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U.S.
UNITED STATES DEPARTMENT OF AGRICULTURE
MISCELLANEOUS PUBLICATION NO. 500

WASHINGTON, D. C. February 1943

THE FLEAS
OF NORTH AMERICA

Classification, Identification, and Geographic Distribution
of These Injurious and Disease-Spreading Insects

By
H. E. EWING, Entomologist
and

IRVING FOX, Collaborator

Division of Insect Identification
Bureau of Entomology and Plant Quarantine

For sale by the Superintendent of Documents, Washington, D.C. - - - - Price 20 cents

UNITED STATES
DEPARTMENT OF AGRICULTURE

Miscellaneous Publication No. 500

Washington, D. C. February 1943

THE FLEAS OF NORTH AMERICA

Classification, Identification, and Geographic Distribution of
These Injurious and Disease-Spreading Insects

By H. E. Ewine, entomologist, and Irvine Fox, collaborator, Division of Insect
Identification, Bureau of Entomology and Plant Quarantine.

CONTENTS

Page Page

PERU PGUINECRMSTY 2) heer he ee a 1 | The order Siphonaptera—Continued.
The economic importance of fleas_____________ 3 The family Hystrichopsyllidae Tira-
The structure of the siphonapteran head__-__- - boschi—Continued.
PRHCRm ENN fees AOS Ee ess 2 6 The subfamily Hystrichopsyllinae
The modified terminal abdominalsegments___ 8 auraposehi: =f beret) 2 epee tL 84
The origin end relationships of the Siphonap- The subfamily Macropsyllinae Oude-
Bi net eet ERS 5 24 28 8 UE Ess Se SNe eae ee 10 WANS OS 7 rok ae Ae eo ea 89
The order Siphonaptera_-____- ee eee 13 The subfamily Leptopsyllinae Roths-
Key to the families and subfamilies of ehldiet eas pape A NOS Veg vo 89
Siete CUPS rs eee ae ry le 14 The family Ischnopsyllidae Wahlgren ____ 96
The family Dolichopsyllidae Baker_______ 15 The family Pulicidae Stephens___________ 101
The subfamily Vermipsyllinae Wag- The subfamily Spilopsyllinae Oude-
Vil OS 5 ow) See Se) a eee 15 midnsS- Eos eee 8 Sie bteg h m Ee So, 102
The subfamily Rhopalopsyllinae @u- The subfamily Chimaeropsyllinae,
OIE ECU [Sse en ed oe or ee 20 new Ssubiamilys ss a 115
The subfamily Dolichopsyllinae The subfamily Pulicinae Tiraboschi__ 116
et SO ese A Pa ee eee 23 The family Stephanocircidae Wagner_____ 119
The subfamily Uropsyllinae Oude- The family Hectopsyllidae Baker ________ 119
ir gel seee se 2 ee ee eee 73 | List of synonyms of North American species
The family Hystrichopsyllidae Tira- ang subsperiess: sere? A LW ir eo ie 123
Loot o> Ee ee ee Ja ISTLCLALUT CCl Cd ams = one «SRE rel yee ON os 124
The subfamily Ctenophthalminae Iastrations, (figs. 4=13) 2.0. 2 © E28
vi: ATU tit ft Se ee eee ee Vo) Paxonowicundes ties (4 eee A eet 139

INTRODUCTION

The stress of war and the resulting concentration of people intensify
flea problems and the danger of serious outbreaks of flea-borne dis-
eases, for fleas figure prominently in the health, efficiency, and comfort
of large numbers of people throughout the world. Such diseases as
bubonic plague and endemic typhus are carried to man by these insects,
which also play an important role as pests of livestock and in
dwellings and warehouses.

1 During the preparation of this work the writers have been aided in various Ways by the
following persons: Karl Jordan, of the Tring Museum, England; N. FE. Good and F. M.
Prince, of the U. S. Public Health Service; and E. A. Chapin, of the U. S. National Museum.
They have also drawn to some extent upon data, published or unpublished (correspondence,
notes with specimens, etc.), left by the late N. C. Rothschild, of England; the late Carroll
Fex, of the U. S. Public Health Service: and B. J. Collins, formerly-of the U. S. Public
Health Service. Other specialists on the Siphonaptera have kindly deposited types at the
U. S. National Museum and in this manner have greatly aided in the determination of the
Synonymy of our North American species. Among them are the following: W. L. Jellison
and G. M. Kohls, of the U. S. Public Health Service: C. A. Hubbard, of Pacifie University ;
and Chi-Ying Liu, of National University of Chekiang, Hangchow, China.

1
467459—43—_1

2 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

The proper identification of fleas forms a sound basis for disease
prevention and flea control. This publication, prepared primarily as a
taxonomic revision, will help those called upon to identify the species
of these insects and recommend control procedures. Full descrip-
tions are given of the genera and higher groups, accompanied in
most instances by figures. ~ For species, subspecies, and varieties
formal descriptions are avoided, but data have usually been supplied
for each as follows: Indication of synonymy, name of type host,
type locality, statement relative to range, and remarks upon identifica-
tion characteristics, nature of original description, references to
figures, published notes on biology or economic importance, and
redescriptions. ;

In the preparation of this work the literature has been reviewed up
to the end of 1939, and in addition data from those papers that were
published before October 1, 1940, are included, if the material upon
which they were based was available for study.

Although the fleas of North America have probably been studied
more extensively than those of any other continent, only two large and
comprehensive papers dealing with them have been published. One of
these was by Baker (4)? in 1904, revising all the American fleas, and
the other by I. Fox (25) in 1940, revising the fleas of the eastern part
of the United States. Thus an up-to-date treatment of all North
American fleas is in order. In a limited way a review of the species
of this continent is here presented, it being based largely upon the
extensive collections of the United States National Museum and of
the Bureau of Entomology and Plant Quarantine of the United States
Department of Agriculture.

Since taxonomic categories are concepts, any extensive use of them
should be preceded by definitions or a statement explaining their limi-
tations. The difference between the genus and the subgenus, as these
two categories are here used, lies in the nature of the characters em-
ployed. Characters which, from a morphological standpoint, would
be considered as of only specific importance, if occurring in all species
of a small natural group, are here regarded as being of subgeneric
value. Characters of the terminalia of both sexes, if they have to do
with the presence or absence of a structure or a change in the relation-
ship of one structure to another, are regarded as supplemental generic
characters, or in certain cases these sex characters alone may constitute
the distinguishing characters of a subgenus. In other words, all first
groupings of related species based exclusively on sex characters con-
stitute subgenera. If, however, these sex characters are correlated
with one or more other characters, even though the latter are rather
insignificant, the group of so-characterized species is regarded under
most conditions as a genus.

The differences implied between a subspecies and a variety, as the
two terms are here employed, have to do not so much with the con-
spicuousness or morphological importance of the characters used as
with their variation. Minor characters which are continuously vari-
able, such as coloration (amount of pigmentation), length of certain
setae, or the shape of some part of the terminalia, are considered of
varietal importance if the variations are correlated with geographical

2 Italic numbers in parentheses refer to Literature Cited, p. 124.

THE FLEAS OF NORTH AMERICA 3

or host distribution. However, if there is a break in the variation
of these characters, they are considered of subspecific importance re-
gardless of whether the differences are correlated with geographical
or host distribution. A subspecific character is considered as usually
being morphologically more distinctive than a varietal character,
and by that token a subspecies is regarded as taxonomically of more
_importance than a variety, although from the nomenclatorial point of
view the two are treated alike.

Characters used for specific purposes less frequently refer to colora-
tion, or length of setae, or even size. They are regarded as essentially
the same as those used for the subspecies, except that they are more
obvious, less variable, or have a greater morphological significance.
Characters resulting from hybridization or local environmental in-
fluences are not here considered as good specific characters.

In preparing this taxonomic review an attempt has been made
to follow a conservative policy in the treatment of the different cate-
gories, avoiding alike the extremes of the “splitter” and those of
the “lumper.” It is the writers’ opinion that in the Siphonaptera the
tendency has been toward too much “splitting.” This is strikingly
shown by the manner in which the genus Ceratophyllus Curtis has
been split up in recent years. On this Jordan (43, p. 70) has com-
mented as follows:

* * * The splitting-up of Ceratophyllus has begun, and there is no means of
stopping the process of dissolution from going on. * * *- If I here inflict on
science a number of new generic names, I plead the excuse that the divisions
of Ceratophyllus I here define would inevitably be named by somebody else,
probably by someone who does not know these fleas, as has happened in other
orders of animals.

In regard to the splitting up of Cerutophyllus and certain other
genera the present authors have dealt rather firmly. Many of the
segregates are retained as subgenera, being in reality excellently
fitted for such a rank, while others are reduced to synonymy.

In the part of this publication dealing with the taxonomy of
species, subspecies, and varieties the paragraph headings “Cotype
Hosts” and “Cotype Localities” have been introduced in those in-
stances in which cotype material came from more than one host and
more than one locality.

The number of described genera and species of North American
fleas has increased very rapidly in recent years. In this paper 61
genera, 14 subgenera, 209 species, and 63 subspecies and varieties of
fleas are listed from North America and the West Indies.

THE ECONOMIC IMPORTANCE OF FLEAS

Fleas are of economic importance in two ways. First, by their
direct attacks on man and his domestic animals they cause irritation,
loss of blood, and in severe cases the loss of life. Second, because
of their blood-sucking habits they are important transmitters of
certain internal parasites and disease-producing organisms.

Fleas are probably best known as household pests. Their presence
in houses, under certain conditions, may cause the occupants great
annoyance. Their occurrence there is due to the presence of their
natural hosts, either in the houses or in nearby quarters. These are

4 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

such domesticated animals as cats, dogs, and pigs. During the early
part of the summer the fleas lay large numbers of eggs on their hosts,
which soon drop off into cracks of the flooring or into debris of vari-
ous kinds. Here the larvae hatch and feed, grow to larval maturity,
and transform into quiescent pupae. Then about the middle of the
summer, frequently near the time when the occupants of the infested
dwelling may be returning from a summer vacation, large numbers
of the second generation of adults appear. It is this generation
which, because of its great numbers, usually causes so much distress.

In the United States fleas are serious pests of dogs, cats, and
poultry. Dogs may be greatly annoyed by fleas and when attacked
spend much time gnawing at the affected parts, particularly the back
near the root of the tail. As a result these parts become raw and
sore, and infection often follows.

Poultry are frequently infested by fleas, and the sticktight flea of
the South attaches permanently to the heads of its hosts. Here it
sucks the blood and causes much irritation. Young chickens may
become so heavily infested that they die from the attacks of these

ests.

: Some fleas are of much economic importance because they are the
transmitters of certain diseases. This is particularly true of those
that transmit plague, one of the worst scourges of man. In the
United States, as well as in many foreign countries, the oriental rat
flea (Xenopsylla cheopis (Rothschild) ) is the most efficient carrier
of the bacterial organism of plague. Also in this country four other
species that readily feed on man are thought to be occasional car-
riers. They are the human flea (Pulex irvitans Linnaeus), the cat
flea (Ctenocephalides felis (Bouché) ), the dog flea (Ctenocephalides
canis (Curtis)), and the rat flea (Vosopsyllus fasciatus (Bosc) ).

Plague organisms, developing in the “gizzard” of the oriental rat
flea, frequently cause an obstruction so that when the flea attempts
to feed again regurgitation results. It is this regurgitation that is
believed to be generally responsible for the transmission of the
organisms to man.

Plague first appeared in the United States in California, in 1900,
and within 10 years was found to exist in native ground squirrels in
several counties. Among these hosts the plague spread slowly until
today diseased rodents have been found in the States of Oregon,
Washington, Nevada, Idaho, Utah, New Mexico, and Montana. The
squirrels act as reservoirs for the disease, and the fleas, becomin
infected with the bacteria obtained from these rodents, may later feed
on rats or man and thus increase the spread of the plague.

Fleas also are vectors of endemic typhus. This disease is a mild
form of typhus found in several of the warm countries of the world.

THE STRUCTURE OF THE SIPHONAPTERAN
HEAD

In many of the more generalized fleas, such as Ctenocephalides
canis or Hoplopsyllus affinis, the division of the forehead into a front
and a pair of genae is evident. The front extends, with its outwardly
curved anterior margin, down to the anteroventral angle of the head.
The genae form the lower margin of the head and extend upward at

-

THE FLEAS OF NORTH AMERICA o

least to the antennal grooves. They fuse anterodorsally, without a
delimiting mark of any kind, with the front.

The front, which is so conspicuous in Ctenocephalides (fig. 1) and
many other genera, becomes so reduced in certain families that it
is very inconspicuous, and in Craneopsyllus wolff huegeli it is vestigial,
having been apparently overlooked in the past. As the front be-
comes smaller, the genae become increasingly larger. This process
has gone so far in the family Stephanocircidae that almost all the
forehead is composed of the greatly enlarged genae.

If it were not for the fact that the front usually can be identified
by two of its structural features, the frontal tubercle and the dermal
pits, confusion frequently would exist in regard to the part it takes in

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Ficure 1.—Head of a flea, female of Ctenocephalides felis, with the different
parts labeled.

the composition of the head. The frontal tubercle, although sometimes
absent, is a well-known structure on the anterior margin of the front.
Usually it is sharply angulate at its apex and is situated in a small
notch, the frontal notch. The dermal pits are small paired pits (fig. 1)
which usually are located near the anterior margin of the front. The
maximum number of pairs on the front is three, as in such genera as
Hoplopsyllus, Actenophthalmus, and Sternopsylla. In Neo psylla,
Ctenophthalmus, and many other genera there are two pairs of dermal
pits on the front. In Crazeopsylla only a single pair persists. The
presence of any of these pits always identifies the front.

When the genae become greatly enlarged at the expense of the front
there is a tendency for each gena to become differentiated into an
anterodorsal and a posteroventral part. This differentiation has pro-

6 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

gressed so far in the family Stephanocircidae that the two parts are
completely separated. At the same time the anterodorsal parts of the
genae still retain their fusion with the front, and these three fused
sclerites form a conspicuous crownlike structure known as the helmet.
Such a profound change in the basic structure of the head sets apart
the family Stephanocircidae very distinctly from all the other families
of fleas.

In the figure of the head and thorax of Ctenocephalides canis given
by Ewing (16, p. 154) the unpaired piercing element of the mouth
parts was labeled, following the practice of certain former workers,
the hypopharynx. This structure has been called the labrum-epiphar-
ynx by some workers, but it should be called the labrum (fig. 1), since
there is no evidence that the epipharynx enters into its composition.

The postantennal portion of the head of a flea (fig. 1) has long been |
known as the occiput, or occipital region. These terms, however, should
be restricted in their application to the posterior part of this post-
antennal region. The part of the head immediately posterior to the
front must be the epicranium. Since the epicranium and the occiput
are not separated by an occipital suture, the extent of each is not
definitely known. The setae situated in oblique rows on the postan-
tennal portion of the head in some fleas may be called the postantennal
setae. Those of the posterior vertical row (fig. 1) may be called the
occipital setae, or setae of the posterior, marginal row.

THE fHORAx

Jordan and Rothschild (47, p. 23) have given an exposition of the
external structure of the thorax of a flea. The description here given
differs from that of those authors largely in the interpretation of the
lateral sclerites of the metathorax.

The body of a flea, including the thorax, has become greatly com-
pressed or laterally flattened. As a result the ventral sclerites have
assumed largely or completely a lateral position. Although much
modified in shape by compression, the generalized nature of the thorax
is easily discerned. It is divided into three distinct segments (fig. 2),
the prothorax, mesothorax, and metathorax, all of which are movably
articulated into the body “chain” of segments. The upper part of each
thoracic segment is covered with a single, undivided sclerite. These
are the pronotum, mesonotum, and metanotum. The posterior margin
of each of these dorsal thoracic sclerites overlaps the anterior margin
of the following dorsal sclerite in the same manner as do the dorsal
abdominal sclerites.

The sternal and pleural sclerites of each thoracic segment have be-
come either fused or united (fig. 2) so as to make a single unit. Such
a unit or section of a thoracic segment is typically made up of a ventral
sternum, and on each side an anterodorsal episternum and a postero-
dorsal epimeron. |

The sternal and pleural sclerites of the prothorax typically are fused
into a movable saddlelike mass (fig. 2), the sternopleura. This sterno-
pleura extends forward and has its longest dimension in an almost
horizontal direction.

The sternal and pleural sclerites of the mesothorax (fig. 2) have
a position and relationship more commonly found in other insects.

THE FLEAS OF NORTH AMERICA v4

The sternum is ventral, and in lateral view usually appears to be
the smallest of the sclerites. It is the most heavily pigmented. In
nearly all genera it is without demarcation from the episternum,
but in some genera, such as Stenistomera and Cratynius, the meso-

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sternum is separated from the mesepisternum on each side by an
inner chitinous ridge, a character of generic importance. The mes-

episternum similarly may be separated from the mesepimeron by an

inner chitinous ridge (fig. 2), the pleural ridge, in many genera of

8 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

fleas such as Ctenocephalides, Spilopsyllus, and Chimaeropsylla; or
the two sclerites may be completely fused, without demarcation, as
in Moeopsylla, Pulex, and others. This character has been used in
separating the subfamily Pulicinae from the other subfamilies of
Pulicidae.

The sternopleural sclerites of the metathorax (fig. 2) show the
greatest diversification of those of any thoracic segment. Though
typically of a similar arrangement to those of the mesothorax, in
many genera the following important diversifications may be noted:

(a) The metepisternum may be divided into an upper and a lower
part by an internal chitinous ridge as in the genus fectofrontia.
This division may be continued to such a degree that the upper part
of the metepisternum becomes detached from the lower part and
transferred to the metanotum, being fused with the latter as in the
subgenus Opisocrostis. |

(b) The metepimeron may be so expanded (fig. 2) as to be the
largest of all the laterally placed sclerites of the body.

