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), 19 ;

Roscommon County (VII-1-1950, R. Dreisbach), 19 ;

Vineyard Lake, Jackson County (VII-26-1954, G. Steyskal),

It.

Minnesota: Biological Station, Itasca State Park (VI26-1963, D. L. Deonier), 3$ 3 , 7 9 9 ; Bohall Lake, Itasca

State Park (VIII-17-1963, D. L. Deonier), 13 ; 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, Hubbard County (VIII-8-1963, D. L. Deonier), 13 , 19 ; Iron

Corner Lake, Itasca State Park (VI-18-1963, D. L.

Deonier), 13,299 ; Mississippi River, sec. 34, T. 145 north,

R. 36 west, Clearwater County (VII-8-1963, D. L. Deonier)

1 3 , 19 ; 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), 13,499 ; Mississippi River

at Sucker Creek, Clearwater County (VII-9-1963, D. L.

Deonier), 13,899 ; 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. Deonier), 13 , 19 ; 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 Biological Station, Lake Itasca, Itasca State Park (VII-28-

1963, D. L. Deonier), 13,399 -

New Hampshire: Mount Washington (no dates—1917, A.

Slosson), 19 .

New York: Dryden Lake, Tompkins County (VIII-8-

1961, D. L. Deonier), 19 ; Ithaca (VI11-28-1894, collector unknown), 13 ; 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. Deonier), 13 , 19 .

North Carolina: Clingsman Dome, Great Smoky Mountains National Park, 6,300-6,800 feet (V-28-1957, J. Vockeroth), 1 9 ; Forney Ridge, Great Smoky Mountains National

Park (VI-26-1941, A. Melander), 19 ; Highlands, Macon

County, 3,800 feet (V-10-1957, J. Vockeroth), 2 9 9 , (VI3-1957, J. Vockeroth), 13 ; Highlands, Macon County,

3,000 feet (VI-1-1957, W. Mason), 2 3 3 , 19 ; Looking

Glass Peak, Pisgah National Forest (VII-19-1957, W. Richards), 1 3 ; Wayah Gap, Macon County, 4,000 feet (VIII16-1957, J. Chillcott), 1 3 ; Wilson's Gap, Highlands, 3,100

feet (V-12-1957, J. Vockeroth), 13 , (V-25-1957, J.

Vockeroth), 13 , 19 ; 3 miles southwest of Glenville, Jackson 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), 13 ; Marmora (VII-5-1951, J. McAlpine), 19 , (VIII-4-1952, J.

McAlpine), 19 ; Normandale, 42°42' N, 80°19' W (V27-1956, J. Vockeroth), 13 ; Ottawa (VIII-8-1923, C.

Curran), 1 9 .

Pennsylvania: Alleghany County (VH-no dates, H.

Smith), 13 ; Castle Rock (V-28-no year, collector unknown), 13 ; Jack Run, Alleghany County (VI-14-1908,

collector unknown), 13 , 19 ; Ohiopyle, Fayette County

(VII-22-1905, H. Kahl), 19 , (VII-28-1905, H. Kahl),

1 3 , (VIH-2-1905, H. Kahl), 13 , 4 9 9 , (VIII-7-1905,

H. Kahl), 2 3 3 , (VIII-11-1905, H. Kahl), 19 , (VHI-no

date-1907, H. Kahl), 8 3 3,399 ; Point Pleasant (V-30 -

1914, collector unknown), 19 ; Swarthmore (V-28-1907,

collector unknown), 19 , (VH-no date-1908, collector unknown), 13 , 19 , (VI-13-1909, E. Cresson, Jr.), 13 ,

(VIII-21-1910, E. Cresson, Jr.), 19 ; 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), 13 -

Quebec: Laniel (VII-3-1944, A. Brooks), 19 ; Old

Chelsea (VIII-31-1958, J. Vockeroth), 1 9 ; Rigaud (VIII3-1919, J. Ouellet), 13 -

South Carolina: Cross Anchor (VII-3-1953, M. Wheeler), 633 , 699 .

Tennessee: Greenbrier Cove, Great Smoky Mountains National Park, 2,000 feet (V-18-1957, J. Vockeroth), 2 9 9 ;

Knoxville (VIII-17-1939, R. Beaman), 13 ; Shelby Forest

State Park (IX-3-1952, M. Wheeler), 13 .

Texas: Huntsville (IV-11-1952, M. Wheeler), 13 -

Virginia: Fairfax (VII-19-1954, M. Wheeler), 13 ;

Falls Church (V-26-1951, W. Wirth), 4 3 3, 2 9 9 , (VII15-1954, W. Wirth), 13 , 19 ; Maywood, Alexandria (V -

21-1922, W. McAtee), 13 , 2 9 9 , (VII-9-1922, W. McAtee), 19 ; Reddish Knob, Augusta County (VIII-29-

1953, W. Wirth), 2 3 3, 5 9 9 .

West Virginia: Cranberry Glades (VII-16-1955, C. Sabrosky),2 3 3 .

IMMATUR E SPECIMENS EXAMINED.—36 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, Clearwater 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 between the male terminalia, this species must be
74 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

placed in the H. griseola species group. The immature 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 considerably 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 development 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 polyphagous. 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 Protozoa, 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 repeatedly touched her abdominal venter to the microcrustacean. 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 specimens 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 attempting to mount. In one observation, copulation lasted

for 10 minutes. The female walked intermittently

during copulation. Modes of species and sex recognition 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 possibly 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 Grigarick'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 temperature 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 attempted 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 damaged 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 temperature 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 pupariation 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 distribution 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 infested Z. aquatica, and a third-instar larva pupariated 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 infested rice growing in quiet water supported this

assumption.

I measured the percent of infestation of Z. aquatica 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 thirdinstar 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 anteroventrally 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 apparently 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; antenna velvety dark brown; apicodorsal antennal

prominent; 2 basal coxals; costal section II: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; antenna 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 mesanepimeron 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:

II:I 1.4-2.2; III:IV 2.5-2.9; V:IV 3.0-3.5; Mi*s

index 1.1-1.5.

ABDOMEN.—Terga semiglossy dark brown medially, light gray laterally and ventrally. Male terminalia: 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 covered by fused surstyli). Anterior margin of fused

surstyli deeply cleft medially; B-index about 0.8.