(c) The metepimeron usually is entirely free from the metanotum
and is clearly separated from the metepisternum by an internal
chitinous ridge, but in the genus Rectofrontia the metepimeron be-
comes completely fused, without demarcation, with the metanotum,
while in the genus Opisocrostis the metepimeron is movably articu-
lated with the metanotum by means of a condyle. Although usually
separated from the metepisternum by an inner vertical ridge, in the
genus Trichopsylloides the metepimeron is completely fused, without
demarcation, with the former sclerite along its upper anterior end.

THE MODIFIED TERMINAL ABDOMINAL
| SEGMENTS

The abdomen of a flea usually has been regarded as being com-
posed of 10 segments, but according to Wagner (81) there are in
reality 12. Wagner’s contention rests to a large extent upon whether
the “cerci” of a flea (present in females) represent the true cerci.
If they do, they indicate the presence of the eleventh segment. This
follows since embryological studies have shown that in general the
cerci of insects represent the appendages of the eleventh segment.
Behind the segment in a flea that bears the stylets, or “cerci,” there
are always present sclerites which Wagner believes represent those
of the twelfth abdominal segment. There are reasons, however, for
not regarding the stylets as true cerci. In the first place they not
only are restricted to the female sex, but are wanting in this sex in
certain genera. Secondly, it has been shown by the recent study of
the embryology of the fleas by Kessel (53) that there are 10 posterior
pairs of ganglia belonging to the abdominal region; while the caudal
segment has no ganglion. Thus it would appear that there are 11
abdominal segments in the flea embryo and probably the same num-
ber in the adult. If this reasoning is correct, then the anal seoment
of Wagner is the eleventh abdominal and the stylets are on the tenth,
while true cerci are wanting.

The first seven segments of the abdomen usually are unmodified in
both sexes.

a

THE FLEAS OF NORTH AMERICA 9

The eighth abdominal segment is practically unmodified in the
males of some genera. In the males of most genera, however, the
eighth sternal plate is much modified. It may be greatly reduced

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FIGURE 3.—Modified terminal abdominal segments of a male flea, Nosopsyllus
fasciatus, with parts separated from one another so as to show their struc-
ture more clearly.

in size, or it may be extended into a long, posteriorly directed process
or have other modifications. The eighth sternal plate may be some-
what rudimentary in the female.

10 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

The ninth abdominal segment is of greatest interest both morpho-
logically and taxonomically because in the male (fig. 3) 1t bears what
is believed to be a pair of segmental appendages, the clasping ap-
paratus. The ninth tergal plate is either fused with the pygidium
in both sexes, or, in the male, it may be fused with the dorsal plate
of the clasping apparatus. Most of the ninth sternal plate of the
male usually consists of a modification and elongation (which may
be rather complicated) of the overlapping posterior margin of this
plate. The same sternal plate is rudimentary or absent in the female.

The clasping apparatus of a flea, which is commonly regarded as
representing a pair of segmental appendages, consists of the follow-
ing parts on each side: The basal body, which is united with the body.
of the other side by the dorsal plate, an inwardly and forwardly
projecting arm known as the manubrium (fig. 3), a dorsal process,
and a posterior movable element known as the movable finger. Wag-
ner regards the basal body, with its movable finger, as representing
a gonopod.

The tenth abdominal segment usually is regarded as being repre-
sented by the pygidial plate. According to Wagner this plate is fused

with the tergal plate of the twelfth segment in the female, and in the

same sex the sternal plate of the tenth segment is absent. The tenth
sternal plate usually is absent in the male, but, according to Wagner,
may be present in the form of a subpygidial sclerite.

The eleventh segment, according to Wagner, has no tergal sclero-
tizations, but the sternal plate may be the subanal plate in the male,
and the “cerci” of the female may be rudiments of the ventral part
of the eleventh segment shifted to a dorsal position.

The twelfth segment, according to Wagner, is the so-called anal
seoment, consisting of a simple tergal and sternal plate in the female
and a simple tergal plate and two cercuslike sternal sclerites or a single
plate in the male.

It appears to the writers, however, that the anal sclerites of the
so-called anal segment are more likely comparable to the epiproct and
paraprocts of thysanurans and orthopterous insects, in which case they
should represent the sternal and tergal plates of segment XI of the
abdomen.

THE ORIGIN AND RELATIONSHIPS OF THE
SIPHONAPTERA

The origin and the relationships of the Siphonaptera have been
moot questions ever since the study of taxonomic entomology began,
and during the last century many papers were published dealing with
these subjects. Unfortunately the theories advanced were based not
only upon wholly insufficient and largely faulty interpretations of the
morphology of the fleas themselves but also of the morphology of in-
sects in general. Those who are interested ina summary of the theories
of these earlier workers are referred to a paper by Packard (61) pub-
lished in 1894 under the title On the Systematic Position of the
Siphonaptera, with Notes on their Structure. The prevailing ten-
dency of these early workers was to regard the fleas as specialized
Diptera or specialized Hemiptera. Some of them, however, considered
the fleas as a distinct order having relationships with either the Dip-
tera, the Hemiptera, the Coleoptera, or the Hymenoptera.

———

THE FLEAS OF NORTH AMERICA ~ Met

Packard himself, although giving an interpretation of the mor-
phology of the fleas now known to be in error in many respects, had
this to say about their relationships (61, p. 354) :

The larva of the Siphonaptera apparently presents the nearest approach of any
of the insects now existing to the shape of the primitive Diptera. It is certainly
a more perfectly developed larva as regards its external structure and also the
position of the brain than any dipterous one, and approaches nearer to our con-
ception of the primitive, ancestral, dipterous larva than any other form.

Packard further says:

As regards the phylogeny of the Siphonaptera we can only say, with our present
imperfect knowledge of their embryology, that they seem to stand nearer to the
Diptera than to any other order, and that they must have diverged from the
ancestral dipterous stem before the existing forms of Diptera had become so
extremely specialized as we now find them to be.

Dahl (9) published in 1898 an interesting paper on the systematic
position of the Siphonaptera. He would derive the fleas, the phorids,
and the dipterous genus Scatopse from a common hypothetical stem
which he called the Archiscatopse. Although, as pointed out by
Ewing (14, p. 348) in 1924, Dahl’s—
interpretation of the degeneration of the Phorid compound eye into the minutely
spined area found in some fleas just above the antennal fossa, is certainly very
speculative * * #*
yet his theory is one of the best proposed, and he has given many data
in support of it. Particularly convincing is his demonstration of the
apparently complete homology between the antenna of certain male
fleas and that of Scatopse. Not only does the antenna of the male flea
have the same two basal segments, but the pseudosegments in the long
club are the same in number as those distad of the second segment of
the antenna in Scatopse.

Martini (56) in 1922 revived the old theory of the basic relationship
of the fleas with the Coleoptera. He pointed out the many homologies
between fleas and the genus Oxytelus and gave figures representing
cross sections of a flea and an oxyteline in which the fundamental
structures are homologized. It should be noted that some of the homol-
ogies mentioned by Martini as existing between structures of fleas on
the one hand and those of the Staphylinoidea on the other are of a
very general nature and doubtless exist between fleas and many other
kinds of insects.

Crampton (7, p. 324) has pointed out the relationships of fleas to
the Trichoptera and states that the latter—

could readily serve as the types ancestral to fleas since they have three-segmented
labial palpi, their prothoracic sclerites are as much like those of the fleas as any
insects, the mesothoracic and metathoracie coxae of Trichoptera are divided into
a eucoxa and meron, and the Trichoptera have cerci and terminal structures sug-
gestive of those of the fleas.

He opposed the theory that fleas could be derived from the Diptera
because— |

The labial palpi of fleas are frequently composed of at least three segments,
while the labial palpi of Diptera (labella) and Mecoptera are never composed
of more than two segments, and therefore could not serve as the prototypes of
the labial palpi of fleas.

He also opposed the theory of the origin of fleas from the Diptera
because the latter have no meron in the metathoracic leg.

12 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

Crampton assigns to the fleas the rank of a superorder, it being
one of the three superorders into which holometabolous insects are
divided. His arrangement of these superorders follows: The
Pansiphonaptera, including the Siphonaptera only; the Panmecop-
tera, including the Neuroptera, Mecoptera, Diptera, Lepidoptera, and
Trichoptera, with their fossil allies; the Pancoleoptera, including the
Coleoptera, Strepsiptera, and Hymenoptera, with their fossil allies
except the Protohymenoptera.

The theory that fleas have been derived from the Trichoptera was
assailed by Tillyard (74, p. 39) who stated that—

There is not a shred of evidence in favor of Fleas having been derived from
an aquatic ancestral type. ... For Fleas to have been derived from Tri-
choptera, it would be necessary to postulate the loss of the ancestral functional
mandibles and the evolution of the present type of elongated spear-like organ
from the nonfunctional vestige of the Trichoptera! This also involves a belief
in the loss and subsequent reappearance of the mandibular muscles, and contra-
venes Dollo’s Law..

* * * Wleas are smooth and shiny, very unlike the soft-haired Trichoptera in
their body-covering. What hairs they possess are specialized into stiff bristles
and combs.

Tillyard (74) claimed that the ancestors of the Siphonaptera must
have possessed short antennae, elongated mandibles, four-segmented
maxillary palpi, a smooth and leathery body, a metathorax not highly
reduced, middle and hind coxae with a meron, and a terrestrial larva
feeding on animal or vegetable débris. The pupa must have been
a. pupa libera, either in a cocoon or a primitive earthen cell. He
believed that the only ancestors that meet these conditions are—

(a) a small, reduced type of primitive Mecopteron, probably of the Upper
Permian family Permochoristidae, or (b) a related form classifiable definitely
within the Paratrichoptera.

The objections of Crampton (7) and Tillyard (74) to deriving
fleas from ancestral Diptera because of the absence of a meron from
the third coxa of the latter is not well founded. Many Diptera have
the second coxa with a well developed meron, although both coxa and
its meron may be partly or largely transferred to the body wall.
Such is the case in the Tabanidae. Other Diptera have the third
coxa well developed, with the evidence indicating that there has been
a reduction and fusion of the eucoxa and the meron.

A condition somewhat similar to this is found-in certain fleas, as,
for example, those of the genus E'chidnophaga. In fleas of this genus
the meron of the mesocoxa is reduced but separated from the eucoxa
by a groove, while that of the metacoxa is partly fused with the
eucoxa. Further, it is to be noted that the third coxa of some phorids
is not very different in its major aspects from the third coxa of cer-
tain fleas, and although there is no suture or ridge dividing it into a
true eucoxa and a meron, there exists an undifferentiated area which
probably represents the meron. ea:

The writers believe that much printed space has been wasted dis-
cussing the value of the labial palpus as a structure indicating the
origin of the Siphonaptera. The segmentation of these appendages
is such that they could have been evolved from those of almost any
group of free-living insects. The most common number of true seg-
ments in the labial palpus of a siphonapteron is five. Yet there are
several genera with four, three, or two segments in each palpus.

THE FLEAS OF NORTH AMERICA 13

Fleas like those of the genus Vermipsylla have many so-called seg-
ments in the labial palpus. Most of these evidently are not true seg-
ments, but pseudosegments, as is indicated by the fact that they are
very short and frequently only partly separated from one another
by sutures that are incomplete and not transverse.

The present writers would derive the fleas and-the Diptera from a
common ancestral stock. However, the fleas are to be regarded as
older and more primitive than the Diptera. There are two reasons
for this, one being that the flea larva is decidedly of a more gener-
alized type than the larvae of most of the Diptera, and the other,
that fleas lack reduction and modification of the metathorax such as
that found in flies. Fleas probably developed from ancestors that

et had functional metathoracic wings and hence were not true
iptera.

~The Order SIPHONAPTERA

Small insects, with a complete metamorphosis; parasitic in adult state on
warm-blooded vertebrates; without wings or wing vestiges; body greatly com-
pressed and frequently provided with 1 or more combs of spines.

Head always divided into 2 regions by an oblique antennal groove and in
some genera also divided into an anterior and a posterior part by the presence
of a deep fronto-epicranial groove. Antennae short, stout, each contained in a
deep sulcus known as the antennal groove; segments beyond the second greatly
shortened and formed into a club. Eyes simple, consisting of a single pair;
degenerate or absent in some genera. Mouth parts adapted for piercing and
bloodsucking. Maxillae each with a filiform, segmented palpus and a sub:
triangular, ventrally projecting, acute, maxillary lobe. Labrum long, flattened.
almost needlelike, together with the mandibles forming a piercing beak.
Mandibles long, piercing blades, with parallel, saw-toothed margins. Labium
with greatly reduced base, but with segmented palpi, grooved on inner surfaces,
the 2 fitting together to form a labial beak.

Thorax composed of 3 freely moving segments. Prothorax with large prono-
tum, frequently bearing a transverse comb of spines; sclerites of sternal and
pleural regions anchylosed, forming a saddlelike structure bearing the front
pair of legs. Mesothorax larger than prothorax. Mesonotum usually somewhat
expanded, never with comb of spines. Mesosternum, mesepisternum, and
mesepimeron anchylosed, but the boundaries of union usually indicated by
internal sclerotized ridges. Metathorax similar to mesothorax except that
the metasternum is greatly reduced, the metepisternum is divided into 2 parts
by a horizontal suture, and the metepimeron is greatly expanded posteriorly.

Abdomen apparently composed of 10 segments, but in reality of 11. Segment
‘I greatly reduced ventrally, without sternum. Tergal plate VII usually with
a clump of 1 to 4 large or stout bristles (antepygidial bristles) on each side
at posterior margin. Dorsum IX with a specialized, saddlelike plate known as
the pygidium, which is studded wtih minute setae. In female, segment XI
with a pair of small, 1-segmented stylets (? cerci) situated dorsally. In
males of most genera sternal plate VIII modified, being either greatly reduced
in size or extended posteriorly into a long process of a variety of shapes.
Sternal plate VIII in the female occasionally somewhat rudimentary. Ninth
abdominal segment in male bearing a pair of claspers. Sternal plate IX in
male greatly modified, consisting usually of a pair of inner vertical processes
and a long, highly modified, posterior or horizontal process or lobe. Male flea
with a complicated protrusible penis and female with 1 or 2 receptacula seminis.

Legs typically insectan. Each with a large, long, flattened coxa; a small,
ringlike trochanter; a large femur; a tibia enlarged distally; a 5-segmented
tarsus, and each tarsus terminated by a pair of subequal, curved claws.

The order contains six families as follows: Dolichopsyllidae,
Hystrichopsyllidae, Ischnopsyllidae, Pulicidae, Stephanocircidae,
and Hectopsyllidae.

14 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

Besides the name Siphonaptera, which should be attributed to
Latreille, two other names have been commonly applied to the fleas,
Aphaniptera Kirby and Suctoria Retzius. In the selection of the
names of orders more consideration should be given to appropriate-
ness and usage than to priority. Hence, in this work the term
Siphonaptera has been used. _

The fleas have been divided into from three to about six families
by most modern workers. All workers recognize the families Puli-
cidae and Ischnopsyllidae, and two other families are usually recog-
nized, ohne containing the genus Ceratophyllus and related genera and
the other containing the degenerate genus 7unga and related genera.

Wagner (S86) has surpassed all others in the number of families
and subfamilies he has recognized for the order. He recognizes 10
families and 28 subfamilies, in addition to many tribes. Since many
of these subfamilies and tribes contain but a single genus each and
many are hard to separate properly on morphological characters, a
more conservative and usable classification appears desirable.

Key To THE FAMILIES AND SUBFAMILIES OF SIPHONAPTERA

1. Thorax not greatly reduced, thoracic terga taken together longer than
first abdominal tergum; gravid females moderately, if at all,
enlar Cedi 2 he Me Ae eR ee apr Si cA oe Pee ee 2

Thorax greatly reduced, thoracic terga taken together shorter than
first abdominal tergum; females sometimes with a reduced number
of abdominal spiracles, and when gravid, abdomen greatly dis-
Temes SFE we a a he ee Hectopsyllidae Baker.
2. Gena not divided by a vertical suture; head without helmet________ 3
Gena greatly enlarged, being extended dorsally in front of antennal
groove and divided by a vertical suture; head with a helmet com-
posed of the greatly reduced front and anterior parts of the greatly
enlarged genae; postantennal region of head with a dorsal incragssa-

COT ee Re ee eee eee Stephanocircidae Wagner.
3. Typical abdominal terga each with more than 1 transverse row of
setae: fronto-epicranial groove frequently present__________________ 4

Typical abdominal terga each with but 1 transverse row of setae;
fronto-epicranial groove usually absent; eyes usually present.
Pulicidae Stephens 12
4. Head seldom elongated, ventral flaps absent. Not parasitic on bats._. 5
Head usually elongated, always with 2 or 8 ventral flaps on each side
adjacent to fronto-genal angle. Parasitic on bats.
Ischnopsyllidae Wahlgren.
5. Cephalic combs absent; front not reduced; abdominal combs rarely
present; abdominal apical spines rarely numerous.
Dolichopsyllidae Oudemans’ 6
Cephalic combs present; front frequently reduced; abdominal combs
frequently present; abdominal apical spines frequently numerous.
Hystrichopsyllidae Baker 9
6. Frontal notch or frontal tubercle, or both notch and tubercle, present

(although ‘sometimes vestie i ail ye 25ers eee eae cf
_Frontal notch and frontal tubercle absent, there being no vestige of
eltHe Taos ees Zee Raed ee Aes en Uropsyllinae Oudemans.
7. Antepygidial bristles absent or very poorly differentiated; abdomen
frequently distended in female___.____________ Vermipsyllinae Baker.
Antepygidial bristles present, conspicuous; abdomen not distended;
labial palpus usually, with 5 cor Jess “Sesments=226 =" 2 es oe 8
S$ Eronotal .comb absent 5." she 2 ee Rhopalopsyllinae Oudemans.