THIRD-INSTAR LARVA.—Body of larva not examined. 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 (Figure 110). Prothoracic end slightly convex in ventral

view; head-lobe scar obovoid or subcircular; maximum 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 Biological Station, Lake Itasca, Itasca State Park, Clearwater 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 Biological 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 aquatica 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: Braconidae) 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 anterior 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 moderate yellow; 8-10 aristal rays; prementum light

gray; apicodorsal antennal prominent; antenna dark

brown; fronto-orbital area and frontal vitta moderate 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 mesokatepisternum; legs light-gray pruinose except darkbrown 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; costalsection 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 protuberant; 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 parafrontale 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; abdominal segment 8 without ventral, transverse row of

setulae, but with 3 annuli of spinules. Body translucent, 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; maximum 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 maximum 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 County, Michigan (VIII-14-1941, C. O. Berg).

ADULT SPECIMENS EXAMINED.—131 (39o"cf,

94$ ? ) from 22 localities:

Iowa: South Pond, Ledges Park, Boone County (VII10-1960, 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), 13 ,

399 .

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 , 19 , (VIII-7-1947, C. Berg), 13 , 19 ; Douglas

Lake, Cheboygan County (VII-3-1941, C. Berg), 19 ;

Nigger Creek, Cheboygan County (VIII-21-1941, C. Berg),

1$.

Minnesota: Bohall Lake, Itasca State Park (VIII-17-

1963, D. L. Deonier), 13 , 19 ; Chamber's Creek, Itasca

State Park (VII-18-1963, D. L. Deonier), 53 3 , 219 9 ;

Douglas Lodge Bay, Lake Itasca, Itasca State Park (VII20-1963, D. L. Deonier), 5$ 3, 89 9 ; Eagles Nest (VI25-1959, W Balduf), 19 ; Headwaters of LaSalle Creek,

Itasca State Park (VIII-6-1963, D. L. Deonier), 19 ;

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), 13,399 .

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 populationdensity of this species increased by perhaps 50 percent 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 populationdensity, 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 columbiana (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; mesofacial 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 moderate 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 localities:

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), (VI13-1955, 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 interfractural 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 mesanepisternum; 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 prominent anteromedial lobe and anterolateral lobe and

irregularly setose except for seriated setae on

posteromedial margin. Postgonite bent slightly anteriad; 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*, 46 ? $ ) from 33 localities:

Arizona: Oak Creek Canyon, 6,000 feet (VHI-no dates,

F. Snow), 1$ (USNM); Rustler Park, Chiricahua Mountains (VIII-27-1933, collector unknown), 19, (X-18-

1958, M. Adachi), 13 , 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); Herkey Camp (V-14-1950, A. Melander), 13,3 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); Sardine Creek, Mono County, 8,500 feet (VI1-3-1951, collector unknown), 19 (USNM), (VII-6-1951, collector unknown), 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 unknown), 19 (USNM); Ohio Pass Road, Gunnison County

(VIII-26-1961, D. L. Deonier), 19 (ISC).

Mexico: Mexico City (V-no date-1888, collector unknown), 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 (XI5-1938, 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 between 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., Equisetum 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; frontoorbital 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 expanded (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: II: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 tapering 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 examined. 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. Prothoracic 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 subterminal. Empty puparium translucent, light yellowish brown.

TYPE.—Holotype male, USNM 43453.

TYPE-LOCALITY.—White Mountains of New Hampshire (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 Waterville, 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),

233 , 1$.

Michigan: Dickinson County (VII-1-1955, R. Dreisbach), 19; Midland County (VI-10-14-1951, R. Dreisbach),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. Aldrich), 13, 69 9; Stinson Lake, White Mountains (VII23-1961, 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 (VII17-1957, 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, (VIII3-1957, C.Hines), 13-

Washington: 15 miles west of Kettle Falls (VII-14-1960,

W. Rapp), 13.

IMMATURE SPECIMENS EXAMINED.—7 from 4 localities:

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 emergence 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 terminalia 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 lightgray 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: II:I 2.2-2.6; III: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 irregularly 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\ 59 ? $ ) from

24 localities:

Illinois: Chicago (no dates—1899, collector unknown),

1 3 ; McHenry (VIII-21-no year, A. Melander), 13 .

Indiana: Chesterton (VI-2-1916 , collector unknown),

IS.

Iowa: Siewer's Springs State Park, near Decorah (IX -

15-1962, D. L. Deonier), 13,299 -

Minnesota: Biological Station, Itasca State Park (VI26-1963, 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), 11 4 3 , 19 9 9 ;

Crookston (IX-5-1936, D. Denning), 19 ; Douglas Lodge

Bay, Lake Itasca, Itasca State Park (VI-20-1963, D. L.

Deonier), 5 3 3 , 4 9 9 ; Iron Corner Lake (VI-18-1963 ,

D. L. Deonier), 13 , 1 9 ; Itasca State Park (VII-14-1938,

P. Schroeder), 19 ; Kabekona Creek, Hubbard County

(VIII-8-1963, D. L. Deonier), 2 9 9 ; Mississippi River at

Sucker Creek, Clearwater County (VII-9-1963 , D. L. Deonier), 19 ; Mississippi River near north entrance Itasca

State Park (VII-9-1963, D. L. Deonier), 13 , 19 ; Prof

Green Trail, Itasca State Park (VII-25-1963 , D. L. Deonier), 13 ; Squaw Lake, Itasca State Park (VIII-14-1963 ,

D. L. Deonier), 2 3 3 ; Two Island Lake, Itasca State

Park (VI-28-1963, D. L. Deonier), 13 ; west side across

from Biological Station, Lake Itasca, Itasca State Park

(VII-28-1963, D. L. Deonier), 13 , (VIII-14-1963, D. L.

Deonier), 8 3 3,20 9 9 .

New Jersey: Trenton (VIII-5-1910, collector unknown)

1 9 .

New York: Dryden Lake, Tompkins County (VIII-8 -

1961, D. L. Deonier), 2 9 9 ; Ithaca (VI-8-1937, collector unknown), 1 9-

Pennsylvania: 2 miles north of Narberth, Montgomery

County (IX-9-1915, collector unknown), 13 .

Quebec: lie Jesus, 4 miles north of Montreal (VIII-19 -

1956, J. Laffoon), 13 .

Virginia: Alexandria (VII-6-1952 , W. Wirth), 19 .

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 following 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 minutes 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 copulating 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 emergence 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; thoracic pleuron nearly all light gray. Male length

1.33-1.67 mm; female 1.70-2.00 mm. Male terminalia 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 lightbrown 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 convex; anterolateral margin of sternum 5 obtusangular

(about 100°); copulobus conical, directed posterolaterad, and irregularly setose, with 4-5 medially

inclined macrochaetae on posteromedial margin.