Pronotal comb..present = fea eee Dolichopsyllinae Oudemans.

THE FLEAS OF NORTH AMERICA iS

9. Front never greatly reduced, forming at least one-half of anterior
margin of head; genae usually not greatly enlarged, and usually
longest horizontally; eyes sometimes present__--_____---_--_------ 10

Front greatly reduced, dorsal or anterodorsal in position, forming less
than one-half of anterior margin of head, possessing 1 or 2 pairs of
dermal pits and never more than a vestigial frontal tubercle; genae
greatly enlarged, longest in a vertical or nearly vertical direction ;
Sees VeEStISiell (OF -nDSCMt 8 la = Sa Leptopsyllinae Rothschild.

10. Labial palpus with few segments (not over 8); cephalic comb seldom
extending along both ventral margin of gena and anterior margin
TuMETTD Rie i Neg) eee HLL coe eee are gs

Labial palpus with many segments (more than 10); cephalic comb
extending along ventral margin of gena and anterior margin of
antennal groove; female with 2 receptacula seminis, 1 smaller than
the other. Found only in Tasmania______-___ Macropsyllinae Baker.

11. Frontal tubercle or frontal ncetch present, although either or both may
be vestigial; genal comb never with more than 6 spines. Small to
medi sven heas os Ctenophthalminae Rothschild.

Frontal tubercle and frontal notch absent; genal comb sometimes with
more than 6 spines. Medium-sized to giant fleas.

Hystrichopsyllinae Tiraboschi.

12. Mesepisternum separated from mesepimeron by an internal, vertical
GT Pi oe ae eo geet ee TS es Oe ee eee eee 13

Mesepisternum not separated from mesepimeron_-_Pulicinae Tiraboschi.

13. Fronto-epicranial suture indistinct or absent; postantennal region of
head but slightly, if at all, incrassate dorsally.

Spilopsyllinae Oudemans.

Fronto-epicranial suture well developed; postantennal region of head
dorsally incrassate; front considerably reduced, bringing fronto-genal
angle up almost to level of eyes. Found only in Africa.

Chimaeropsyllinae, new subfamily.

The Family DottcHopsyiiipar Baker

Head seldom divided into an anterior and a posterior part by presence of fronto-
epicranial groove. Genae not greatly enlarged and not divided by suture. Cephalic
combs absent. Forehead without helmet, seldom elongated, and always without
ventral flaps. Eyes either well developed, vestigial, or absent, Thorax not greatly
reduced ; thoracic terga taken together longer than abdominal tergum I, Typical
abdominal terga each with more than 1 transverse row of setae. Abdominal apical
spines frequently present but rarely numerous. Abdominal combs absent.
Females without a reduced number of abdominal spiracles, and when gravid not
greatly distended,

This family contains four subfamilies, as follows: Vermipsyllinae,
Rhopalopsyllinae, Dolichopsyllinae, and Uropsyllinae.

The subfamily Dolichopsyllinae was established by Baker (4, p. 127)
in 1905, and in 1909 Oudemans (59, p., 156) raised it to family rank,
attributing the family name to Baker. The family Ceratophyllidae
Rothschild (66, p. 58), based on Ceratophyllus Curtis, should be re-
garded asasynonym of Dolichopsyllidae. The family as here described
does not include the subfamily Ctenophthalminae which was included
in it by Ewing (15, p. 166) 11929. There is some doubt as to whether
this subfamily constitutes a natural group, but its genera can readily
be placed according to the arrangement given in this publication.

The Subfamily VERMIPSYLLINAE Wagner

Head not divided into an anterior and a posterior part by a frontoepicranial
groove. Frontal tubercle present. Eyes present or absent. Labial palpus long
and usually with more than 6 segments. Pronotal comb present in only 1 genus
(Trichopsylloides). Antepygidial bristles absent, or very poorly differentiated.
Abdomen of gravid females sometimes considerably distended.

16 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

The subfamily Vermipsyllinae contains four genera, all but the
type genus being represented in the North American fauna.

The subfamily dates from 1889, when it was established by Wagner
(77, p. 205). It has been recognized as a family by Dalla Torre (70, p.
18) in his catalog of the fleas, and likewise by Wagner (80, p. 39)
in the latter’s catalog of Palaearctic Siphonaptera. In the structure
of the head Vermipsyllinae has much in common with Pulicidae, but
the characters of the body proper ally it with the Dolichopsyllidae.

Key to THE NortH AMERICAN GENERA OF VERMIPSYLLINAE

1.) Pronotal comb absent 3.eyes present-_2_ *s = ee ae 2
Pronotal comb present ; eyes absent. Known only from the western part
ofthe United States-—-- == 22. see Trichopsylloides Ewing.
2. Labial palpus with less than 7 segments______-____ Chaetopsylla Kohaut.
Labial palpus with 7 to 10 segments__=~-—~___=____ Arctopsylla Wagner.

The Genus CuHartTopsyLia K ohaut
(Fig. 4, EZ)

Chaetopsylla Kohaut, 1903, Allattani K6zlemények 2:37. Type, Chaetopsylla
rothschildi Kohaut (by subsequent designation).

Frontal tubercle small to vestigial, not angulate but buttonlike. Antennal
club either rounded or somewhat pointed at apex. Eyes somewhat reduced,
poorly pigmented near center. Labial palpus short, with less than 7 segments.
Postantennal region of head with 2 or 8 oblique rows of setae. Pronotal comb
absent. Upper sclerite of metepisternum separated from metanotum by an inner
pigmented ridge. Metepimeron somewhat reduced and sometimes anchylosed to
Some extent with upper sclerite of metepisternum. Abdomen of female with
somewhat enlarged spiracles and when gravid sometimes considerably distended.
Antepygidial bristles absent (not differentiated). Sternum VIII of male large,
broad, practically undifferentiated; sternum IX of male narrowed, elongated,
considerably modified.

The name 77ichopsylla frequently has been used instead of Chaetop-
sylla Kohaut, the latter being considered as a synonym of the former.
But according to the rules of nomenclature there is apparently no
Justification for such usage. The type species of TZ'richopsylla
Kolenati was first selected by Baker (4, p. 371) in 1904 as
pencilliger Grube. Jordan and Rothschild (50, p. 63) in 1920
pointed out that Kolenati, when he established the genus
Trichopsylla, had misidentified penicilliger Grube, one of the in-
cluded species, and stated that—

His penicilliger has no pronotal comb and therefore is not penicilliger Grube ;
from the descriptions of the genus and the species it is evident that the species
erroneously identified as penicilliger is a Vermipsylla.

Hence Jordan reduced Vermipsylla Schimk. to synonymy with J7ri-
chopsylla Kolenati. At the same time Chaetopsylla Kohaut and On-—
copsylla Wahlgren were sunk as synonyms of Trichopsylla.

Wagener (80, p. 39) in 1930 did not accept this synonymy but claimed
that Chaetopsylla Kohaut wasa valid genus. He identified penicilliger
Grube as a species of Ceratophyllus Curtis, and stated that the system-
atic position of the species Kolenati identified as penicilliger Grube
was unknown. It was identified by Jordan (42, p. 293) in 1932 as
Trichopsylla homoeus Rothschild and listed as the genotype of T'7i-
chopsylla Kolenati. Again Jordan sunk Oncopsylla Wahlgren and
Chaetopsylla Kohaut as synonyms of 7'richopsylla.

THE FLEAS OF NORTH AMERICA 17

This substitution of Trichopsylla homoeus Rothschild for 7. peni-
cilliger Grube as the type of 7'richopsylla can hardly hold under Opin-
ion 65 of the International Commission on Zoological Nomenclature.
The “special Committee for recommendation as to action,” whose re-
port was adopted by the Commission and ordered published, stated :

The Committee is of the opinion that as a specimen is the type of a species, so
a species is the type of a genus, and hence that when an author names a par-
ticular species as type of a new genus it is to be assumed that it has been cor-
rectiy determined. If a case should present itself in which it appears that an
author has based his genus upon certain definite specimens rather than upon
a species it should be submitted to the Commission for consideration.

The genus Chaetopsylla is given first place in this work because
preference is desired for it over Arctopsylla as the most generalized
genus of fleas. The point at issue between the two genera is the num-
ber of segments in the labial palpi, there being a smaller number in
Chaetopsylla. The most common number of segments in the labial
palpi of the more generalized insects appears to be three. In fleas it
is five, but four or two are not uncommon. In those genera in which
the number of segments has been increased to more than five the
evidence of secondary segmentation is easily detected. As pointed out
by Tillyard (74, p. 40)—
if we examine the labial palpi of those types which possess a large number of

palpal segments, we are forced to the conclusion that these supposed segments
are actually only secondary annulations of more primitive segments * * *,

CHAETOPSYLLA FLORIDENSIS (I. Fox)

Trichopsylla floridensis I. Fox, 1939, Wash. Ent. Soc. Proc. 41: 45, pl. 6, fig. 6.
Chactopsylla floridensis Jellison and Good, 1942, U. S. Pub. Health Serv. Natl.
Inst. Health Bul. 178: 40.

Type host—Unknown.

Type locality —Gainesville, Fla.

Range.—Known only from the type locality.

The original description of floridensis was based on specimens taken
in garden truck leafmold and is accompanied by a drawing of the
male terminalia. This species differs from Chaetopsylla lotoris
(Stewart) particularly in the chaetotaxy of the head and met-
epimeron.

CHAETOPSYLLA GLoBIcEPs (Taschenberg)

Pulezx globiceps Taschenberg, 1880, Die Flihe, p. 66, pl. 2, figs. 10, 10a, and 11.
a lea globiceps Wagner, 1930, Katalog der palaearktischen Aphanipteren,
p. 40.

Cotype hosts—Vulpes vulpes (L.) (fox) and Taxidea taxus
(Schreber) (American badger).

Type locality.—(%). :

Range.—In North America known only from Greenland, as reported
by Meinert (57, p. 187) in 1895.

Wagner, 1930, gives Pulex vulpes Motschulsky as a questionable
synonym.

The head of Chaetopsylla globiceps is hardly globular or spherical.
However, the upper and front margins of the head do make a continu-
ous, broadly rounded head outline. Females of globiceps have the
abdomen somewhat distended and the receptaculum seminis with a
hearty sclerotized, subspherical head and a rather long, sausagelike
tall.

467459432

18 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

CHAETOPSYLLA LoToRIs (Stewart)
(Fig. 4, £)

Trichopsylla lotoris Stewart, 1926, Insecutor Inscitiae Menstruus 14: 122, 1 fig. ;
I. Fox, 1940, Fleas of the Eastern United States, p. 33, pl. 8, figs. 30, 37,
and 38, and pl. 9, fig. 42.

Chaetopsylla lotoris Wagner, 1936, Canad. Ent. 68: 195.

Type host—Procyon lotor (L.) (raccoon).

Type locality.—Olcott, N.Y. _

Range.—Eastern part of the United States.

This flea probably has the raccoon as an exclusive true host. That
it should have existed so long on such a common and well-known host
as the raccoon without discovery is remarkable. It is particularly
significant that the host is related to the bears and that Chaetopsylla
setosa Rothschild occurs on our black bear.

CHAETOPSYLLA SETOSA Rothschild
Chaetopsyllus setosus Rothschild, 1906, Canad. Ent. 38: 321, figs. 41 and 42.

Cotype hosts—Canis latrans Say (coyote) and Ewarctos americanus
(Pallas) (black bear). ‘o

Coty pe localities—Eagle River, Sicamous, and Mabel Lake (British
Columbia), Canada.

Range.—Known only from Canada.

The original description of setosa was based only on the female.
A single male, taken from Ursus horribilis bairdi (grizzly bear), and
considered as possibly this species, was described and figured by
Wagner (83, p. 334, fig. 2) in 1936.

The Genus ArcropsyLta Wagner
(Fig. 4, D)

Arctopsylla Wagner, 1930, Katalog der palaearktischen Aphanipteren, p. 40; 1933,
Konowia 12: 94. Type, Ceratophyllus tuberculaticéps Bezzi (by original
designation).

The genus Arctopsylla was proposed by Wagner in 1930, but with-
out description. It was described by him in 1933.

Frontal tubercle vestigial. Antennal club pointed apically. Eyes present,
well developed. Labial palpus long, extending to or beyond apex of trochanter
I, with 7 to 10 segments. Postantennal region of head with 3 rows of setae, the
posterior row submarginal. Pronotal comb absent. Upper sclerite of metepi-
sternum not distinctly separated from metepisternum proper but separated from
metanotum by an internal sclerotized ridge. Metepimeron not fused along
upper part with metepisternum. Abdomen of gravid female but little distended.
Antepygidial bristles not differentiated from the others. Sternum VIII of
male large, broad, unmodified; sternum IX much reduced but otherwise not
greatly modified.

ArcTopsyLLA urst (Rothschild)
(Fig. 4, D)

Pulez ursi Rothschild, 1902, Ent. Rec. and Jour. Variation 14: 62, pl. 2, fig. 1.
Arctopsylla ursi Wagner, 1930, Katalog der palaearktischen Aphanipteren, p. 40
(as a synonym of A. tuberculaticeps (Bezzi) ).
Type host—Ursus horribilis Ord (grizzly bear).
Type locality—Calgary, Alberta.
Range-—Known only from Canada and Alaska.

THE FLEAS OF NORTH AMERICA 19

Wagner (80, p. 40), in his catalog of the Palaearctic fleas, gives
ursi as a synonym of tuberculaticeps Bezzi, but the single male speci-
men of wurst available to the authors for study appears to be distinct.
The absence of antepygidial bristles and the presence of an un-
modified sternum VIII of the male at once mark this species, the only
one of its genus in North America, as very unusual.

The Genus TricHopsyLLowwEs Ewing
(Fig. 4, A)
Trichopsylloides Ewing, 1938, Wash., Ent. Soc. Proc. 40: 94. Monotypic. Type,
Trichopsylloides oregonensis Ewing.
Phaneris Jordan, 1939, Novitates Zool. 41: 317. Monotypic. Type, Phaneris
hubbardi Jordan.

Frontal tubercle minute, angulate apically. Antennal club rounded apically.
Eyes absent. Labial palpus not reaching to apex of trochanter I, 5-segmented.
Postantennal region of head with 3 rows of setae, lowest seta in each row
longest. Pronotal comb composed of long, well-pigmented spines. Upper sclerite
of metepisternum more or less anchylosed with metanotum, but always showing
at least a vestigial inner ridge along its upper boundary. Metepimeron fused
along upper part of anterior end with metepisternum. Abdomen of gravid
female not greatly distended. Antepygidial bristles either undifferentiated from
other nearby setae or but slightly differentiated. Sternum VIII of male large,
unmodified; sternum IX produced into a pair of large, semiclavate, setigerous
lobes.

In the type species there is no evident differentiation of the
antepygidial bristles from the other nearby setae, yet in Phaneris
hubbardi Jordan, a specimen of which was lent to the author of
Trichopsylloides for examination, there is a definite but slight dif-
ferentiation of the antepygidial bristles. The presence in this genus
of a pronotal comb and five-segmented palpi and the absence of eyes
indicate its close relationship with the subfamily Dolichopsyllinae.

TRICHOPSYLLOIDES HUBBARDI (Jordan), new combination

Phaneris hubbardi Jordan, 1939, Novitates Zool. 41: 318, figs. 268 and 269,

Type host—Aplodontia rufa (Raf.) (mountain beaver).

Type locality —Springwater, Oreg.

Range.—Known only from Oregon.

Further study with an abundance of material may show that
hubbardi is only a synonym of oregonensis Ewing. It is similar to
Trichopsylloides oregonensis except for the clear differentiation of
the antepygidial bristles and the fact that the upper sclerite of the
metepisternum is more clearly demarcated from the metanotum in

hubbardi.
TRICHOPSYLLOIDES OREGONENSIS Ewing
(Fig. 4, A)

Trichopsylloides oregonensis Ewing, 1938, Wash. Ent. Soc. Proc. 40: 94.

Type host—Aplodontia pacifica Merr. (mountain beaver).

Type locality.—Delake, Oreg.

Range.—Known only from Oregon and Washington.

This unusual flea is readily identified by the lack of clear differen-

tiation of the antepygidial bristles. The species is of medium size,
the length of the male being 1.8 mm. and that of the female 2.7 mm.

20 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

The Subfamily RHOPALOPSYLLINAE Oudemans

Fronto-epicranial groove usually poorly developed. Frontal notch present,
also frontal tubercle, although the latter may protrude but slightly beyond
the surface of the front. Antennal club long or short, Symmetrical or asym-
metrical. Labial palpus 4- or 5-segmented. Eyes unreduced and pigmented or
reduced and pale. Pronotal comb absent. Antepygidial bristles present. Ab-
domen in gravid females not greatly distended. Parasitic on both birds and
mammals.

This subfamily contains three genera, only one of which (Rhopa-
lopsyllus) is represented in North America.

Rhopalopsyllinae was established by Oudemans (59) in 1909 for
the single genus Rhopalopsyllus Baker, 1905. In it the present writ-
ers place two additional genera, Parapsyllus Enderlein, 1903, and.
Tetrapsyllus Jordan, 1931. Members of this subfamily are confined
to the Southern Hemisphere, except for a few species of Rhopa-
lopsyllus.

In the characters of the head, thorax, and tarsi the Rhopalopsyl-
linae show relationships with the Pulicidae. They differ. from the
latter rather markedly in the characters of the abdomen. Further-
more, in the Rhopalopsyllinae the posterior borders of the labial
palpi usually are well sclerotized, while in the Pulicidae usually they
are either poorly sclerotized or membranous.

Wagner (86, p. 70) places the three genera of this subfamily in
two subfamilies, leaving only the type genus in the Rhopalopsyllinae.
Such a division hardly appears justified on the ground of either
morphological characters or geographical distribution. Furthermore,
the genera Parapsyllus Enderlein and Tetrapsyllus Jordan, included
in the proposed subfamily Parapsyllinae, really differ more between
themselves than the two together differ from Rhopalopsyllus Baker,
and if the subfamily Rhopalopsyllinae, as here used, is to be broken
up, it should be divided into three subfamilies. This number could
hardly be justified.

The genus Rhopalopsyllus Baker is the only North American genus.
of this subfamily.

The Genus RHopALopsyLLus Baker

(Fig. 4, B)

Rhopalopsyllus Baker, 1905, U. S. Natl. Mus. Proc. 29: 129. Type, Pulex lutzii
Baker (by original designation).

Rothschildella Enderlein, 1912, Zool. Anz. 40: 72. Monotypic. Type, Roth-
schildella cryptoctenes Enderlein.

Frontal tubercle angulate apically, varying much in size and sometimes en-
tirely contained in frontal notch. Antennal club short, asymmetrical, with seg-
mentation less marked on front side. Upper margin of antennal groove with a
long comb of closely set microsetae. Eyes well developed and well pigmenied.
Labial palpus 5-segmented, varying much in length. Postantennal region of
head with 3 rows of setae, the posterior row being submarginal. Upper scler-
ite of metepisternum separated from the metanotum by an inner sclerotized
ridge. Abdomen of gravid female not greatly enlarged. Sternum VIII of
abdomen of male enlarged, but otherwise little modified; sternum IX extending
distally as a single, median, unbranched, setigerous process.

This genus was divided into three genera by Jordan (46) in 1939.

The chief characters used for this division are: The shape of the
prosternum, the length of the mesosternum in comparison with its.

THE FLEAS OF NORTH’ AMERICA a4 |

height, and the length of the last segment of each tarsus, but more
particularly the length of the last segment of tarsus II in compari-
son with the length of the second segment of tarsus II.

That the three genera derived from Baker’s Rhopalopsyllus repre-
sent natural groups of species there can be little doubt. However,
the writers have found it hard to place certain species into the newly
derived genera because of the intergradation of characters and their
individual variation; hence, the genera are here given the status of
subgenera. One subgenus, Tiamastus Jordan, does not occur in
North America.

The Subgenus RHOPALOPSYLLUS Baker

The subgenus Rhopalopsyllus Baker as here considered was de-
fined by Jordan in 1939 (46, p. 442) and has Rothschildella Ender-
lein as a synonym.

Prosternum humped or extending downward anteriorly between the coxae.
Mesosterhum as high as long. Last segment of tarsus III longer than second
segment of tarsus II. Pygidium with strong projecting anterodorsal margin.
Head of receptaculum seminis without hump, being broadly rounded both above
and below.

Type species—By original designation (for the genus Rhopa-
lopsyllus), Pulex lutzii Baker.

Three North American species are included in this subgenus.

RHOopPALOPSYLLUS (RHOPALOPSYLLUS) AUSTRALIS AUSTRALIS (Rothschild)

Pulez australis Rothschild, 1904, Novitates Zool. 11: 615.
Rhopalopsyllus australis australis Jordan and Rothschild, 1923, Ectoparasites
1: 327, fig. 338.

Type host.—Tayassu pecari Fisch. (white-lipped peccary).

Type locality—Santa Andrea, Tabasco, Mexico.

Range.—Mexico and Central and South America.

In addition to the type variety, Rhopalopsyllus (R.) australis is
represented in South America by A. (f#.) a. tupinus Jordan and
Rothschild (Peru, Bolivia), and #. (#.) a. tamoyus Jordan and
Rothschild (Paraguay, Brazil).

RHOPALOPSYLLUs (RHOPALOPSYLLUs) CACICUs sAEVUS Jordan and Rothschild

Rhopalopsyllus cacicus saevus Jordan and Rothschild, 1923, Ectoparasites 1:
325, fig. 322.
Type host—Felis pardalis mearnsi Allen (Mearns’ ocelot).
Type locality.—La Cabima, Panama.
fange.—Central and South America.
The typical variety of Rhopalopsyllus (R.) cacicus is from Peru.

RHOPALOPSYLLUS (RHOPALOPSYLLUS) MEsUS Wagner

Rhopalopsyllus mesus Wagner, 1933, Berlin Zool. Mus. Mitt. 18: 349, figs. 9-12.

Type host.—‘Saugetier mit dem einheimischen Namen: Ha-
patschins.”

Type locality.—Twenty-five kilometers east of Cartago, Costa Rica.

Range.—Known only from Costa Rica.

ay MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

The female of mesus is easily identified by the peculiar S-shaped
receptaculum seminis, which has a slender head and a rather stout

tail.
The Subgenus POLYGENIS Jordan

Polygenis Jordan, 1939, Novitates Zool. 41: 444. Type, Pulex roberti Rothschild
(by original designation).

Labial palpus not reaching apex of trochanter I. Prosternum not humped
or extending downward anteriorly between the coxae. Mesosternum not so high
as long. Last segment of tarsus III not longer than second segment of tarsus
II. Head (body) of receptaculum seminis usually produced dorsally or ante-
riorly into a hump.

One of the outstanding characters of most species of this subgenus
is the shape of the head of the receptaculum seminis of the female.
It is produced into a hump above or in front. Usually this hump is
angulate distally.

RHoPALopsYLLUs (PoLyGENIs) DUNNI Jordan and Rothschild

Rhopalopsyllus dunni Jordan and Rothschild, 1922, Ectoparasites 1: 269, figs.
261 and 262.

Type host—Sigmodon hispidus Say and Ord (cotton rat).

Type locality —F¥ort Clayton, Canal Zone, Panama.

Range.—Central America.

According to Jordan and Rothschild (52) this species is closely
related to bohlst Wagner, but the movable finger is shorter, not reach-
ing to the apical bristle of the clasper, and also broader and without
a tooth on the dorsal side near the apex.

RHopPALoPsyLLus (PoLycENIs) Gwyn C. Fox
(Fig. 4, B)

Rhopalopsyllus gwyni C. Fox, 1914, U. S. Pub. Health Serv. Hyg. Lab. Bul.
97: 10, pl. 3, figs. 1-6.

Type host.—‘Rats.”

Type locality —Brunswick, Ga.

fange.—Southeastern part of the United States.

At one time considered as a probable synonym of bohlsi Wagner,
this species was shown by Jordan (42, p. 292) in 1982 to be distinct.
It and Lhopalopsyllus (Polygenis) sigmodoni Stewart are the only
wee of the subgenus Polygenis known to occur in the United

tates.

RHopPALopsyLLus (PoLYGENIs) KLAGESI SAMUELIS Jordan and Rothschild

Rhopalopsyllus klagesi samuelis Jordan and Rothschild, 1923, Ectoparasites
1: 381, figs. 342 and 344.
Type host—‘*Didelphis marsupialis” (opossum).
Type locality—San Esteban, Venezuela.
Fange.—Central and South America.
_ The typical variety of Rhopalopsyllus (Polygenis) klagesi occurs
in Venezuela and Brazil.

THE FLEAS OF NORTH AMERICA 23

RHopALOpPsYLLUs (POLYGENIS) sIGMODONI Stewart

Rhopalopsyllus sigmodoni Stewart, 1930, Canad. Ent. 62: 175, pl. 15, figs. 1
and 2. ;

Type host.—Sigmodon hispidus Say and Ord (cotton rat).

Type locality—Houston, Tex.

Range—Known only from Texas.

This species is very near gwyni C. Fox, but the movable finger
of the clasper appears to be more slender, and there may be a diiler-
ence in the shape of the receptaculum seminis of the two species. A
study of a large series of specimens coming from all the Gulf States
would probably show the Georgia forms merging with those of
Texas. The species fed readily on the arm of the describer.

The Subfamily DOLICHOPSYLLINAE Baker

Head either divided or not divided by development of fronto-epicranial
groove. Frontal notch, or frontal tubercle, or beth notch and tubercle present
(although sometimes vestigial). Eyes either well developed and pigmented,
reduced and poorly pigmented, vestigial, or absent. Labial palpus varying
much in length, frequently surpassing the apex of trochanter I. Pronotal
comb always present, but in a few genera composed of very short, poorly pig-
mented spines. Antepygidial bristles varying in number on a side from 1 to 5
and usually in size or number as between the sexes. Plantar bristles on last
segment of tarsi varying much in number and position, the first pair being
frequently shifted ventrally so that they are out of line with the others.

This subfamily contains more genera than any other subfamily, 20
being represented in North America.

Baker (5, p. 127), in 1905, established the subfamily Dolichopsyl-
linae as one of four subfamilies of Pulicidae. He separated it from
the subfamilies Anomiopsyllinae and Pulicinae on the basis of the
number of antepygidial bristles present. The characters of the Doli-
chopsyllinae are unusually variable. Its closest affinity appears to
be with the Rhopalopsyllinae, from which it is especially differen-
tiated by the presence of a pronotal comb.

One synonym of Dolichopsyllinae is known, Ceratophyllinae, pro-
posed by Dampf (11, p. 19) in 1908 and recognized by Rothschild
(66, p. 58) in 1915 and Wagner (85, p. 4) in 1936. A number of
other families or subfamilies are partly synonymous with the Dolli-
chopsyllinae as here defined.

Key To THE NortH AMERICAN GENERA OF DOLICHOPSYLLINAE

1. With a row or patch of spinelets on inner surface of coxa III_______- 2

. Without a row or patch of spinelets on inner surface of coxa III____ 4
2. Segment V of each tarsus with 5 pairs of lateral plantar bristles;
Sel GEL? LL 25 | ieee GRR Bee dete Mee ML ne Or ee a Odontopsyllus Baker.

Segment V of tarsus III with only 4 pairs of lateral plantar bristles___-— 38

3. Frons with 3 rows of setae and additional small ones
Delotelis Jordan.

Frons with only 2 rows of setae________________ Catallagia Rothschild.
4. None of the setae on front pigmented and thickened toward the
DE tee ae alls BNE TE Sols BSD 17 Ese ie pr ee 5

Some or all of the setae on front pigmented and spinelike or thick-
SL a a ee ees = Se 19

24

MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

5. Female with 4 to 5 antepygidial bristles on each side. Male with 3

< 9

10.

Ae
12.
13.
14.

15.

16.

18.

19.

large antepygidial bristles on each side and a conspicuous median
process on tergum VII projecting over pygidium
Dolichopsylla Baker.
Female with less than 4 antepygidial bristles. Male seldom with 3
large antepygidial bristles and never with process on tergum VII__-
Last segment of each tarsus with 6 pairs of lateral plantar bristles,
the third being shifted ventrally and its place taken by a more
slender -lateral>pait=—.-2 2) 2 a ee eee Dasypsyllus Baker.
Last segment of each tarsus never with more than 5 pairs of lateral
plantar’ ‘bristles_2_ “2 = ee
Tibia III with a comblike row of 3 or 4 shorter, outer setae between
long, middle, and apical pristles; movable finger of male with 1
or more black, Spinelike Setae__.=-=~ =") === Amphipsylla Wagner.
Tibia Ill without such a comblike row. of setaee. 2-3 Se eee
Femur I never with more than a single seta on outer surface________
Kemur 1 with several setae on outer Surfacel=22 2" 2) eee
Segment I of tarsus III longer than segments II to IV
Tarsopsylla Wagner.
Segment I of tarsus III shorter than segments II to IV____---_--____
Movable finger with a Ssubmarginal row of 4 to 7 short, black, sub-
equal, spinelike setae. Ventral margin of sternum X of female
MOU ANS A Ce a fe ag eg Ee Orchopeas Jordan.
Movable finger with 3 to 4 unequal, submarginal, black, spinelike setae.
Ventral margin of sternum X of female usually distinctly angulate
Opisodasys Jordan.
Front with 3 rows of setae. Female with 2 receptacula seminis
Atyphloceras Jordan and Rothschild.
Front seldom with 3 rows of setae. Female with 1 receptaculum

Eyes vestigial or absent; 4 or 5 setae in ocular row
Dactylopsylla Jordan (Subgenus: Foxrella Wagner.)
Eyes present, although sometimes distinctly reduced in size, never
vestigial; usually only iS Setae, in ocilan, rOw=- = eee
Pronotal comb with short spines, not heavier in pigmentation than
pronotum itself. Circumpolar.
Mioctenopsylla Rothschild (Syn.: Boreopsyllus Ewing).
Pronotal comb with usual long, heavily pigmented spines____________
Inner surfaces of coxae II and III with thin setae from base to apex;
labial palpus usually extending heyond apex of trochanter [_-_______
Inner surfaces of coxae II and III with thin setae at most on distal
half; labial palpus not reaching apex of trochanter IJ________________
Sternum VIII of male broad, unreduced, bearing several to many setae.
Basal abdominal sternum of female without patch of lateral setae.
Restricted to North Ameri@as- 32) 25a eee Thrassis Jordan.
Sternum VIII of male much reduced, narrowed and usually extended into
a horizontal process... 22 3 Se ee ee eee 2 ee
Pronotal comb with less than 12 spines on a side_________ Oropsylla Wag-
ner and Toff (Subgen.: Diamanus Jordan) Opisocrostis Jordan).
Pronotal comb with 12 or more spines on a side____Amphalius Jordan.
Sternum VIII of male vestigial, without setae. Old World species
introduced into North America_________ Nosopsyllus Jordan.
Sternum VIII of male modified and bearing 1 or more setae____________
Pronotal comb usually with less than 12 spines on a side; movable finger
of male with 1 or more black, spinelike setae, Parasitic on mammals
Trichopsylla Kolenati (Synonyms: Meg-
abothris Jordan, Malaraeus Jordan) (Subgen.: Pleochaetis Jordan).
Pronotal comb always with more than 12 spines on a side; movable
finger of male usually without black, spinelike setae. Parasitic on
iTS 2-3 28-5 ste we as (Sn eh pa: Evi ene te Ge Ceratophyllus Curtis.
Only anterior submarginal row of frontal bristles pigmented and spine-
like; front with anterior margin broadly rounded
Ctenophyllus Wagner.
All cephalic setae with black pigment and more or less spinelike; front
with anterior margin subangulate. Nearctic Stenistomera Rothschild.

11

10

12

13

14

15
nis

16

18

—s-_

THE FLEAS OF NORTH AMERICA 25

The Genus Opontopsy.Lius Baker

Odontopsyllus Baker, 1905, U.S. Natl. Mus. Proc. 29: 181. Type, Pulex multi-
spinosus Baker (by original designation).

Frontal tubercle small, angulate at apex. Eyes well developed and well
pigmented. Labial palpus not reaching apex of coxa I, d-Segmented, Pronotal
comb with the usual long, black spines, which are acuminate apically. Coxa
III with a patch of spinelets on inner surface; segment I of posterior tarsus
shorter than II, III, and IV taken together; last segment of each tarsus with
5 pairs of lateral plantar bristles; apical bristle on segment II of tarsus III
not reaching apex of segment V. Abdomen with apical spines not extending
down almost to level of spiracles. Antepygidial bristles 8 on each Side in
female, 2 in male. Female with but 1 receptxculum seminis and short or long
stylet. In male, sternal plate VIII large, but slightly modified and bearing
a bunch of subapical setae; sternal plate IX with vertical processes, distal
lobe, and median apophysis; movable finger broad, with some long setae but
no pigmented spines.

This genus was described by Baker to include four species, two of
which, charlottensis Baker and telegoni Rothschild, have been re-
moved to other genera. It now includes only three North American
species, of medium to large size, having rabbits and hares for their
natural hosts.

OpoNToPsyLLUs DENTATUs (Baker)

Ceratophyllus dentatus Baker, 1904, U. S. Natl. Mus. Proe. 27: 390.

Odontopsyllus dentatus Baker, 1905, U. S. Natl. Mus. Proce. 29: 131.

Ceratophyllus ponerus Rothschild, 1909, Novitates Zool. 16: 54, pl. 8, fig. 5.
(New synonymy.)

Type host—Lynzx canadensis Kerr (Canada lynx).

Type locality —Moscow, Idaho.

Range.—Western part of the United States.

Ceratophyllus ponerus Rothschild was described from a single
female. In the United States National Museum are five males and
two females which were identified as C. ponerus by Karl Jordan.
These specimens were taken in the State of Washington. A com-
parison of the male specimens with the male type of Odontopsyllus
dentatus (Baker) shows that the two species are the same. The type
host of dentatus is evidently not a natural one. The natural hosts
are rabbits.

Odontopsyllus dentatus (Baker) has been erroneously synonymized
with Nosopsyllus fasciatus (Bosc) by Dalla Torre (10, p. 13).

ODONTOPSYLLUS MULTIsPINosuUs (Baker)

Pulez multispinosus Baker, 1898, N. Y. Ent. Soc. Jour. 6: 54.

Ceratophyllus multispinosus Baker, 1904, U. S. Natl. Mus. Proc. 27: 389, pl. 12,
figs. 1-5.

Odontopsyllus multispinosus Baker, 1905, U. S. Natl. Mus. Proc. 29: 129; I.
a Fleas of the-Eastern United States, p. 47, pl. 7, figs. 29, 30,
and 31.

Type host.—Sylvilagus floridanus (Allen) (cottontail rabbit).

Type locality—Raleigh, N. C.

Range.—Eastern part of the United States.

This large, dark-brown flea can be easily distinguished by the
characters of the movable finger of the male, which is very broad,

triangular, and possesses a row of about five long setae on its lower
margin. :

26 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

ODONTOPSYLLUS SPENCERI Dunn

Odontopsyllus spenceri Dunn, 1923, U. S. Pub. Health Serv. Rpts. 38 :. 2765-2766.

Type host.—Lepus bairdii Hayden (snowshoe rabbit).

Type locality.—Bitterroot Mountains west of Hamilton, Mont.