Postgonite short, bent anteriad; postgonite uncus

fusiform and straight; distiphallus short, almost uniform 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, collector 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. Melander), 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 III: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 lightgray 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: II:I 1.8-

2.2; III: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; anterolateral 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 examined. 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; clypealarch index 1.6-1.7. Clypeal arch sloping smoothly

along about 20° angle relative to lower frontoclypeal

margin. Mouth-hook beak longer than base; maximum 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; maximum breadth: minimum breadth 5.0-6.0; anal-plate

index 1.8-2.4. Prothoracic end semicircular in ventral view; head-lobe scar subcircular; maximum

puparial breadth 7.5-8.0 times maximum breadth

of head-lobe scar and 2.5 times maximum prothoracic breadth; prothorax with many ventral

rugulae; mesothorax and metathorax prominently

rounded laterally; lateral spinules inconspicuous in

midlength of puparium; abdominal segment 8 without ventral, transverse row of setulae and with only

2-3 distinct annuli of spinules (immediately perispiracular); anal plate ovoid, with anterior margin

slightly or distinctly convex. Empty puparium translucent, 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. Mason), 1 $.

California: Big Bear Valley (VIII-9-1933, collector unknown), A$S, 65 9 ; Buena Park (V-19-1914, A. Melander), 2SS, 1$ ; Carpenteria (VIII-11-1950, A. Melander), 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. Grigarick), 7$S, 29 9, (IX-21-1957, A. Grigarick), IS,

(IX-22-1955, A. Grigarick), 19, (IX-27-1955, A. Grigarick), 2$S, 49 9 ; Green Valley (VII-26-1944, A.

Melander), 1 $ ; Jenks Lake, San Bernardino (VIII-3-

1942, A. Melander), 29 9, (VII-22-1957, J. Hall), 19 ;

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), 19 ; Springbrook State

Park, Guthrie County (VI-23-1961, D. L. Deonier), 13 .
NUMBER 6 8 85

Massachusetts: Nantucket (VII—20—year unknown, collector 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), 13 .

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, collector unknown), 13,2 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), 19 , (VII-29-1949, E.

LeRoux), 13 , 19, (VII-30-1949, E. LeRoux), 13 , 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 Yellowstone 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, during 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 terminalia 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 moderate yellow; frontal vitta and fronto-orbital area

yellowish gray or strong yellowish brown; 14-17

postoculars.

THORAX.—Ppn and mesonotum strong yellowishbrown; 3^4 adc and 2 pdc; pleuron light gray or

strong yellowish brown; 1-2 basal coxals (1 macrochaetous); legs light-gray pruinose except darkbrown 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: II:I 2.5-2.9; III: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. Postgonite 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), 15 .

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; epistomal 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 olivebrown; 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) ; costalsection ratios: 11:1 1.8-2.2; III:IV 3.5-4.0; V:IV

3.0-3.6; Mi . 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 pregonite 2.0 times that of postgonite, pregonite extending 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 localities:

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); Victorville, 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 postoculars.
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 darkyellow 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: II:I 2.2-2.5; III:IV 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), 19 .

Idaho: Lake Coeur d'Alene (VII-15-1926, E. Nast),

1 9 ; Potlatch (VI-17-1911, collector unknown), 1 $ ; Soldier 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), 19 ;

Beaver Creek, Newport (no dates, J. Aldrich), 39 9 ; Breitenbush Springs, 2,222 feet (VII-6-1934, H. Scullen), 19 ;

Gold Beach (VII-27-1951, collector unknown), 29 9 ;

Hood River (no dates, Childs), 13 , 599 ; Independence

(VII-25-1934, N. Larson), 19 ; Marshfield (VI-27-no

year, J. Aldrich), 2$ 3 , 19 ; Newport (VI-9-1925, E. Van

Dyke), 13 ; Sear Lake, Fort Lewis, Pierce County (IX-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 date1917, A. Melander), 1$, 19, (VIII-27-1917, A. Melander), IS, (VI-28-1925, A. Melander), 2tt; Lake

Stevens, Everett (VIII-5-1917, A. Melander), 19 ; Seaview (VI-26-1925, A. Spuler), 1 $ ; Tacoma (VIII-27-

1911, collector unknown), 13 ; 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 intergradational 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 parafrontale velvety black; apicodorsal antennal prominent; 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: II:I 2.2-3.5; III: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°); copulobus 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, 528$$ )

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),

1 5, (VII-3-1947, L. Beamer), 29 9 ; Coconino County

(VIII-1&-1927, R. Beamer), 13 ; Hassayampa River,

Wickenburg (VI-29-1953, W. Wirth), 19 ; 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), 13 ; Sunnyside Canyon, Huachuca Mountains (VI1-9-1940, L. Lipovsky), 19.

British Columbia: Atlin, 2,200 feet (VI-13-1955, H.

Huckel), 19 , (VIII-11-1955, B. Gibbard), 19 ; Okanagan 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. Melander), 13 , (X-2-1946, A. Melander), 13 , 19 ; Auburn

(V-12-1953, J. Hall), 1 <$ , 1 9 ; Barton Flats (VIII-3-1942,

A. Melander), 59 9 ; Barton Store (VII-4-1946, A. Melander), 13 ; Berkeley Hills, Alameda County (IV-11-

1908, collector unknown), 19, (IV-20-1908, collector unknown), 33 <$, 59 9 ; Big Bear Lake (V-24-1935), A.

Melander), 19 ; 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), 13 , 19 , (VII-28-1940, L.

Kuitert), 13 , (VII-28-1940, L. Lipovsky), IS, 19 ;

Bowerman Meadow Trinity County (VI-3-1951, A. McClay), 19 ; Cajon (VIII-9-1944, A. Melander), 19 ;

Campo (VII-18-1940, R. Beamer), 1 9 ; Carson Pass (IX11-1938, M. Cazier), 1 $ ; Cassel, Shasta County (VII-15-

1955, E. Schlinger), 19 ; Cedar Pass, Modoc County (V15-1948, W. Wirth), 19 ; 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), 19 ; Clio, Plumas County (VII-9-1916, H.