Range.—Known only from Montana.

A rare flea and apparently not yet figured. According to the orig-
inal description the male “is close to the male described as dentatus
by Baker, differing from it mainly by having more bristles in the
upper genal row and on the dorsal surface of the ninth sternite.”

The Genus Denorte.is Jordan

Delotelis Jordan, 1937, Novitates Zool. 40: 267. Monotypic. Type, Cerato-
phyllus telegoni Rothschild.

Material not being at hand for the description of this genus, the
original description by Jordan is quoted in full:

6 2. Like Catallagia Roths. 1915; but frons with three rows of bristles
(2, 3, 7, or 1, 4, 7) and additional small bristles, some of which are placed
below vestigial eye; stigma-cavity of tergum VIII narrow; hindtarsal segment
V with one ventral bristle in between first lateral pair. In ¢ abdominal
sternum VIII with dense apicimarginal row of bristles; process P of clasper
much shorter than movable digitoid F. In @ duct of bursa copulatrix short;
orifice of body of spermatheca terminal, apex of tail of spermatheca obliquely
truncate.

This genus contains but a single species.

DELoTELIs TELEGONI (Rothschild)

ee telegoni Rothschild, 1905, Novitates Zool.-12: 172, pl. 9, figs. 27
an S
Delotelis telegoni Jordan, 1937, Novitates Zool. 40: 267.

Cotype hosts—Microtus drummondii (Aud. and Bach.) (meadow
mouse) and Hvotomys gapperi (Vigors) (red-backed mouse).

Cotype localities—Horse Creek, Upper Columbia Valley, British
Columbia, and Kicking Horse Canyon, Alberta.

Range.—Known from Alberta and British Columbia.

In the original description of this species it was compared several
times with charlottensis, now Catallagia charlottensis (Baker), with
which it was said to be closely related. The two species evidently
ae eee different, since they are now placed in different genera by

ordan.

The Genus Caratiacta Rothschild
(Fig. 5, C)

Catallagia Rothschild, 1915, Hctoparasites 1: 41, Type, Pulexr charlottensis
Baker (by original designation).

Frontal tubercle small to vestigial, frequently completely concealed in frontal
notch. Forehead with 2 rows of lateral setae, none of which is spiniform.
Eyes vestigial. Labial palpus not reaching apex of coxa I, 5-segmented. Post-
antennal region of head with 3 rows of setae. Pronotal comb with usual long,
dark-brown spines, which are either narrowly rounded or angulate at apex.
Coxa II with a row or patch of spinelets on inner surface; last segment of
posterior tarsus with 4 pairs of lateral plantar bristles. Abdomen with apical
spines which do not extend down to near level of spiracles. Tergal plate VII
without posterior median process. Antepygidial bristles 3 on each side of

THE FLEAS OF NORTH AMERICA yer

abdomen in both sexes. Female with but 1 receptaculum seminis. In male,
sternal plate VIII produced posteriorly into a semihyaline flap; sternal plate
IX produced posteriorly into a spiniferous and setiferous process; movable
finger of clasper broad at base and angulate apically, bearing several to many
small setae.

This genus is confined to northern North America, the most south-
ern locality being San Francisco. A key to the males and females,
before Delotelis was split off, was published by Ewing (/7, p. 176) in
1929.

CaTALLAGIA BOREALIS Ewing

Catallagia borealis Ewing, 1929, Wash. Biol. Soe. Proc. 42: 127.

Type host —Microtus pennsylvanicus (Ord) (meadow mouse).
Type locality—Basin Pond, Mount Katahdin, Maine.
Range-—Known only from Maine.

At the time of its description this species was known only from a
single female. Catallagia onaga Jordan may possibly be a synonym
of borealis. The head of the female and sternal plate VII of the
same sex have been figured by I. Fox (25, pl. 28, figs. 146 and 147).

CATALLAGIA CHAMBERLINI Hubbard

Catallagia chamberlini Hubbard, 1940, Pacific Univ. Bul. 37 (3): [4], 3 figs.

Type host—(?%). “Normal host is the Deer mouse.”

Type locality—Rocky Point (north of Klamath Falls), Oreg.

Range.—Given by Hubbard (37) as “Cascade and Siskiyou Moun-
tains of Oregon, with encroachments into the Willamette Valley.”

The original description of this species was based on 25 specimens
and is accompanied by a drawing of the ninth sternal plate of the
male, the characters of which are distinctive.

CATALLAGIA CHARLOTTENSIS (Baker)
(Fig. 5, C)

Tuphlopsylia charlottensis Baker, 1898, N. Y. Ent. Soc. Jour. 6: 56.
Caiailagia charlottensis Rothschild, 1915, Ectoparasites 1: 45, figs. 44 and 46.
Type host.—*Mouse nest.”
Type locality —Massett, Queen Charlotte Islands.
Range.—Known from British Columbia, Alberta, and Oregon.
This species, the type of its genus, was redescribed and the geni-
talia of the male figured by Rothschild (1915). It is very closely
related to decipiens Rothschild. (See remarks under the latter
species. )
CATALLAGIA DEcIPIENS Rothschild

Catallagia decipiens Rothschild, 1915, Ectoparasites 1: 43, figs. 45 and 47.

Type host—Peromyscus sp. (white-footed mouse).

Type locality—Horse eae Upper Columbia Valley, British
Columbia.

Range.—Known from British Columbia, Alberta, and the Great
Basin of the western part of the United States.

This flea is very similar to charlottensis, from which it can best be
separated on characters of the male genitalia. In decipiens the proc-

98. MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

ess of the clasper extends to about the tip of the movable finger,
while in charlottensis it extends considerably beyond that.

CATALLAGIA MONERIS Jordan

Catallagia moneris Jordan, 1937, Novitates Zool. 40: 267, fig. 50.

Type host—Marmota flaviventris (Aud. and Bach.) (woodchuck).

Type locality —Ravalli County, Mont.

Range.—Known only from Montana.

In his original description Jordan states that this species “Agrees
in all details with C. decipiens Roths. 1915, from Alberta and British
Columbia, except in the size and shape of the spermatheca.” The
shape of this spermatheca, or receptaculum seminis, is different from
that of any other female of Catallagia, as far as known, in having the
head not widened toward the tail and longer than the latter. Only
a single female is known.

CATALLAGIA MOTEI Hubbard

Catallagia motei Hubbard, 1940, Pacific Univ. Bul. 37 (3): [4], 2 figs.

Type host—Not mentioned by describer.

Type locality —Banks, Oreg.

Range.—Known from Oregon and Washington.

The original description of mote? was based entirely on females
and is accompanied by a drawing of the receptaculum seminis and
seventh sternal plate.

CATALLAGIA ONAGA Jordan

Catallagia onaga Jordan, 1929, Novitates Zool. 35: 172, fig. 7; I. Fox, 1940, Fleas
of Eastern United States, p. 103, pl. 28, fig. 145.

Type host—Blarina brevicauda (Say) (short-tailed shrew).

Type locality—Adirondack Mountains, N. Y.

Range.—Known only from New York State.

This species is probably a synonym of Catallagia borealis Ewing,
being based on the male of that species, but until the sexes of either
borealis or onaga have been properly associated, it appears best to
give onaga standing as a distinct species.

CATALLAGIA SCULLENI Hubbard

Catallagia sculleni Hubbard, 1940, Pacific Univ. Bul. 37 (3): [8], 3 figs.

Type host—(*%). “Normal host is the Deer mouse.”

Type locality —F¥orest Grove, Oreg.

Range.—According to a statement accompanying the original de-
scription: “The range extends from the Pacific Ocean east over the
Coast Range into the Puget Sound Trough in Oregon and Washing-
ton.

This species was described from 35 specimens. The original de-
scription is accompanied by a drawing of the ninth sternal plate of
the male, which is characterized by having 4 black, short, spine-
like, subapical setae and a ventral, marginal fringe of long setae.

THE FLEAS OF NORTH AMERICA 29

CATALLAGIA WYMANI (C. Fox)

Odontopsyllus wymani C. Fox, 1909, Ent. News 20: 241, figs. 1-5.
Catallagia wymani Rothschild, 1915, Ectoparasites 1: 42.

Type host.—Microtus californicus (Peale) (California field
mouse).

Type locality—San Francisco, Calif.

Range.—Known only from California.

This species can be distinguished from all others of its genus by
the characters of sternal plate IX of the male, each posterior proc-
ess of this structure having but a single subapical, ventral spine and
being fringed throughout most of its ventral margin wth a row of
subequal setae.

The Genus DoticHopsyLLa Baker
(Fig. 5, H, F)

Dolichopsylla Baker, 1905, U. S. Natl. Mus. Proc. 29: 185. Monotypie. Type,
Ceratophyllus stylosus Baker.

Frontal tubercle small, angulate apically. Forehead with 2 rows of lateral
setae, none of which is spiniform. Eyes absent. Labial palpus long, extending
to or beyond apex of trochanter I, 5-segmented. Postantennal region of head
with 2 rows of setae. Pronotal comb with usual, long, black spines, which are
either narrowly rounded or angulate apically. Coxa III without a row or patch
of setae on inner surface; segment I of posterior tarsus not longer than II,
III, and IV taken together; last segment of tarsus III with 5 pairs of lateral
plantar bristles, the first being shifted slightly ventrally. Abdomen with apical
spines not extending down almost to line of spirac'es. Tergal plate VII with
a posterior median process. Female with 4 antepygidial bristles on each side,
male with 3. Female with a single receptaculum seminis. Male with sternal
plate VIII produced posteriorly into a pair of Ssemihyaline lobes, each having
a fimbriated ventral margin; sternal plate [TX much modified distally, with
crescentiec inner vertical lobes, and a dorsoventrally thickened median apo-
physis ; movable finger of clasper broad, flat, subtriangular, with long setae but
no pigmented spines.

This genus contains but a single species. It is the type genus of

the family Dolichopsyllidae, the largest of all the families of
Siphonaptera.

DoticHopsyLia styLosa (Baker)
(Fig. 5, B, F)

Ceratophyllus stylosus Baker, 1904, U. S. Natl. Mus. Proc. 27: 418, pl. 14, figs.
1-7, and pl. 15, figs. 1 and 2.
Dolichopsylla stylosa Baker, 1905, U. S. Natl. Mus. Proc. 29: 1385.

Type host.—Aplodontia rufa (Raf.) (mountain beaver).

Type locality.—Astoria, Oreg.

Range.—Known only from Oregon.

This species is a large flea, females frequently measuring 6 mm.
in length. It is at once identified by its large size plus the generic
characters. In the United States National Museum there is a fine
series of six specimens of stylosa representing type material, each
specimen mounted on a separate slide. Excellent figures accompany
the original description of the species.

30 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

The Genus DasypsyLt_us Baker
(Fig. 5, D)

Dasypsylius Baker, 1905, U. S. Natl. Mus. Proc. 29: 129. Monotypic. Type,
Ceratophyllus perpinnatus Baker.

Frontal tubercle minute, angulate apically. Forehead with 3 rows of lat-
eral setae, none of which is spiniform. Eyes well developed and heavily pig-
mented. Labial palpus not reaching to apex of coxa I, 5-segmented. Post-
antennal region of head with 3 rows of setae. Pronotal comb with long, dark-
brown, approximate, apically angulate spines. Coxa III. without a row or
patch of spinelets on inner surface. Last segment of each tarsus with 6 pairs
of lateral plantar bristles, the third pair being shifted ventrally and its place
taken by a more slender lateral pair. Abdomen with a few apical spines, all
situated dorsally. Tergal plate VII without a posterior median process. A
single antepygidial bristle, flanked by a pair of microsetae, on each side in
both sexes. Female with 1 receptaculum seminis and a pair of short, stout
stylets. In male, sternal plate VIII greatly shortened and bearing posteriorly
a pair of semihyaline lobes, each with a fimbriated ventral margin; sternal
plate IX with a pair of internal vertical processes, a conspicuous median apo-
physis, and a pair of long, setigerous, posterior processes; movable finger of
clasper broad, flat, bearing both long setae and pigmented spines.

Dalla Torre (10, p. 12) in his catalog placed this genus as a
synonym of Ceratophyllus Curtis.

DasyPsYLLUS GALLINULAE PERPINNATUs (Baker)
(Fig. 5, D)
Ceratophyllus perpinnatus Baker, 1904, U. S. Natl. Mus. Proc. 27: 391, pl. 13,

figs. 1-6.
Dasypsyllus perpinnatus (Baker), 1905, U. S. Natl. Mus. Proc. 29: 129.
Dasypsyllus gallinulae perpinnatus Wagner, 1930, Katalog der palaearktischen
Aphanipteren, p. 14.

Type host—Unknown.

Type locality——Queen Charlotte Islands.

Range.—Known only from British Columbia.

This subspecies was regarded by Baker as a species in his original
description, which was based on a single male and accompanied by
six detail drawings. It is a medium-sized flea (length of male, 3
mm.) of light-brown color. The type variety of gallinulae occurs
in Europe.

The Genus AmpuipsyLta Wagner
(Fig. 6, C)

Amphipsylla Wagner, 1909, Mitt. Kaukasischen Mus. 4: 196, 201. Type, Amphi-
psylla schelkovnikovi Wagner (by subsequent designation).

Front tubercle vestigial. Forehead with 3 rows of lateral setae, middle row
frequently incomplete, and none of the setae in any row sSpinelike. Eyes small
to vestigial. Labial palpus medium in length, usually not reaching apex of
coxa I. Postantennal region of head with 3 rows of setae. Pronotal comb with
usual long, black, or dark-brown spines. Coxa III without a row or patch
of spinelets on inner surface; tibia III with a comblike row of 3 or 4 shorter
outer setae between long middle and long apical bristles; last segment of
tarsus III with 5 pairs of lateral plantar bristles, the first pair situated in
between the bristles of the second pair. Abdomen with apical spines, nearly all
of which are situated dorsally. Tergal plate VII without posterior median
process. Three antepygidial bristles on each side in both sexes. Female with

but 1 receptaculum seminis. In male sternal plate VIII broad, not modified; —

sternal plate IX with internal vertical processes and much modified posterior
processes ; movable finger of clasper broad, flat, with 1 or more pigmented spines
in addition to a few setae.

THE FLEAS OF NORTH AMERICA 31

This genus is represented in America by only three species, but it
is well represented in the Old World, where it ranges over much of
Europe and central and northern Asia. The species are parasitic
chiefly-on rodents of the family Muridae and carnivores that feed on
these rodents. :

AMPHIPSYLLA EWING! I. Fox
(Fig. 6, ©)
Amphipsylla ewingi I. Fox, 1940, Wash. Ent. Soc. Proc. 42: 61, pl. 10, fig. 1.

Type host—“Alaska short-tailed mouse.”

Type locality —Golovin, Alaska.

Range.—Known only from Alaska.

This is a medium-sized, dark-brown species which is known only
from male specimens. The movable finger of the clasper has two
black, posterior, submarginal spines, the upper one being much the
shorter and somewhat stouter than the other and situated near the
posterodorsal angle. Sternal plate VIIT of the male has a distinctive
comblike row of long setae on each side near the posterior margin.

AMPHIPSYLLA NEOTOMAE I. Fox

Amphipsylla neotomae I. Fox, 1940, Wash. Acad. Sci. Jour. 30: 273, fig. 3.

Type host—Neotoma desertorum Merr. (wood rat).

Type locality—Dos Palos, Calif.

Range—kKnown only from the type locality.

The original description of neotomae was based on a single male
and is accompanied by a figure of the clasper. ‘This is the only species
of its genus to be reported from the United States. It differs from
the other two species of Amphzpsylla known from North America in
the structure of the male terminalia.

AMPHIPSYLLA SIBIRICA POLLIONIS (Rothschild)

Ceratophyllus pollionis Rothschild, 1905, Novitates Zool. 12: 171, pl. 9, figs. 28,
31, and 82.
Amphipsylla sibirica pollionis Wagner, Canad. Ent. 68: 195.

Cotype hosts—Microtus drummondii (Aud. and Bach.) (meadow
mouse) and Hvotomys saturatus Rhoads (red-backed mouse).

Type locality—Red Deer, Alberta.

Range.—Known only from Alberta.

This subspecies is similar -to ewingi I. Fox, but in it the movable
finger is much broader at the top and the upper spine on the same is
longer and not so stout as in ewing?. Sternal plate VIII has a patch of
long setae on each side instead of the submarginal comblike row of
EWING.

The Genus TarsopsyLta Wagner

(Mig. 6;, 5)

Ctenonotus Kolenati, 1863 (not Fitzinger, 1843), Soc. Ent. Rossicae Horae 2: 34.
Monotypic. Type, Ctenonotus octodecimdentatus Kolenati.
Tarsopsylla Wagner, 1927, Konowia 6: 108.

The name Zarsopsylla was proposed by Wagner to replace Ctenono-
tus Kolenati, 1863, which was found to be preoccupied.

oe MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

Frontal tubercle angulate apically and concealed in frontal notch. Forehead
with 3 short rows of lateral setae, none of which is spiniform. Eyes present, pig-
mented. Labial palpus not reaching to apex of coxa I, 5-Segmented. Postantennal
region of head with 3 rows of setae, first 2 incomplete. Pronotal comb with
usual, long, black spines. Femur I with not more than a Single lateral seta;
coxa FII without a row or patch of setae on inner surface; segment I of tarsus
III longer than II, III, and IV taken together; segment V of tarsus III with 5
pairs of lateral plantar bristles, pair I shifted ventrally. Abdomen without
apical spines. Tergal plate VII without posterior median process. Male with 2
antepygidial bristles on a side. Female with a single receptaculum seminis. In
male sternal plate VIII modified, being much narrowed and curved upward Cis-
tally; sternal plate IX with inner vertical processes and anterior median
apophysis; movable finger of clasper long, flattened, without pigmented spines,
but with several long setae.