Dyar), 1 S ; Coalinga, Fresno County below 500 feet (VI1-3-1907, 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, collector unknown), IS; Dellecker, Plumas County (VII-4-

1951, E. Schlinger), 19 ; Ebbett's Pass, Alpine County

(IX-11-1938, M. Cazier), 23 3 , (VII-22-1957, E. Mezger), 19, (VIII-24-1957, E. Mezger), 19 , (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 (VII23-1955, E. Schlinger), 4 3 $, 59 9 ; Felton, Santa Cruz

Mountains (V-20-25-1907, J. Bradley), IS, 15 ; Garden

Valley, Eldorado County (V-3-1952, E. Schlinger), 1$,

2 9 9 ; Glacial Point Road, Yosemite National Park (VII6-1947, A. Melander), 19 ; Grass Lake, Luther Pass, Eldorado 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 Junction, Lassen County (VII-4-1949, J. MacSwain), 19 ,

(VII-16-1953, E. Schlinger), 1 <J ; Hat Lake, Lassen National Park (VI1-2 3-1950, P. Arnaud), 19 ; Herkey Camp

(V-14-1950, A. Melander), IS, 19 ; near Hobart Mills,

Nevada County (X-10-1952, E. Schlinger), 19 ; Hope

Valley, Alpine County (VII-18-1948, L. Quate), 19 ;

Idylwild (VI-4-1935, P. Oman), 19 ; Jacumba (VII-17-

1940, D. Hardy), 29 9 ; Jenks Lake (VIII-18-1950, A.

Melander), 19 ; Keen Camp (VI-7-1942, A. Melander),

2S S, 19 ; Laguna Mountains (VII-6-1929, L. Anderson),

1 9 , (VII-6-1929, R. Beamer), 19 ; Lake Alpine (IX11-1938, M. Cazier), 3 3 3 , 29 9 ; Lake City, Modoc

County (X-l 1-1952, E. Schlinger and J. Hall), 13 , 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$, 19 ; Lake Tahoe (VIII-11-1940, E.

Kenaga), 19 , (VIII-11-1940, L. Kuitert), 19 , (VIII11-1940, L. Lipovsky), 3$ S, 39 9 ; LaPosata Creek, San

Diego (IV-10-1950, L. Quate), 19 ; Leavitt Meadow,

Mono County (VII-6-1951, A. McClay), 19 , (X-ll1952, J. Hall), 19 ; Likeley, Modoc County (VII-6-1951,

A. McClay), 19, (X-ll-1952, E. Schlinger), 29 9, (X11-1952, 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 Mountain Camp, San Jacinto Mountains (VII-1-1952, A. McClay), 19 ; Mason Creek R. S., Modoc County (X-12-

1952, E. Schlinger), 2 3 3,499 ; Mono Lake, Mono County

(VII-31-1940, E. Kenaga), 1 9 ; Monterey County (VII-5-

1896, collector unknown), 13 , 19, (VII-8-1896, W.

Wheeler), 13 , (VII-9-1896, W. Wheeler), 13,19 , (VII17-1896, G. Hough), 19 ; Morro Bay (VI1-27-1940, A.

Melander), 1 9 ; Ortega Highway, El Cariso Camp (V-26-

1944, A. Melander), 13 ; Pacific Grove (V-7-1906, J.

Aldrich), 39 9, (V-13-1906, J. Aldrich), 19 (Vl-no

dates, W. Mann), 19 : Palm Springs (XII-14-17-1917, J.

Bradley) ,233 , (XI-19-1943, A. Melander), 1 $, 1 9, (XI22-1943, A. Melander), 13 , 29 9 ; Palomar Mountain

(VI-14-1948, A. Melander), 13,299 ; Pinecrest Lookout,

Tuolumne County (VII-7-1948, P. Arnaud), 13 , (VII30-1948, P. Arnaud), 19 ; 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), 13 ,

(X-28-1944, A. Melander), 19 ; Sage Hen, 5 miles northwest 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 Bernardino 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), 19 ,

(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), 13 , (VII-6-1951, A. McClay),

2 9 9, (VII-11-1951, A. McClay), 15 , 2 9 9, (VII-12-

1951, A. McClay), 19, (VII-18-1951, A. McClay), 2 9 9,

(VII-29-1954, J. Downey), 19 ; Sequoia National Park

(VIII-6-1940, L. Kuitert), 19 ; Siberian Outpost, Tulare

County, 9,500-10,500 feet (VII-3-1915, collector unknown), 19 ; Sonora Pass, 9,624 feet (VII-12-1951, A.

McClay), 19 , (VII-17-1951, A. McClay), 69 9 ; South

Fork, Santa Ana River (VII-29-1942, A. Melander), 13 ,

1 9 , (VIII-4-1945, A. Melander), 19 ; South Fork Camp,

Barton Flats (VIII-31-1944, A. Melander), 13 , (IX-3 -

1944, A. Melander), 19 ; South Fork Camp, Santa Ana

River, San Bernardino County (VI-28-1952, R. Beamer

et al.), 19 ; Stanford University, Santa Clara County

(XII-29-1897, collector unknown), 19 ; Strawberry,

Tuolumne County (VI-15-1951, A. McClay), 13 , 19 ,

(VI-19-1951, A. McClay), 13 , 19 , (VI-22-1951, A.

McClay), 29 9 ; Sugarloaf, Barton Flats (IX-17-1945, A.

Melander), 13 , 59 9 ; Sugarloaf Mountain (V-l 1-1947,

J. Sperry), 19 , (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), 19 ; Thousand Springs, San Bernardino Mountains (X-24-1946, A. Melander), 59 $ ; Tioga Pass (VII31-1940, 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), 19 ; Truckee (VII5-1936, A. Pritchard), 1,5, (VII-6-1927, E. Van Duzee),

3$ $, 39 9 ; Tuolumne Meadows (VIII-1-1940, R.

Beamer), 13 , 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), 19 , (IX-4-1950, A.

Melander), 2$ $, 19 ; 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), 13 , 19 ; (VIII-1-1940, R. Beamer), 13 ,

(VIII-1-1940, D. Hardy), 13 , 29 9, (VIII-1-1940, E.

Kenaga), 19 , (VIII-1-1940, L. Kuitert), 19, (VIII1-1940, L. Lipovsky), 2 3 3 , 29 9 ; Yuba Pass, Sierra

County (VIII-20-1953, E. Schlinger), 19 ; 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), 19 ; 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 (VIII13-1934, C. Sabrosky), 19 ; 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), 19 ; Juliaetta (no dates, J. Aldrich), 19 ;

Moscow Mountain (VIII-23-1908, collector unknown),

\$, 39 9 ; Potlatch (IX-21-1918, A. Melander), 13 ,

29 9 ; Priest Lake (IX-3-1919, A. Melander), 19 ; Swan

Lake, Bancock County (VI-19-1952, E. Schlinger), 19 ;

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), 19 ; 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), 19 ; Tajique (VI-28-1947, R. Beamer), 19.