Besides the type species, which occurs on squirrels in eastern and
central Europe and northern Asia, Zarsopsylla includes a single North
American species, also found on squirrels.

TARSOPSYLLA COLORADENS!IS (Baker)
(Fig. 6, B)

Pulex coloradensis Baker, 1895, Canad. Ent. 27: 110.
Ceratophyllus coloradensis Baker, 1804, U. S. Natl. Mus. Proce. 27: 417, 441, pl.
25, figs. 6-9.

Lype host—“Fremont’s chickaree.”

Type tocality.—Georgetown, Colo.

Range.—Known from Colorado and British Columbia.

A medium-sized, dark-brown flea, easily identified by the fact that
segment I of tarsus III is longer than II, III, and IV taken together.
Apparently this is a rare species, of which the United States National
Museum contains only a single specimen, the type.

The Genus Orcuopgas Jordan
(Fig. 7, C)

Orchopeas Jordan, 19388, Novitates Zool. 39: 71. Type, (Pulex wickhami
Baker) =Pulex howardii Baker (by original designation).

Frontal tubercle small, angulate apically, and largely concealed in tubercular
notch. Forehead with 1 to 2 complete rows of lateral setae, the third (front)
row occasionally represented by 1 or 2 setae in some species. Eyes conspicuous,
black. Labial palpus not reaching to apex of trochanter I, 5-segmented. Post-
antennal region of head with but 1 row of setae, the marginal row. Pronotal
comb with usual, long, black spines. Femur I with not more than 1 lateral seta;_
coxa III without a row or patch of spinelets on inner surface; segment I of
posterior tarsus shorter than II, III, and IV combined; last segment of each
tarsus with 5 lateral plantar setae, first pair displaced ventrally. Abdomen
with but a few apical spines, al situated dorsally. Tergal plate VII without a
posterior median precess; 3 antepygidial bristles on each side in female, 2 in
male. Female with but a single receptaculum seminis, and with ventral margin
of sternal plate X angulate near middle. In male sternal plate VIII much
modified, with apical membranous appendage; sternal plate IX with conspicuous
inner vertical processes, 2 pairs of highly modified distal lobes, and anterior
median apophysis; movable finger of clasper broad, expanded dorsally, with

posterior, submarginal row of 4 to 7 upwardly directed, short, black, subequal]
spines.

This genus is exclusively Nearctic in distribution.

THE FLEAS OF NORTH AMERICA 33

OrcCHOPEAS CAEDENS (Jordan)

Ceratophyllus caedens Jordan, 1925, Novitates Zool. 32: 104, figs. 16 and 17.
Ceratophyllus caedens durus Jordan, 1929, Novitates Zool. 35: pl. 1, fig. 3.
Orchopeas caedens Jordan, 1933, Novitates Zool. 39: 71.

Type host—Martes americana (Turt.) (marten).

Type locality—Banff, Alberta.

Range—United States, Canada, and Alaska.

The writers are following Wagner (84, p. 199) in the synonymy
given above. Under the name of Ceratophyllus caedens durus Jor-
dan, this species was reported from south Labrador by Eidmann (72,
p. 100), the specimens having been determined by Jordan.

OrcHOPEAS DIETERI (C. Fox)

Ceratophyllus nepos dieteri C. Fox, 1929, Ent. News. 40: 218.
Orchopeas dieteri I. Fox, 1940, Wash. Ent. Soc. Proc. 42: 66, pl. 10, figs. 5 and 6.
Type host—Lynzx rufus (Schreber) (bobcat).
Type locality—lLos Angeles County, Calif.
Range.—Known only from California.
I. Fox (1940) redescribed dieteri as a species and gave drawings
of the parts of the terminalia of both sexes.

OrcHovEas HowArpt (Baker), new combination

Pulex howardii Baker, 1895, Canad. Ent. 27: 110.

Pulez wickhami Baker, 1895, Canad. Ent. 27: 109; I. Fox, 1940, Fleas of Eastern
United States, p. 59, pl. 15, figs. 75-77.

Pulex gillettei Baker, 1895, Canad. Ent. 27: 109.

Cotype hosts—*“Squirrel,” “gray or fox squirrel,” “nest of field
mouse.”

Cotype localities—tIthaca, N. Y.; Tallulah Falls, Ga.; Lincoln,
Nebr.

Range.—United States east of the Great Plains.

The name wickhami has long been used for this species and has
page precedence over howardii.. However, Baker (3, p. 37) in 1899
selected howardii for the name of the species. He states:

Puler howardii Baker. In the group which includes this flea and fasciatus
and its allies, the males determine the snecies. For this reason wickhami and
gillettet are not well founded. They were separated on characters whose values
could not have been correctly estimaied at the time. They should be reduced to
synonyms of howardii.

The life history of this fiea (under the name of wickhamz) has been
studied by Sikes (70), who gives a description of the eggs and first
three larval instars.

A chalcidoid parasite, reared from a larva of howardii, was de-
scribed by Waterston (89) under the name of Batramlia fuscipes.
Parasites of this type are very rarely reported from ectoparasites.

OrcHopEas LABIATUs (Baker)

Ceratophyllus labiatus Baker, 1904, U. S. Natl. Mus. Proc. 27: 402, pl. 19,
figs. 6-9.
Orchopeas labiatus Jordan, 1933, Novitates Zool. 39: 72.
Type host—Lynx canadensis Kerr (Canada lynx).
Type locality—Moscow, Idaho.

467459—43——3

94 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

Range.—Known only from Idaho.

This species is based upon a single female which has not yet been
associated with its opposite sex. According to Jordan (40, p. 29)
it is very closely allied to nepos (Rothschild) and caedens (Jordan).
I. Fox (26, pl. 19, fig. 7) gives a detail drawing of sternal plate VII
of the female.

OrcHOPEAS LATENS (Jordan)

Ceratophyllus latens Jordan, 1925, Novitates Zool. 32: 105, figs. 19 and 20.
Orchopeas latens Jordan, 1933, Novitates Zool. 39: 72.

Type host—“Gray squirrel.”

Type locality—Santa Cruz County, Calif.

Range-—Known only from California.

Jordan’s description of this species was based on one male and
two females from the Baker collection. It is accompanied by a
drawing of the clasper of the male and one of sternal plate VII of
the female.

Orcuopgas LEucopus (Baker)

Ceratophyllus leucopus Baker, 1904, U. S. Natl. Mus. Proc. 27: 401.

Ceratophyllus aeger Rothschild, 1905, Novitates Zool. 12: 166, pl. 6, figs. 5, 7,
and 9.

nae leucopus Jordan, 1933, Novitates Zool. 39: 72.

Type host—Peromyscus leucopus (Raf.) (white-footed mouse).

Type locality—Peterboro, N. Y.

Range.—KEastern part of the United States and Canada.

The original description of this species, one of the most common
and one of the best known fleas in America, was based upon a single
female. Ovrchopeas leucopus is very near O. howardii (Baker) but
the clasper has a broad, truncate process and a more slender movable
finger. I. Fox (25, pp. 64 and 159) has redescribed and figured the
species.

OrcHopgas NEPOs (Rothschild)

Ceratophyllus nepos Rothschild, 1905, Novitates Zool. 12: 168, pl. 7, figs. 18
and 14.
Orchopeas nepos Jordan, 1933, Novitates Zool. 39: 72.

Type host.—S pilogale phenax latifrons Merr. (spotted skunk).

Type locality—Chilliwack, British Columbia.

Range.—Known only from British Columbia.

The original description of this species was based on one male and
one female. It is accompanied by drawings of part of the ter-
minalia of the female and a side view of the male claspers. The
species is closely related to howardii (Baker) but differs from that
species in that the process of-the clasper is rounded and not expanded
apically instead of being truncate and expanded; also the distal half
of the movable finger is not nearly so swollen as in howardii.

ORCHOPEAS SEXDENTATUS SEXDENTATUS (Baker)
(Fig. 7, C)
Ceratophyllus sexdentatus Baker, 1904, U. S. Natl. Mus: Proc. 27: 403, pl. 26.

figs. 8-14.
Orchopeas sexdentatus Jordan, 1933, Novitates Zool. 39: 72.

THE FLEAS OF NORTH AMERICA | 35

Type host—Neotoma sp. (wood rat).

Type locality—Boulder Creek, Calif. :

Range.—The species is widely distributed in the United States and
Canada, forming several subspecies. This particular subspecies is
known only from California. ;

The original description of serdentatus was based upon a single
lot of material from the type locality. It is accompanied by several
detail figures, but does not properly treat of the characters now con-
sidered of subspecific value. The various subspecies of seadentatus
parasitize different species of Veotoma.

ORcHOPEAS SEXDENTATUs AGILIs (Rothschild)

Ceratophyllus agilis Rothschild, 1905, Novitates Zool. 12: 167, pl. 7, figs. 16-18.
Ceratophyllus serdentaius agilis Jordan, 1929, Novitates Zool. 35: 30.

Type host—Neotoma cinerea (Ord) (wood rat).

Type locality—Banff, Alberta.

Range-—Known from British Columbia and Alberta.

Rothschild’s description of agilis was based on a large series of
both sexes taken from several localities and is accompanied by three
detail drawings. He states:

Both sexes differ from serdentatus in the longest apical bristle on the hinder

side of the first hindtarsal segment reaching nearly to the apex of the second
segment.

ORCHOPEAS SEXDENTATUS CASCADENSIS Jordan

Orchopeas sexdentatus cascadensis Jordan, 1939, Novitates Zool. 41: 317,
fig. 267.

Type host—Neotoma fuscipes fuscipes Baird (wood rat).

Type locality.—Odell Lake, Oreg.

Range-—Western part of the United States.

The description was based on females and accompanied by a figure
illustrating the variation in sterum VII, particularly its process above
the sinus. This process is acutely angulate apically in cascadensis
whereas it is long and obtuse in agilis.

ORCHOPEAS SEXDENTATUS NEVADENSIS (Jordan)

sneha sexrdentatus nevadensis Jordan, 1929, Novitates Zool. 35: 30, pl. 1,
S:

Type host —Mustela arizonensis (Mearns) (weasel).

Type locality.—Pine City, Calif.

Range.—Known from California and Utah. -

The female of Orchopeas sexdentatus nevadensis is not always dif-
ferentiated from that of O. s. sexdentatus, but the male is distinct,
having the anterior lobe of sternal plate [TX longer and the spinelike
seta situated beyond its middle. The type host of nevadensis is not
a natural one. The subspecies probably is a natural parasite of wood
rats.

ORCHOPEAS SEXDENTATUS PENNSYLVANICUs (Jordan)

Ceratophyllus sexdentatus pennsylvanicus Jordan, 1928, Novitates Zool. 34: 184,
figs. 8 and 9.

Type host—Neotoma pennslyvanica Stone (wood rat).
Type locality—Rolling Rock Club, Ligonier, Pa.

36 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE.

Range.—Kastern part of the United States.

The original description of this subspecies is accompanied by a
drawing of the terminalia of each sex. In the female the upper lobe
of sternal plate VII, which is long and slender, is dilated apically.
The male genitalia differ from those of the other subspecies in minor
details. A redescription of the subspecies is given by I. Fox (28,
DOD):

OrRcCHOPEAS SEXDENTATUS SCHISINTUs (Jordan)

Ceratophyllus serdentatus schisintus Jordan, 1929, Novitates Zool. 35: 31,
pled, figs +5:

Type host—Neotoma sp. (wood rat).

Type locality—Arizona.

Range.—Known only from Arizona.

The original short description of this subspecies is here quoted in
full:

Apparently differs in the Q only: the upper lobe of sternite VII broad and
short, the lower one absent or quite short. Exopodite of ¢ with 5 or 6 spini-
forms.

The Genus Opisopasys Jordan
(Fig. 7, B)

Opisodasys Jordan, 1933, Novitates Zool. 39: 72. Type, Ceratophyllus vesperalis
Jordan (by original designation).

Frontal tubercle small to vestigial, angulate apically, and largely concealed
in frontal notch. Forehead with a single row of setae, the ocular row, but
each of other 2 rows frequently represented by a single large seta. Eyes well
developed, black. Lahial palpus 5-segmented, not reaching beyond apex of
trochanter I. Postantennal region of head with a single row of setae, the
posterior or marginal row; second row usually represented by a Single large
seta. Pronotal comb with usual long, black spines. Femur I with not more
than a single lateral seta; coxa III without a patch or row of spinelets on
inner surface; segment I of posterior tarsus shorter than II, III, and IV com-
bined; last segment of each tarsus with 5 pairs of lateral, plantar setae, pair
I being displaced ventrally. Abdomen with a few apical spines, all dorsal.
Tergal plate VII without posterior median process. Antepygidial bristles 3 or
4 on each side in female; 2 on each side in male. Female with a single recep-
taculum seminis and ventral margin of sternal plate X not distinctly angulate.
In male sternal plate VIII much modified, with or without apical membranous
appendage; sternal plate IX with conspicuous, inner, vertical processes, an
anterior median apophysis, and a pair of expanded, setigerous, posterior lobes:
movable finger of clasper large and flat, bearing 3 or 4 unequal pigmented spines,
which usually are directed backward or downward.

This genus is very clearly near to Orchopeas Jordan, from which,
however, it is easily distinguished by the nature of the pigmented
spines on the movable finger of the clasper in the male. All species
are North American and probably all are Nearctic.

Opisopasys ENopLus (Rothschild)

Ceratophyllus enoplus Rothschild, 1909, Novitates Zool. 16: 53, pl. 8, fig. 6.
Opisodasys enoplus Jordan, 1933, Novitates Zool. 39: 72.

Type host—‘California grey squirrel.”

Type locality—Kings River, Calif.

Range.—Known only from California.

Rothschild based his description of enoplus upon a single male.
The species is characterized by having two pigmented, spinelike setae

THE FLEAS OF NORTH AMERICA 37

on the movable finger, the upper being acutely pointed apically and
the lower rounded apically.

Opisopasys KEENI (Baker)
(Fig. 7, B)
Pulexr keeni Baker, 1896, Canad. Ent. 28: 234.
Ceratophyllus keeni Baker, 1904, U. S. Natl. Mus. Proc. 27: 400, pl. 15, figs. 7-12.
Opisodasys keeni Jordan, 1933, Novitates Zool. 39: 72.
Type host—Peromyscus maniculatus keeni (Rhoads) (white-
footed mouse). a
Type locality—Queen Charlotte Islands, British Columbia.
Range.—Western part of the United States and Canada. ae
In this species the frontal tubercule and notch are either vestigial
or absent. When vestigial they are easily overlooked.

Opisopasys PsEUDARCToMys (Baker)

Ceratophyllus pseudarctomys Baker, 1904, U. S. Natl. Mus. Proce. 27: 399,
pl. 24, figs. 1-7.

Ceratophyllus acasti Rothschild, 1£05. Novitates .Zool. 12: 168, pl. 7, figs. 19
and 20.

Opisodasys pseudarctomys. Jordan, 1933, Novitates Zool. 39: 72.

Type host—Marmota monax (.) (woodchuck).

Type locality—Newport, Herkimer County, N. Y.

Range.—United States and Canada.

Baker’s description of this common and well-known flea is accom-
panied by seven drawings, one being of the head of the male and
one the head of the female. The movable finger of the clasper in
pseudarctomys is striking in appearance and identifies the species. It
is roughly sickle-shaped, with a long, stout, backwardly curved,
blunt-pointed, apical, spmelike seta, and two short setae on the lower
extremity, the upper being sharply pointed and the lower stout and
rounded at the tip.

I. Fox (25, p. 57) has recently redescribed pseudarctomys.

Opisopasys ropustus (Jordan)

Ceratophyllus robustus Jordan, 1925, Novitates Zool. 32: 105, fig. 18.
Opisodasys robustus Jordan, 1933, Novitates Zool. 39: 72.
Opisodasys spatiosus I. Fox, 1940, Wash. Ent. Soc. Proc. 42: 65, pl. 10, fig. 2.

Type host—Unknown.

Type locality—White River, Cooley, Ariz.

Range.—Southwestern part of the United States.

The original description of this species was based entirely on the
female. The male was described in 1939 by Jordan (45, p. 316),
who examined specimens taken in the Catalina Mountains, Ariz., on
Sciurus arizonensis catalina.

OPISODASYS VESPERALIS (Jordan)

Ceratophyllus vesperalis Jordan, 1929, Novitates Zool. 35: 28, pl. 1, figs. 1 and 2.
Opisodasys vesperalis Jordan, 1933, Novitates Zool. 39: 72.
Type host—Glaucomys sabrinus alpinus (Rich.) (flying squirrel).
Cotype localities —“Okanagan and Okanagan Landing,” British
Columbia.

38 MISC. PUBLICATION 500, U. S.:DEPT. OF AGRICULTURE

~ Range.—Known only from British Columbia.

The original description of vesperalis is accompanied by a draw-
ing of the clasper of the male and the seventh sternal plate of the
female. According to Jordan this species is close to pseudarctomys
Baker. However, the male genitalia are very different, the clasper
being without the posterior process, the movable finger being almost
straight instead of crescentic, and the upper spinelike seta on the
same being swordlike instead of rodlike.

The Genus ATYPHLOCERAS Jordan and Rothschild
(Fig. 6, £)

Atyphloceras Jordan and Rothschild, 1915, Ectoparasites 1: 59. Type, Cerato-
phyllus multidentatus C. Fox (by original designation).