Oregon: Crater Lake (IX-16-1934, A. Melander), 19 ;

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 (VIII15-1930, F. Wynd), 19 ; Crater Lake National Park, Sun

Creek Meadow, 6,500-7,000 feet (VIII-26-1930, H.

Scullen), 19 ; 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),

1 9.

Texas: Henkes Pond, Kerrville (IV-4-1955, W. Wirth),

13,19 .

Utah: Cedar City (VIII-13-1939, P. Oman), 13,799 ;

Echo (X-16-1943, G. Knowlton), 2 $ 6 ; Eden (VI-21-

1937, D. Hardy), 19 ; 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. Knowlton), 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), 19 ;

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), 13 ;

Klickatack River, Glenwood (VI-27-1917, A. Melander),

1 & ; Mount Rainier National Park, Berkeley Park (VIII23-1934, A. Melander), 1 9 ; Mount Rainier National Park,

Paradise Park (VIH-no date-1917, A. Melander), 19 ;

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&&, 19 ;

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), 19 ; Uniontown (VI-26-1932, J. Aldrich),

1 9 .

Wyoming: Kemmerer (VIII-14-1950, M. Wheeler),

1 $ ; Yellowstone National Park, Geyser Basin (VIII-7 -

1919, A. Melander), 1$, 2 9 9 ; Yellowstone National

Park, Norris Basin (VII-26-1923, A. Melander), 19 .

REMARKS.—Except for H. decens, the male of

which is unknown, this species probably could be

confused only with H. melanderi. Hydrellia melanderi 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 moderateyellow 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 anterior interfractural costals; costal-section ratios:

II:I 2.0-2.3; III: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 obtusangular (about 100°) ; copulobus lanceolate; posterior 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 \ 37$? )

from 11 localities:

Connecticut: Southwest corner of Sleeping Giant State

Park, near Hamden (VI1-30-1961, J. Laffoon), 19 .

Illinois: Chicago (VIII-29-1896, W. Wheeler), 19 -

Iowa: Goose Lake, Hamilton County (VII-14-1960, D.

L. Deonier), 1 £ ; Ledges State Park, Boone County (VIII16-1960, 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 , (IX-22-1962 , D.

L. Deonier), 1$, 19 ; Spring Lake, Greene County

(IX-19-1961, D. L. Deonicr), 4 3 $, 12 9 9 ; Springbrook

State Park, west «/2 sec. 33, T.81 north, R.31 west, Guthrie

County (VII-19-1960, D. L. Deonier), 19 .

Kansas: Marais des Cygnes Wildlife Refuge, Linn County

(IX-5-1961 , D. L. Deonier), 5$ $, 8 9 9 .

Maine: Bar Harbor, Mount Desert Island (VII-15-1933 ,

W. Procter), 1$ ; Mount Desert Island (VII-25-1935 , W.

Procter), 2 9 9 .

Michigan: Augusta (X-4-1953 , collector unknown), 19 .

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 distiphallus 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 parafrontale not velvety; thoracic pleufon bronzed moderate 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 parafrontale very dark brown; fronto-orbital area and

frontal vitta moderate olive-brown; 12-16 postoculars.

THORAX.—Most of thorax and legs bronzed moderate 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. Postgonite 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 (VIH-no 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 (VI29-1928, C. Curran), 1$, 35$ .

Pennsylvania: Ohiopyle, Fayette County (VIH-no date1907, 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 coxals. 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 moderate 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; clypealarch index 1.8-2.0. Clypeal arch angled 20°-25°

in relation to lower frontoclypeal margin. Mouthhook beak distinctly longer than base; maximum

mouth-hook base thickness about 2.0 times that of

beak; mouth-hook light spot small, elliptical. Prothorax and mesothorax sparsely spinulose; abdominal segment 8 without ventral, transverse row of

setulae, but with many irregular dark spinules ventrally. 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; maximum 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 translucent, light brown.

TYPE.—Holotype female, ANSP (not numbered).

TYPE-LOCALITY.—Spencer Lake, New York (VI30-1907, 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), 19 , (V-20-

1940, C. Berg), 19, (VII-17-1941, C. Berg), 19 , (II2-1942, 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), 19 ; Whitmore

Lake, Washtenaw County, (III—5—1941, reared by C. Berg),

2$ $, (111-11-1941, reared by C. Berg), 19 .

Minnesota: Squaw Lake, Itasca State Park (VIII-14-

1963, D. L. Deonier), 19 ; Two Island, Itasca State Park

(VI-28-1963, D. L. Deonier), 13 , (VIII-8-1963, D. L.

Deonier), 7 9 9, (VIII-18-1963, D. L. Deonier), 19 ;

1.3 miles south of main entrance, Itasca State Park (VI26-1963, D. L. Deonier), 79 9.

New York: Ithaca (no dates or collector), \$.

Ontario: London (V-20-1958, W. Judd), 19 , (V-21-

1958, W. Judd), 19 , (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), IS ,

399 .

Washington: Orcas Island above Mountain Lake (VIII18-1925, A. Melander), 19 .

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 (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 prominent face bordered by the

nonseriated primary facials together with its relatively 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 instance, 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 collected several adults by the lighted-receptacle method.

Berg found eggs in masses one layer deep concealed 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 stadium 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 "Apparently 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 hymenopterans 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. columbiana 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 lightgray 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 anterior margin of sternum 5 broadly notched; anterolateral 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, Tennessee, 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 physiognomically 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 medially, light gray laterally and ventrally. Male terminalia: 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, truncate, long, and directed laterad; distiphallus tapering to mucronate apex (except small preapical

constriction) ; distiphallus carinate and finely lamellate 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 mesanepisternum 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: II:I 2.0-2.5; III:IV 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 (45°-50°);

anterior and lateral margins of sternum 5 distinctly

more heavily sclerotized; copulobus truncate posteriorly and irregularly setose. Postgonite bent anterolaterad and postgonite uncus slightly laterad; distiphallus 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 (VH-no

date-1917, A. L. Melander).

SPECIMENS EXAMINED.—181 (70o*d\ 111?? )

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, 19 ; 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), 19 ; 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), 13 , 19 , (VI-18-

1953, E. Mason), 13 , (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), 13 ; Moresby

Camp, Queen Charlotte Islands (V-31-1957, E. MacDougall), 1 3 ; peat bog at Pitt Meadows (VII-9-1953,

W. Mason), 1,5, 29 9 ; Vancouver (VI-20-1958, H. and

A. Howden), 19 ; Victoria (VII-10-1924, A. Melander),

1 3 , 299 .