Frontal tubercle angulate apically, inconspicuous. Forehead with 3 rows
of lateral setae, none of the latter being spiniform. Eyes much reduced or
vestigial. Labial palpus with 5 to 8 segments. Postantennal region of head
with 3 oblique rows of lateral setae in addition to posterior marginal row.
Pronotal comb with usual, long, black spines. Femur I with several iateral
setae; coxa III without a row or patch of setae on inner surface; segment I of
posterior tarsus shorter than’ II, III, and IV taken together; last segment of
tarsus III with 5 pairs of lateral plantar bristles, the first pair not being shifted
ventrally. Abdomen with apical spines extending down almost to level of spir-
acles. Tergal plate VII without a posterior median process. Three ante-
pygidial bristles usually on each side in female, nnd 1 large antepygidial
bristle in addition to 1 or 2 small ones in male. Female with 2 equal recepta-
cula seminis. In male sternal plate VIIf large and practically unmodified ;
sternal plate IX with large inner vertical processes, no median apophysis or a
vestigial one, and a long, setigerous, distal lobe; movable finger of clasper
long, flat, with conspicuous setae but no pigmented spines.

Species of this genus are found exclusively on small North Ameri-
can mammals.

ATYPHLOCERAS ARTIUs Jordan

Atyphloceras artius Jordan, 1933, Novitates Zool. 39: 69, fig. 19.

Type host—Peromyscus sp. (white-footed mouse).

Type locality.—British Columbia.

Range.—Known only from British Columbia.

This species was described from a single female and was said to be
close to multidentatus C. Fox, “but the bursa copulatrix of the same
long shape as in A. echis J. & R. 1915.”

ATYPHLOCERAS BISHOPI Jordan

Atyphloceras bishopi Jordan, 1933, Novitates Zool. 39: 63, figs. 11 and 12.

Type host—Blarina brevicauda (Say) (short-tailed shrew).

Type locality—F airport, N. Y.

Range.—Known only from New York.

As indicated in the original description, this species is identified
by having the labial palpus composed of five segments instead of a
larger number and by the distance of the frontal tubercle from the
fronto-genal angle “being more than one-third as long as the distance
of the anterior oral angle from the nearest point of anterior margin
of antennal groove.”

THE FLEAS OF NORTH AMERICA 39

ATYPHLOCERAS ECHIs Jordan and Rothschild

Atyphloceras echis Jordan and Rothschild, 1915, Ectoparasites 1: 59, fig. 64.

Type host—Mus. sp. (mouse).

Type locality—Paradise, Ariz.

Range.—Southwestern part of the United States.

The original description of this species was based on a single female.
The male is unknown.

ATYPHLOCERAS FELIX Jordan

Atyphloceras feliz Jordan, 1933, Novitates Zool. 39: 69, fig. 18.

Type host—Peromyscus trueit (Shufeldt) (white-footed mouse).

Type locality—Cuddy Valley, Ventura County, Calif.

Range—Known only from California.

The original description of this species, based on a single male
specimen, is accompanied by a drawing of the male terminalia. In
this species the labial palpus is seven-segmented.

ATYPHLOCERAS MULTIDENTATUS (C. Fox)
(Fig. 6, F)

Ceratophyllus multidentatus C. Fox, 1909, Ent. News 20: 107, 3 figs.
Atyphloceras multidentatus Jordan and Rothschild, 1915, Ectoparasites 1: 459.

Type host—‘Nest of Neotoma sp.” (wood rat).

Type locality—San Francisco, Calif.

Range.—Western part of the United States.

This is a medium-sized species in which the body of the penis is
much swollen and heavily pigmented.

The Genus DacryLopsyLia Jordan
(Fig. 5, A)

Dactylopsylia Jordan, 1929, Novitates Zool. 35: 37. Type, Dolichopsylla bluei
C. Fox (by original designation).

Frontal tubercle very small, angulate apically. Forehead with 2 rows of
lateral setae, none of which is spiniform. Eyes vestigial. Labial palpus 5-
segmented. Postantennal region of head with but a single row of setae, the
marginal row. Pronotal comb with the usual long, black spines. Mesosternum
not separated from mesepisternum by an inner ridge. Femur I with several
lateral setae; coxa III without a row or patch of spinelets on inner surface;
segment I of posterior tarsus shorter than II, III, and IV taken together; last
segment of each tarsus with 5 pairs of lateral plantar bristles, the first pair
of which is shifted ventrally. Abdomen with but few apical spines. Tergal
plate VII without a posterior median process. Antepygidial bristles 3 on each
Side in female, 1 in male. Female with 1 receptaculum seminis. Modified seg-
ments of male of 3 different types which are described under the 3 included
subgenera.

Although the type species of this genus was originally designated
under the generic name of Odontopsylla, it was originally described
by C. Fox (22, p. 195) in Dolichopsylla. The genus Dactylopsylla as
here recognized is composed of three subgenera. These subgenera are
practically identical except for the modified segments of the male.

40 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

The Subgenus DACTYLOPSYLLA Jordan

This subgenus is here diagnosed for the first time.

With the characters of the genus Ductylopsylla and in addition: In male,
sternal plate VIII large, divided posteriorly into a pair of large, flat lobes;
sternal plate IX with a pair of large, inner, vertical processes, a conspicuous,
anterior, median apophysis, and a pair of large, conspicuous, posterior processes,
each being divided into a proximal and a distal lobe by a broad ventral sinus;
movable finger of clasper long, slender, hooked backward at apex; penis with
dorsal portion of outer sheath modified into a long, downwardly curved, finger-
like process.

Type species.—Same as for the genus Dactylopsylla, Dolichopsylla
bluec C. Fox.

Although at first glance the modified segments of the male in this
subgenus appear to be of an entirely different type from those of the
other subgenera of Dactylopsylla, yet upon closer study the differences
observed are found to be in the shape and size of parts rather than
in the general plan of the terminalia or in the relations of the parts
to one another. This point is further discussed under the description
of the subgenus Yowxella Wagner.

DactytopsyLia (DacryLopsyLia) BLUEI (C. Fox)
(Fig. 5, A)

Dolichopsyllus bluei C. Fox, 1909, Ent. News 20: 195, figs. 1-4.
Dactylopsylla bluei Jordan, 1929, Novitates Zool. 35: 38.

Type host—Mustela xanthogenys Gray (weasel).

Type locality—San Francisco, Calif.

Range.—Known only from California.

The original description of this species, which is based on females
only,-is accompanied by a drawing of parts.

DacryLopsytia (DactryLopsyLLa) comis Jordan

Dactylopsylla comis Jordan, 1929, Novitates Zool. 35: 38, pl. 2, fig. 26.

Type host—Thomomys fuscus (Merr.) (pocket gopher).

Type locality—Okanagan Landing, British Columbia.

Range.—Known only from British Columbia.

The male of comis is unknown. According to the original descrip-
tion of the female the apical margin of sternal plate VII is rounded
and not produced into an angular lobe as in bluei.

DacryLopsyLLa (DactTyLopsyLLa) sTiMsont (C. Fox)

Ceratophyllus stimsoni C. Fox, 1914, U. S. Pub. Health Sery. Hyg. Lab. Bul.
97: 12, pl. 4, figs. 4-6.
Dactylopsylia stimsoni Jellison and Good, 1942, U. S. Pub. Health Serv. Natl.
Inst. Health Bul. 178: 49.
Type host—Thomomys sp. (pocket gopher).
Type locality.—tLos Angeles, Calif.
Range.—Known only from California.
The original description of stiémsoni was based on the male sex.
It is accompanied by drawings of the head, terminalia, and posterior
tarsus.

THE FLEAS OF NORTH AMERICA A]

The Subgenus FOXELLA Wagner

Forella Wagner, 1929, Konowia 8: 314, footnote. Type, Pulex ignotus Baker
(by original designation).

With the characters of-the genus Dactylopsylla and in addition: In male,
sternal plate VIII somewhat reduced, ending posteriorly in a pair of small,
simple lobes, each of which bears a large, heavily pigmented, marginal bristle
at its base; sternal plate IX with a pair of large, inner, vertical processes, a
conspicuous, anterior, median apophysis, and a pair of large, posterior proc-
esses, each being divided into a proximal and a distal lobe by a broad ventral
sinus; movable finger of clasper long, slender, but not hooked backward at

apex.

As already stated (see paragraph following the description of the
subgenus Dactylopsylla), the male ter minalia of the three subgenera
of the genus Dactylopsylla are essentially the same in general plan.
In Fozella sternal plate VIII is much more reduced than in the
type subgenus and possesses a very large, pigmented bristle at the
base of each distal lobe, the movable finger is not hooked backward
at the apex, and the outer sheath of the penis is less modified.

DactyLopsyLLa (FoxELLa) IGNoTA IcNoTa (Baker), new combination

Pulez ignotus Baker, 1895, Canad. Ent. 27: 110.
Typhlopsylla americana Baker, 1895, Canad. Ent. 27: 189.
yeh cares ignotus Baker, 1904, U. S. Natl. Mus. Proc. 27: 416, pl. 21,
s.
Forella ignotus Wagner, 1930, Konowia 8: 315, footnote; I. Fox, 1940, Fleas
of Eastern United States, p. 39, pl. 11, figs. 52, 54, and 55.

Type host—Unknown.

Type locality—Ames, Iowa.

Range.—The species is common in the western part of the United
States and Canada, forming several varieties. This particular
variety occurs in Iowa, Minnesota, and Illinois.

’ This variety is a rodent flea, yet Jellison (34, p. 794) found 55
individuals in a nest containing 5 half-grown fledglings of the bur-
rowing owl, Speotyto cunicularia (Molina), in the sylvatic-plague
area of Montana. In addition to the specimens of Dactylopsylla
ignota ignota, Jellison found in this same nest rodent fleas of 5 other
species, making a total of 109 flea specimens. These observations
show how important the burrowing owl may become in disseminat-
ing fleas occurring on plague-infested rodents.

DactyLopsyLta (FoxELLA) IGNOTA ALBERTENSIs (Jordan and Rothschild), new
combination

Ceratophyllus ignotus albertensis Jordan and Rothschild, 1915, Ectoparasites
1: 56, figs. 58 and 60.

Cotype hosts—Geomys sp. (pocket gopher), Mustela sp. peaiek
and Lynx canadensis Kerr (Canada lynx).

Type locality —Blackfalls, Alberta.

Range—Known only from Alberta.

In this variety the eighth sternal plate of the abdomen in the
male is shorter than in the varieties zgnota, apachina, recula, and
franciscana and has very short apical lobes. The movable finger is
provided with four long setae on its posterior margin.

42. ‘MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

DacryLopsyLta (FoxELLa) IGNoTA APACHINA (C. Fox), new combination

Ceratophyllus apachinus ©, Fox, 1914, U. S. Pub. Health Serv. Hyg. Lab.

Bul. 97: 14, pl. 5, figs. 1-3.
Ceratophyllus ignotus apachinus Jordan and Rothschild, 1915, Ectoparasites

1: 56, figs. 59 and 62.

Type host— Prairie dog.”

dope locality —“Apache County, N. Mex.”

Range.—Western part of the United States. —

The type locality of this variety was given as in the State of New
Mexico, but since there is no Apache County in New Mexico it is
probable that the specimens really came from Apache County, Ariz.
In this variety the eighth sternal plate of the male is intermediate
in relation to that of the varieties ignota and albertensis. The
movable finger of the clasper has three long setae on the posterior
margin.

DacryLopsyLLa (FoxELLA) IGNOTA FRANCISCANA (Rothschild), new
combination

Ceratophyllus franciscanus Rothschild, 1910, Ent. Monthly Mag. (2) 21: 88,
pl. 3, figs. 1 and 2.

Ceratophyllus ignotus franciscanus Jordan and Rothschild, 1915, Ectoparasites
1: 58.

Type host.—Thomomys bottae (Eyd. and Gerv.) (pocket gopher).

Type locality—San Francisco, Calif.

Range.—Known only from California.

This variety differs from the others in having a shorter process
of the clasper and a shorter movable finger. The process of the
clasper is about as broad at its base as it is high.

DacryLopsyLLA (FoxELLA) IGNOTA RECULA (Jordan and Rothschild),
new combination

Ceratophyllus ignotus recula Jordan and Rothschild, 1915, Hctoparasites 1: 58,
figs. 59 and 61.
Type host-—Mustela arizonensis (Mearns) (weasel).
Type locality —Okanagan Landing, British Columbia.
Range.—Known only from British Columbia.
This variety is characterized by having only 16 to 18 spines in the
pronotal comb and a broad emargination near the middle of the
posterior margin of the movable finger of the clasper.

DactyLopsyLLa (FoxELLA) IGNOTA UTAHENsIs (Wagner), new combination

Fozella ignotus utahensis Wagner, 1936, Ztschr. f. Parasitenk. 8: 655.

Type host—Thomomys sp. (pocket gopher).

Type locality—Wellsville, Utah.

Range—Known only from Utah.

In this variety there are three or four antepygidial bristles on each
side of the abdomen, the two lower ones being well developed. The
eighth sternal plate of the male is as in Dactylopsylla ignota apachina
(Jordan and Rothschild).

THE FLEAS OF NORTH AMERICA 43

DactyLopsyLLa (FoxELia) MExicaNA I. Fox, new combination

Fozrella mexicana I. Fox, 1939, Iowa State Col. Jour. Sci. 13: 337, pl. 1, figs. 1 and 4.

Type host —“Pocket gopher.”

Type locality —Cerro Potosi, Nuevo Leon, Mexico.

Range.—Known only from the type locality.

The original description of this species was based on two females
and is accompanied by a drawing of the head and terminalia. Ac-
cording to the original description of mexicana it is “readily separated
from Fozxella ignota (Baker) by the distinctive.chaetotaxy of the
head and by the genitalia.”

The Subgenus SPICATA I. Fox

Spicata I. Fox, 1940, Wash. Acad. Sci. Jour. 30: 275. Monotypic. Type, Dac-
tylopsylla (Spicata) rara I. Fox.

With the characters of Dactylopsylla and in addition: In male sternal plate
VIII produced posteriorly into a strong, upwardly curved process which bears
a ventral fringe of setae, the 6 postericr of which are heavily pigmented and
enlarged into stout bristles; sternal plate IX with a pair of long, inner, vertical
processes, a conspicuous median apophysis, and a pair of large, posterior pro-
cesses, each being divided into a proximal and a distal lobe; movable finger of
clasper long, slightly curved, not hooked backward at apex; manubrium narrow
and tapering distally.

As stated by I. Fox in the original description:

* +* * In the type subgenus [Dactylopsylla] sternite VIII is large but not
armed with heavy pigmented bristles. In Fogella this sternite is small and
bears but a Single long bristle near the apex. The manubrium of the new
subgenus [Spicata], being narrow and tapering distally, is markedly different
from both Fozella and Dactylopsylia (Dactylopsylla) where this structure is
short, broad, and triangular.

DactyLopsyLia (SpicaTA) RARA I. Fox

Dactylopsylla (Spicata) rara 1. Fox, 1940, Wash. Acad. Sci. Jour. 30: 275, figs.
5 and 6.

Type host—Thomomys fossor Allen (pocket gopher).

Type locality —Jackson County, Colo.

Range.—Known only from the type locality.

The original description of this species was based on a single male
and is accompanied by drawings of part of the male terminalia.

The Genus MiocTENopsytia Rothschild

(Fig. 5, B)

Mioctenopsylla Rothschild, 1922, Rpt. Norwegian Exped. Novaya Zemlya 1921,
No. 4, p. 3. Monotypic. Type, Mioctenopsylla arctica Rothschild.

Boreopsyllus Ewing, 1929, A Manual of External Parasites, pp. 168 and 201.
Monotypic. Type, Amphipsylla hadweni Ewing.

Frontal tubercle smail, almost hyaline, angulate apically. Forehead with
2 rows of lateral setae, none of which is spiniform. Eyes large, black. Labial
palpus Ssegmented, not reaching to apex of coxa I. Postantennal region of
head with 1 incomplete row of setae in addition to a complete marginal row.
Pronotal comb with very short, sharply pointed spines which are not heavier
in pigmentation than the pronotum itself. Femur I with several lateral setae;
coxa III without a row or patch of spinelets on inner surface; segment I of
posterior tarsus shorter than II, III, and IV taken together; last segment of

44 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

posterior tarsus with 5 pairs of lateral plantar bristles, the first pair not being
shifted ventrally. Abdomen with but few apical spines. Tergal plate VII with
a small, posterior, median process. Female with 1 or 2 antepygidial bristles
on each side of abdomen and a single receptaculum seminis. In male, sternal
plate VIII with large, fringed, membranous flap; sternal plate IX with large,
inner, vertical processes, an anterior median apophysis, and long, 2-lobed, posterior
processes ; movable finger of clasper large, broad, flat, somewhat rectangular and
with 3 pigmented, marginal spines.

This genus, not being given in Dalla Torre’s catalogue, was over-
looked at the time Boreopsyllus was established.

MiocTENOPSYLLA ARcTICA Rothschild
(Fig. 5, B)

Mioctenopsylla arctica Rothchild, 1922, Rpt. Norwegian Exped. Novaya Zemlya
1921, No. 4, p. 4, figs. 1-5.
Amphipsylla hadweni Ewing, 1927, Wash. Biol. Soe. Proce. 40: 89.

Type host.—Rissa tridactyla (.) (Atlantic kittiwake).

Type locality —Gribovil Fjord, Novaya Zemlya.

Range—Novaya Zemlya, Alaska, and probably most of coastal
regions inside of Arctic Circle.

Notwithstanding the fact that Amphipsylla hadweni Ewing was
described from North America, while Jioctenopsylla arctica Roths-
child came from Novaya Zemlya, the two forms appear to be identi-
cal. The species should be regarded as circumpolar. It is well illus-
trated by the figures which accompany Rothschild’s original deserip-
tion.

The Genus Turassis Jordan
(Fig. 7, 4)

Thrassis Jordan, 1933, Novitates Zool. 39: 72. Type, Ceratophyllus acamantis
Rothschild (by original designation).