Colorado: Bear Lake, Estes Park, 1,000 feet (VII-11-

1934, A. Melander), 1 9 ; Cameron Pass (VIII-19-22-1940,

C. Sabrosky), 13 ; Marshall Pass, 10,856 feet (VII-28-

1906, collector unknown), 13, 19 ; Rocky Mountain

Biological Laboratory, Gunnison County, 9,500 feet (VIII26-1961, D. L. Deonier), 13.

Idaho: Pottsville (VIII-23-1916, A. Melander), 13-

Manitoba: Churchill (VII-8-1952, J. Chillcott), 19 ,

(VII-30-1952, J. Chillcott), 13 ; 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. Chillcott), IS, (VII-10-1952, J. Chillcott), 19 ; Mile 505 of

Hudson Bay Railway (VI-29-1952, J. Chillcott), 19 ,

(VII-16-1952, J. Chillcott), 19, (VII-26-1952, J. Chillcott), 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, 19 ; Ilwaco (VII-no date-1917,

A. Melander), 2 3 3,299 , (VIII-27-1917, A. Melander),

1 $ , (Vl-10-1918, A. Spuler), 2 3 3,299 , (VI-18-1918,

A. Spuler), 63 3 , 79 9, (VII-no date-1918, O. Miner),

2 9 9, (VI-28-1925, A. Melander), 13 ; Longmire (VI27-1935, A. Melander), 13 ; Mount Constitution (VII31-1909, collector unknown), 13 , 19 ; Olga (V-17-1910,

collector unknown), 19 ; Olympia (VII-17-1922, A.

Melander), 13 ; Pluvius (VII-16-1922, A. Melander),

13,19 ; Point Gamble (VIII-16-1910, collector unknown),

1 3 ; Poulaho (VIII-17-1910, collector unknown), 13 ;

Roche Harbor (VII-3-1909, A. Melander), 13 ; Seaview

(VI-28-1925, A. Spuler), 33 3 , 3 9 9, (VI-30-1925, A.

Spuler), 13 ; 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 II: 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 yellowish brown; 3-4 adc and 2 pdc; pleuron light

gray; legs light-gray pruinose except moderateyellow 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: II:I 3.0-3.7; III: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 convex, 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 occasionally anastomosing; spaces between ridges regularly micropunctate. Micropylar protuberance infundibulate, exposed in dorsal view.

FIRST-INSTAR LARVA.—Length 0.38-1.20 mm;

maximum breadth 0.06-0.20 mm. Clypeal arch distinctly 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 Figure 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 frontoclypeal 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, triangular, 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; maximum 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 translucent, 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), 13 .

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 , 4799 ;

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 Hydrellia spinicornis without access to the holotype; however, paratype comparison was made, and it seems unlikely 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 exceedingly 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 contained late pupae. Mines in stems and stolons containing puparia had an escape slit either lateral to

or slightly anterior to the operculum of the puparium, 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 localities 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 II: 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 parafrontale 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:

II:I 2.4-3.2; III: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 anterolaterad; postgonite uncus straight (directed anterolaterad) ; 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 (VIII27-1911, A. L. Melander).
98 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

SPECIMENS 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, 19 .

Wyoming: 12 miles northwest of Lusk (VH-no date1895, 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 index 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 olivebrown; 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: II:I 1.7-

2.3; III: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; copulobus 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; distiphallus 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, USNM 70542.

TYPE-LOCALITY.—Two Island Lake, Itasca State

Park, Becker County, Minnesota (VI-28-1963,

D. L. Deonier).

PARATYPES.—I l l (24o* d \ 87 ? 9 ) from 19 localities:

Connecticut: Avon Old Farms, Avon (VI-27-1929, C.

Curran), 29 9 (AMNH).

Florida: DeFuniak Springs, Walton County (VIII-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 Whitewater (VI-22-1958, C. Miller), 19 (ISC).

Massachusetts: Mashbee (VII-28-1950, collector unknown), 29 9 (USNM and ISC); Woods Hole (VIIIno 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), 15,2 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$, 129 9, (IX8-1952, E. Smith), 19, (IX-9-1952, J. McAlpine), 5<$ $,

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 date1907, H. Kahl), 19 (ISC).

Yukon: Reflection Lake, mile post 1160 of Alaska Highway (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.rocotyle 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 ventrally. 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 (VII19-1960, D. L. Deonier), 2 9 9.

Maine: Mount Desert Island (VII-12-1935, W. Procter),

19.

REMARKS.—I have studied the holotype, but careful 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 darkbrown 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; costalsection ratios: II: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; postgonite 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 entirely black like the mouth-hook (Figure 87). Mouthhook 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; analplate index 2.0. Prothorax and mesothorax damaged, 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 terminal. 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), 13 , 12 ; 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), 13 , 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), 12 ; 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), 13 , 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), 13 ; San Diego (XI-18-1916, H. Dyar), 12.

Colorado: Alamosa (VIII-1-no year, M. T. and G. M.

James), 12 ; Creede, 8,844 feet (VHI-no date-1914, S.

Hunter), 12 ; Creede (VII-1-1936, L. Tuthill), 12,

(VII-10-1939, L. Tuthill), 12 ; Deckers (VI11-25-1950,

collector unknown), 12 ; Electra Lake, about 37°33' N,

107 °48' W, 8,400 feet (VI-28-VII-1-1919, collector unknown), 23 3 ; Estes Park (VIII-11-1952, R. Dreisbach),

1 2 ; Grand Junction (IX-4-1938, D. Hardy), 13 , 32 2 ;

Tennessee Pass, 10,240 feet (VII-8-no year, J. Aldrich),

12, (VII-24-1917, collector unknown), 12 ; Wray, about

40° N, 102°10' W (VIII-17-19-1919, collector unknown),

12.

Delaware: Rehoboth (VI-25-1939, A. Melander), 12 ;

(VIII-4-1941, A. Melander), 13-

Florida: Baxter (IX-no date-1954, M. Wheeler), 12 ;

Fruitville (VIII-11-1930, R. Beamer), 12.

Idaho: Moscow (X-9-1907, collector unknown), 12 ;

Soldier Creek, Priest Lake (VIII-22-1919, A. Melander),

1 3 , (VIII-22-1920, A. Melander), 12 ; 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. Shackleford), 2 2 2, (VII-10-1926, V. Smith), 12 ; Meredosia

(V-29-1917, collector unknown), 12 ; Peoria VIII-26-

1917, J. Aldrich), 12 ; Springfield (IX-7-1952, M.

Wheeler), 12.

Indiana: Lafayette (X-22-1915, collector unknown), 13-

Iowa: Ames (IV-16-1927, collector unknown), 13 , 19 ,

(IV-24-1927, collector unknown), 12, (V-12-1927, collector unknown), 12 ; Ames, Skunk River (VII-17-1960,

D. L. Deonier), 1 2 ; Ames, Izaak Walton League Reserve,

Story County (IV-28-1960, D. L. Deonier), 12 , (VI17-1960, D. L. Deonier), 12 , (IV-23-1962, D. L.

Deonier), 13 , 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), 13 , 12 ; Fraser Dam, Boone County

(IX-4-1960, D. L. Deonier), 12 , (VII-6-1961, D. L.

Deonier), 13 , 12 ; 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), 13 , (VI-14-1960,

J. Laffoon), 12, (VIII-16-1960, D. L. Deonier), 12 ,

(VI11-18-1960, D. L. Deonier), 13 , 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), 12 ; Little Wall Lake, Hamilton County (VII13-1960, D. L. Deonier), 13,322 , (VIII-27-1960, D. L.

Deonier), 33 3 , 32 2, (VI-18-1961, D. L. Deonier),

4 3 3 , 222 ; 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), 12 ; near Missouri River, 7 miles

southwest of Whiting, Monona County (VI-6-1960, D. L.

Deonier), 13 ; 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 , 1499 , (IX-19-1962,

D. L. Deonier), 2 3 3 , 19 ; Springbrook State Park,

Guthrie County (VII-19-1960, D. L. Deonier), 13 , (VI23-1961, D. L. Deonier), 1 3 ; West Okoboji Lake, Dickinson County (IX-3-1949, J. Laffoon), 19 ; 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), 19 ; 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), 13 ,

(VIII-2-9-1937, D. Denning), 19 ; Herchmer (VIII-1 -

1937, D. Denning), 13 , 29 9 ; Whitewater Lake, 4 miles

north of Whitewater (VII-17-1958, J. Chillcott), 19 .

Maryland: Plummer's Island, Potomac River (IV-28-

1914, R. Shannon), 19 ; unspecified locality (no dates, H.

Loew), 13 .

Michigan: Baraga County (VIII-27-1952, R. Dreisbach), 19 ; Bay County (VIII-11-1951, R. Dreisbach),

1 9 ; Cass County (VIII-2-1953, R. Dreisbach), 13 , 19 ;

Cheboygan County (VII-26-1935, L. Penner), 19 ; Vineyard 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), 19 ;

Chisago County (no dates, O. Oestlund), 29 9 ; Cook

County (VIII-21-1938, H. Milliron), 39 9, (VIII-21-

1938, H. Peters), 19 ; Eaglenest (VIII-26-1959, W.

Balduf), 13 ; Freeborn County (IX-3-1937, R. Daggy),

1 3 , 19 ; Haydenville (IX-11-1935, A. Pritchard), 13 ,

2 9 9 ; Headwaters of LaSalle Creek, Itasca State Park

(VIII-6-1963, D. L. Deonier), 13 , 29 9 ; Itasca State

Park (VI-20-1938, H. Milliron), 19 ; Mille Lacs (VI-2-

1937, D. Denning), 19 ; St. Paul (V-25-1926, S.

Kepperley), 19 ; Washington County (V-7-1938, H. Milliron), 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 (VII10-1962, 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), 13 , 19 ; Dickinson's

Pond, Lamar County, sec. 3, T.4 north, R.I4 west (VII11-1962, D. L. Deonier), 5 3 3,699 ; near U.S. Highway

90, sec. 20, T.7 south, R.5 west, gravel pit (VI-9-1962,

D. L. Deonier), 13 , 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), 13 ; 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), 13 ; 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. Vockeroth), 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 Mountains 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), 19 .

Pennsylvania: Ohiopyle, Fayette County (VIII-7-1905,

H. Kahl), 13 , 29 9, (VIII-11-1905, H. Kahl), 33 3 ,

19 ; Swarthmore (VI11-22-1909, E. Cresson), 19 .

Quebec: Gaspe (VIII-9-1937, C. Alexander), 19 ;

Gaspe Bay (VII-18-1931, J. Aldrich), 19 ; Great Valley,

Gaspe (VII-20-1931, J. Aldrich), 3 9 9 ; La Ferme (VIIno date-1943, A. Roberts), 19 ; Notre Dame du Portage

(VIII-17-1957, W. Mason), 19 ; Rupert House (VII-25-

1949, D. Gray), 19 .

Tennessee: Dale Hollow Reservoir, 6 miles south of Byrdstown, 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), 19 ; Reelfoot State Park, Reelfoot Lake (VIII-16-1962, D. L.

Deonier), 333 3 , 389 9 ; Samburg, Reelfoot Lake (VIII11-1962, 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 , (X19-1950, M. Wheeler), 13 , (XI-19-1950, collector unknown), 19 , (XII-1-1951, M. Wheeler), 19 ; Double

Lake (IV-11-1952, collector unknown), 1 $ ; Galveston

(IX-13-1953, M. Wheeler), 19 ; Goliad (IX-5-no year,

Townsend), 19 ; Guadalupe River, Gonzales (IV-22-1956,

W. Wirth), 2$ 2; Rio Hondo, Cameron County (VIII2-1950, M. Wheeler), 2$ $, 3 9 9 ; San Antonio (111-28-

1942, A. Melander), 29 9 ; Sinton (XII-25-1945, R. Beamer), 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), 19 .

Washington: Union Flat (IX-8-1918, collector unknown), 1 S •

Wyoming: Lander (VIII-16-1950, M. Wheeler), 19 ;

Yellowstone National Park, Biscuit Basin (VIII-2-1934, A.

Melander), 49 9 ; Yellowstone National Park, northwest

entrance (VII-27-1923, A. Melander), 19 ; Yellowstone

National Park, Yellowstone Lake (VII-18-1923, A. Melander), 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 recognized 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 latifolia, 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 hypnoides; 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 indicated 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 greenishyellow, 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 overwintered in the plants or had recently hatched, for

the adults did not emerge until 28 days after collection 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; apicodorsal 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 posterior half of each segment). Male terminalia:

median third of posterior margin of sternum 5 deeply,

rectangularly recessed and congruent with distiphallus; anterolateral margin of sternum 5 with

inconspicuous dorsal process projecting laterad;

copulobus slightly convex posteriorly, without macrochaetae, 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 moderately prominent lateral convexities; B-index about

2.0; C-index 0.6-0.8. Syntergum 9+10 rounded posteriorly. 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 mouthhook base thickness about 2.2 times that of beak;

mouth-hook light spot touching margin of mouthhook 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 spinulose 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 (VIII3-1931, 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 3,799 ,

(IX-19-1961, D. L. Deonier), 63 4 , 39 9 , (V-4-1962,

D. L. Deonier) 19 ; Springbrook State Park, Guthrie

County (VI-23-1961, D. L. Deonier), 14 , 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 5,19 .

New York: Dryden Lake, Tompkins County (VIII-8-

1961, D.L. Deonier), 19.

North Carolina: Highlands Biological Station, Highlands, Macon County (VI-15-26-1968, D. L. Deonier),

83 3,20 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 deposited 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 specimens. 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 February and crawled around the bottom of the dish

until I placed it on the fresh leaf. It started excavating 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 interfractural 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 terminalia 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 infrequently 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: II:I 2.2-

2.8; III: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 irregularly 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, 64$? )

from 24 localities:

Connecticut: Goose Island (VII-21-1913, collector unknown), 1 $ ; Greenwich (VI-21-1916, E. Kalmbach), 1 $ ;

New Haven, west shore of harbor (VII-29-1961, D. L.

Deonier), 3$$, 18$$ , (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 $ $ ; Horseneck 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$, (VII21-1954, M. Wheeler), 5$ $, 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 (VU-n o

other data), 4SS, 2 9 9 , (VII-6-1927, A. Melander),

1$, 19 (VII-26-1932, C. Curran), 19 .

Nova Scotia: Lockeport (VII-18-1958, J. Vockeroth),

1 9 ; Smith's Cove (VI-24-1955, G. Steyskal), 19 .

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), 19 .

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 immatures 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; apicodorsal 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: II:I 2.2-2.5; III: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. McDunnough), 1<5, 19 .

California: Glacier Point Road, Yosemite National Park

(VII-6-1947, A. Melander), 19 ; Sequoia National Park

(VIII-6-1940, D. Hardy) ,14,299 .

Colorado: Bear Lake, Routt County, 8,500 feet (VIII19-1935, D. Wilbur), 1 $ ; Lake Brennan, 10,000 feet (VII15-1934, C. Alexander), 31 $ <}, 27 9 9 ; Nederland, Boulder County (VII-10-1937, H. Peters), 19 ; Tennessee Pass

(VII-25-1917, J. Aldrich), 19 .

Wyoming: Kemmerer (VIII-14-1950, M. Wheeler), 1$,

399 .

REMARKS.—The male terminalia of H. wilburi

and H. platygastra appear to be identical. The two

species may be conspecific, but since they are sympatric 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

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Allen, Pamela

1957a. The Larval Morphology of Agromyzidae (Diptera). Proceedings of the Royal Entomological

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Balachowsky, A., and L. Mesnil

1935. Les Insectes Nuisibles aux Plantes Cultivees. Volume I. Paris: Busson.

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1949. Limnological Relations of Insects to Plants of the

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1954. Les Insectes Aquatiques d'Europe (Genres: Larves,

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1894. I Ditteri del Trentino. Saggio di un Elenco Delle

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1904. Neue Beitrage zur Dipterenfauna der Balkanhalbinsel. Wissenschaftliche Mitteilungen aus Bosnien und der Herzegowina, 9:519—581. (Herausgegeben vom Bosnisch-Herzegowinischen Landesmuseum in Sarajevo).

Sturtevant, A. H.

1925. The Seminal Receptacles and Accessory Glands of

the Diptera, with Special Reference to the Acalypteratae. Journal of the New York Entomological

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1926. The Seminal Receptacles and Accessory Glands of

the Diptera, with Special Reference to Acalypteratae. 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 Society, 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

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1916. Ueber Wasserhymenopteren. Zeitschrift fiir Wissenschaftliche Insektenbiologie, 12:49-54.

Thompson, W. R.

1943. A Catalogue of Parasites and Predators of Insect

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1961. Contribution a l'Etude de l'Ecologie et du Peuplement des lies de la Region Marseillaise. Colloques

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124 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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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, Clearwater 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, Clearwater 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 radial vein

R4 + 5, combined fourth and fifth radial 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

--HI 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

ORM3+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 (Fallen), male: head, front view ( x 32). 140,

H. griseola (Fallen), male: cibarium of probosis, front view ( X 32). 141, H. griseola (Fallen),

male: cibarium, posterior view ( X 80). 142, H. griseola (Fallen), male: cibarium, right lateral

view ( X 80).

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omit the number. For easy reference, number also the taxa and their corresponding headings

throughout the text; do not incorporate page references in the key.

In synonymy, use the short form (taxon, author, date, page) with a full reference at the end

of the paper under "Literature Cited." Begin each taxon at the left margin with subsequent lines

indented about three spaces. Within a taxon, use a period-dash (.—) to separate each reference.

Enclose with square brackets any annotation in or at the end of the taxon. For references within

the text, use the author-date system: "(Jones, 1910)" or "Jones (1910)." If the reference is

expanded, abbreviate the data: "Jones (1910, p. 122, pi. 20: fig. 1)."

Simple tabulations in the text (e.g., columns of data) may carry headings or not, but they

should not contain rules. Formal tables must be submitted as pages separate from the text, and

each table, no matter how large, should be pasted up as a single sheet of copy.

For measurements and weights, use the metric system instead of (or in addition to) the

English system.

Illustrations (line drawings, maps, photographs, shaded drawings) can be intermixed

throughout the printed text. They will be termed Figures and should be numbered consecutively;

however, if a group of figures is treated as a single figure, the individual components should be

indicated by lowercase italic letters on the illustration, in the legend, and in text references:

"Figure 9b." If illustrations (usually tone photographs) are printed separately from the text as

full pages on a different stock of paper, they will be termed Plates, and individual components

should be lettered (Plate 9b) but may be numbered (Plate 9: figure 2). Never combine the

numbering system of text illustrations with that of plate illustrations. Submit all legends on pages

separate from the text and not attached to the artwork.

In the bibliography (usually called "Literature Cited"), spell out book, journal, and article

titles, using initial caps with all words except minor terms such as "and, of, the." (For capitalization of titles in foreign languages, follow the national practice of each language.) Underscore

(for italics) book and journal titles. Use the colon-parentheses system for volume, number, and

page citations: "10(2) :5-9." Spell out such words as "figures" and "plates" (or "pages" when

used alone).

For free copies of his own paper, a Smithsonian author should indicate his requirements on

"Form 36" (submitted to the Press with the manuscript). A non-Smithsonian author will receive

50 free copies; order forms for quantities above this amount with instructions for payment will be

supplied when page proof is forwarded.