Frontal tubercle small, apically angulate. Forehead with but a single row
of lateral setae, the ocular row. Eyes well developed, fully pigmented. Labial
palpus 5-segmented, extending to or beyond apex of trochanter I. Postan-
tennal region of head with a single row of setae, the marginal row. Pronotal
comb with usual, long, black spines. Femur I with several lateral setae;
coxa IiI without a row or patch of spinelets on the inner surface; segment
I of posterior torsus not longer than II, III, and IV taken together; last seg-
ment. of each tarsus with 5 pairs of lateral, plantar bristles, the first pair being
in line with the others. Abdomen with but few apical spines, all of which
are situated dorsally. Tergal plate VII with either no posterior, median process,
or a very small one. Two or 8 antepygidial bristles on each side of abdomen
in female; 1 long and 2 minute setae on each side in male. Female with 1
receptaculum seminis, the head of which is spherical or subspherical; stylet
with 2 or 8 prominent lateral bristles. In male, sternal plate VIII large,
broad, without long, filamentous, apical appendage; sternal plate IX with
long, Slender, elbowed, inner vertical processes, a somewhat enlarged anterior.
apophysis, and a posterior process which usually has 2 lobes and a pair of
pigmented spines; movable finger of clasper broad, flat, curved, with a few
long setae on posterior margin.

The postericr process of the ninth sternal plate of the male in
this genus has never been properly studied. Male specimens should
have the modified abdominal segments dissected so as to show the
structure of this process and also that of the penis, which is of almost

equal taxonomic importance. Thrassis is exclusively a Nearctic

genus. Its species parasitize various rodents.

THE FLEAS OF NORTH AMERICA 45

Turassis ACAMANTIS. (Rothschild)
(Fig. 7, A)

Ceratophyllus ucamantis Rothschild, 1905, Novitates Zool. 12: 156, pl. 8,
figs. 24 and 25.
Thrassis acamantis Jordan, 1933, Novitates Zool. 39: 72.

Cotype hosts —Mephitis occidentalis spissigrada (Bangs) (skunk),
Marmota flaviventrts (Aud. and Bach.) (woodchuck), Austela vison
energumenos (Bangs) (weasel), Canis latrans Say (coyote). t

Cotype localities —Okanagan, Sumas, and Eagle River, British
Columbia.

Range.—Western part of the United States, Canada, and Alaska.

This widely distributed species is of medium size and has marmots
for natural hosts. In it the posterior process of the ninth sternal
plate of the male bears near the ventral margin of its proximal lobe
a pair of Jong, flat, apically acuminate, modified, pigmented spines,
the shape of which is characteristic of the species.

THRASSIS ARIZONENSIS ARIZONENSIS (Baker)

Pulex arizonensis Baker, 1898, N. Y. Ent. Soc. Jour. 6: 55.
Ceratophyllus proximus Baker, 1904, U. S. Natl. Mus. Proc. 27: 412, pl. 19,
figs. 1-5.
Thassis arizonensis Jordan, 1933, Novitates Zool. 39: 73.
Type host—“Nest of silvery mouse.”
Type locality—Tucson, Ariz.
Range.—Western part of the United States.
The original description of the species ar?zonensis was based upon
a single male and unaccompanied by figures.

THRASSIS ARIZONENSIS LITTORIS (Jordan)

Ceratophyllus arizonensis littoris Jordan, 1929, Novitates Zool. 35: 31, pl. 1,
fig. 6.

Type host —‘Citellus turdicaudatus” (ground squirrel).

Type locality —San Diego, Calif.

PRange.—KKnown only from California.

The original description of /2ttoris is based upon a single female
which is like the type subspecies, “but the ventral, widened area of
tergite VIII with 21 bristles on one side and 20 on the other, in-
stead of from 14 to 16 as in C. a. arizonensis.”

Turassis BaccHI (Rothschild )

Ceratophyllus bacchi Rothschild, 1905, Novitates Zool. 12: 159, pl. 9, fig. 34.
Thrassis bacchi Jordan, 1933, Novitates Zool. 39: 73.

Type host —Citellus tridecemlineatus (Mitch.) (18-striped ground
squirrel).

Type locality—Red Deer, Alberta.

Range—Western part of the United States and Canada.

The original description of this species, based exclusively on the
male, is accompanied by drawings of the terminalia.

46 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

THRASSIS DESERTORUM Stewart

Thrassis desertorum Stewart, 1937, Canad. Wnt. 692273, fies. Wand 2:

Type host—Ammospermophilus leucurus (Merr.) (antelope
squirrel).

Type locality —Riverside County, Calif.

Range.—Known only from California.

A comparison of a male paratype of Thrassis desertorum with the
male holotype of 7. arizonensis shows that the two species are very
close to each other. Probably a further study of a large series of
specimens of each species will indicate the reduction of desertorwm to
a subspecies or variety of 7’. arizonensis.

Turassis FoTus (Jordan)

Ceratophyllus fotus Jordan, 1925, Navitates Zool. 32; 107, fig. 30.
Ceratophyllus citelli Stewart, 1928, Canad. Ent. 60: 150, figs. 3 and 4.
Thrassis fotus Jordan, 1933, Novitates Zool. 39: 73.

Type host.—Citellus tridecemlineatus (Mitch.) (13-striped ground
squirrel).

Type locality —Colorado Springs, Colo.

Range.—Western part of the United States.

The original description of this species is accompanied by a figure
of the clasper of the male and sternal plate VIII of the same sex.

TuRAssIs FRANCISI (C. Fox)

Ceratophyllus francisi C. Fox, 1927, Amer. Ent. Soc. Trans. 53: 210.
Thrassis francisi Jordan, 1933, Novitates Zool. 39: 73.

Type host.—Citellus mollis (Kenn.) (ground squirrel).

Type locality —Utah.

Range.—Known only from Utah.

In this species each spine of the pair situated on the proximal
lobe of the posterior process of sternal plate IX of the male is greatly
flattened and bladelike, with an exceedingly long and finely pointed
apex. Wagener (83, p. 343) in 1935 redescribed this species, illustrat-
ing the terminalia of the two sexes.

THRASSIS GLADIOLIS GLADIOLIS (Jordan)

Ceratophyllus gladiolis Jordan, 1925, Novitates Zool. 32: 108, figs. 32 and 33.
Thrassis gladiolis Jordan, Novitates Zool. 39: 73.

Type host —‘Citellus turdicaudatus” (ground squirrel).

Type locality —San Diego, Calif.

Range.—Known only from California.

The original description of gladiolts is accompanied by a drawing
of the clasper of the male and the posterior process of the sternal
plate IX of the same sex.

THRASSIS GLADIOLIS cADUCUS (Jordan)

Ceratophyllus gladiolis caducus Jordan, 1930, Novitates Zool. 35: 268-269, fig. 2.

Type host-——Ammospermophilus levcurus (Merr.) (antelope
squirrel).

THE FLEAS OF NORTH AMERICA 47

Type locality—Vernal, Utah.

Range-—Known only from Utah.

Very similar to type variety but: “Exopodite F [movable finger]
a little broader than in @. g. gladiolis; anterior angle of paramere
slightly less projecting * * * .”

THRASSIS HOWELLI HOWELLI (Jordan)

Ceratophyllus howelli Jordan, 1925, Novitates Zool. 32: 109, figs. 34 and 35.
Thrassis howelli Jordan, 1933, Novitates Zool. 39: 73.

Cotype hosts —Mustela arizonensis (Mearns) (weasel) and Jfar-
mota flaviventris sierrae How. (woodchuck).

Type locality—Pine City, Mono County, Calif.

Range.—F¥or the species as a whole, western part of the United
States; for this particular subspecies, California only.

Thrassis howelli is near acamantis Rothschild, but sternal plate
VIII (a figure of which accompanies the original description) is
much more slender than in the latter species. The receptaculum
seminis is as In acamantis.

TurAssIs HOWELLI UTAHENSIS Wagner

Thrassis howelli utahensis Wagner, 1936, Ztschr. f. Parasitenk. 8: 340, fig. 12.

Type host—Marmota flaviventris engelhardti (Allen) and MM. f.
nosophora How. (woodchucks).

Cotype localities —Salina and Logan, Utah.

Range.— Utah.

This subspecies is said by its author to unite Thrassis howelli
Jordan with 7. acamantis Rothschild. Drawings of the eighth ster-
num and tergum of the male accompany the original description.

THRASSIS PANDORAE Jellison

aaa genterae Jellison, 1937, U. S. Pub. Health Serv. Rpts. 52: 727, figs. 1,
, and 4.

Cotype host—Citellus elegans (Kenn.) (ground squirrel).

Type locality —P. and O. Ranch, Beaverhead County, Mont.

Range——Known only from Montana.

In this species the posterior process of the ninth sternal plate of
the male has, in addition to the usual two ventral lobes, a third
ventral lobe which is situated at its base. The basal lobe is flattened,
subdiscoidal, and without setae. The receptaculum seminis has a
seoried oveb somewhat asymmetrical head and a button on the apex
of the tal.

Turassis pANsus (Jordan)

Ceratophyllus pansus Jordan, 1925, Novitates Zool. 32: 109, fig. 38.
Thrassis pansus Jordan, 1933, Novitates Zool. 39: 73.

Type hosts—Citellus sp. (ground squirrel) and Mephitis sp.
(skunk).

Type locality —Paradise, Ariz.

Rangé.—Known only from Arizona.

The original description of this species, which is accompanied by
a aes of the distal end of the penis, is based exclusively on the
male.

48 MiSC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

Turassis PETIOLATUS (Baker)

Ceratophyllus petiolatus Baker, 1903, U. S. Natl. Mus. Proc. 27: 415, pl. 18,
figs. 7-11.
Thrassis petiolatus Jordan, 1933, Novitates Zool. 39: 73.
Type host.—Lynx canadensis Kerr (Canada lynx).
Type localityx—Moscow, Idaho.
Range.—Western part of the United States and Canada.
The type host of this species is undoubtedly not a natural one,
as the species occurs normally on rodents. The original description
of it is accompanied by illustrations but is based on a single male.

THRAssIs SPENCERI Wagner

Thrassis spenceri Wagner, 1936, Ztschr. f. Parasitenk. 8 : 655. .

Type host—Marmota sp. (woodchuck).

Type locality—Birch Island, British Columbia.

Range.—Known only from British Columbia.

This species 1s, according to its describer, most nearly related to
Thrassis acamantis Rothschild, from which it differs in having
shorter labial palpi and in a few other characters.

THRASSIS STANFORDI Wagner

Thrassis stanfordi Wagner, 1935, Ztschr. f. Parasitenk. 8: 344, figs. 15-18.

Cotype hosts—Marmota flaviventris engelhardti (Allen) and
M. f. nosophora How. (woodchucks). '

Cotype localities—F¥ish Lake, Saiima, and Logan, Utah.

Range.— Utah. ‘

This species belongs to a group of species in which the labial palpi
are very long and the parameres are rounded distally.

The Genus Oropsytta Wagner and loft
C7)

Vropsylla Wagner and Ioff, 1926, Rev. de Microbiol. et d’Epidémiol. Saratoy
5: 86 and 120. Type, Ceratophyllus silantiewi Wagner (by original desig-
nation).

Frontal tubercle small, cither rounded or angulate apically. Forehead with
1 complete ocular row of lateral setae and usually with an additional in-
compleie row. Eyes well developed, black. Labial palpus 5-segmented and
extending beyond apex of coxa I. Postantennal region of head with but a
single row of lateral setae, the posterior, marginal row. Pronotal comb with
usual long, black spines. Femur I with several lateral setae; coxa III with-
out a row or patch of spinelets on inner surface; segment I of posterior
tarsus not longer than II, III, and IV taken together; last segment of
posterior tarsus with 5 pairs of lateral, plantar bristles none of which is
shifted ventrally. Abdomer with few apical spines, all of which are dorsal
or nearly dorsal in position. Tergal plate VII without posterior, median proc-
ess. Female with 2 to + antepygidial bristles on each side of abdomen;
male with 2 on each side, 1 of which may be very small. Female with a
single receptaculum seminis. In male, sternal plate VIII much reduced and
modified; sternal plate IX with conspicuous, inner, vertical processes and an
anterior, median apophysis.

The genus Ovopsy/la, as here recognized, is composed of three
subgenera, the type subgenus Oropsy//a and the subgenera Diamanus

THE FLEAS OF NORTH AMERICA 49

Jordan and Opzsocrostis Jordan. The genus as a whole is Holarctic
-in distribution, but the subgenus Opzsocrostis is confined to the
Nearctic Region.

The Subgenus OROPSYLLA Wagner and Ioff

Typical subgenus of genus Oropsylla Wagner and Ioff. Type, Ceratophyllus
silantjewi Wagner.

With the characters of the genus Oropsylla and in addition: Frontal tubercle
acutely angulate apically and lavgely concealed in frontal notch. Labial pal-
pus long, extending beyond apex of trochanter I. In male, setae on segment
II of antenna not reaching middle of club. Basal abdominal sternal plate
without paich of bristles on side. Female with 3 or more antepygidial bristles,
male with 1 long and 1 minute bristle. Female with head of receptaculum
seminis ovate or pyriform and the tail short. In male, sternal plate VIII
narrow, rodlike, without apical membranous appendage; sternal plate IX with
posterior process divided into 4 pair of long, broad lobes, each of which is
itself divided into a proximal and a distal lobe by a deep ventral incision;
movable finger of clasper broad, flattened, somewhat clavate: spring of penis
with a single. complete convolution.

Several species, both Nearctic and Palaearctic, belong in this sub-
genus. It is particularly characterized by the length of the labial
palpus, which may extend beyond the middle of femur I in some

species.

OropsyLLA (OropsyLLa) ALASKENsIS (Baker)

Ceratophyllus alaskensis Baker, 1904, U. S. Natl. Mus. Proc. 27: 394.
Oropsylla alaskensis Wagner, 1929, Konowia 8: 313.

Type host —Citellus barrowensis (Merr.) (ground squirrel).

Type locality —Point Barrow, Alaska.

Range.——kKnown only from Alaska.

The type locality of this species represents the farthest-north
record for the order in continental North America. It is a flea above
medium size and is seldom found in flea collections.

OropsyLLa (OropsyLia) arctomys (Baker)

Ceratophyllus arctomys Baker, 1904, U. S. Natl. Mus. Proc. 27: 411, pl. 22,
figs. 1-6.

Oropsylla arctomys Wagner, 1929, Konowia 8: 315: I. Fox, 1940, Fleas of
Eastern United States, p. 45, pl. 9, figs. 40, 41, and 43.

Type host —Mermota monax (l.) (woodchuck).
Type locality.—FPeterboro, N. Y.
Range—Northern part of the United States. Canada, and Alaska.

OropsyLta (OropsyLia) IDAHOENSIS (Baker)
(Fig. 7, D)

Ceratophyllus idakoensis Baker, 1904, U. S. Natl. Mus. Proc. 27: 413, pl. 18,
figs. 1-6.

' Ceratophyllus poeantis Rothschild, 1905, Novitates Zool. 12: 155, pl. 7, figs.
22 and 23.

Ceratophulius bertholfi C. Fox, 1927, Amer. Ent. Soc. Trans. 53: 211.

Oropsylla idahoensis Wagner, 1930, Konowia 8: 314.

Type host.—Citellus columkianus (Ord) (ground squirrel).
Type locality—Moscow, Idaho.

467 459—_43—-—-4

50 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

Range—Western part of the United States, Canada, and Alaska.
_ The original description of this species appears to be in error in
stating that the frontal tubercle is lacking. An examination of the
type specimens on which it is based shows that the male and female
each have a minute, angulate, almost hyaline frontal tubercle.

OropsyLLta (ORopsyLLA) RUPEsTRIS (Jordan)

Ceratophyllus rupestris Jordan, 1929, Novitates Zool. 35: 382, pl. 1, figs. 8 and 9.
Oropsylla rupestris Wagner, 1929, Konowia 8: 314.

Cotype hosts—Mustela longicauda Bonap. (weasel) and Citellus
richardsonii (Sab.) (ground squirrel).

Type locality —Calgary, Alberta.

Range.—Western Canada.

According to the remarks given by Jordan accompanying the orig-
inal description, rwpestris 1s similar to arctomys Baker, but smaller,
and “the bristles on the whole less numerous, the lowest bristle of the
posterior row of the abdominal tergite II posterior to the stigma,
not directly below it.” The original description is accompanied by
a drawing of the genitalia of both sexes.

The Subgenus DIAMANUS Jordan

Diamanus Jordan, 1933, Novitates Zool. 39: 73. Type, Puler montanus Baker
(by original designation).

Labial palpus very long. In male, bristles of antennal segment II short, not
reaching middle of club. Sternal plate VIII of male very short and largely or
completely contamed inside of VII; sternal plate IX with each posterior process
setigerous and conspicuously bilobed because of a deep ventral emargination ;
movable finger of clasper long, flattened, somewhat swordlike, and curved for-
ward apically. Female with 2 subequal antepygidial bristles on each side of
abdomen, and with dorsolateral bristle of stylet smaller than ventral bristle.

This subgenus includes but two species, the Nearctic montanus
Baker and the Palaearctic mandarinus Jordan and Rothschild. It is
separated from the other subgenera of Oropsylla entirely on sex
characters.

OropsyLta (DiAMANUsS) MONTANA (Baker)

Pulex montanus Baker, 1895, Canad. Ent. 27: 32.
Ceratophyllus montanus Baker, 1904, U. S. Natl. Mus. Proc. 27: 411, pl. 22,

' figs. 7-8, and pl. 23, figs. 1-5.

Ceratophyllus acutus Baker, 1904, Invert. Pacifica 1: 40.

Oropsylla montanus Wagner, 1929, Konowia 8: 314; Stanford, 1931, Utah Acad.
Sci., Arts and Letters Proe. 1931: 154.

Diamanius montanus Jordan, 1933, Novitates Zool. 39: 72.

Type host— ‘Large mountain grey-squirrel (Sciurus aberti?).”

Type locality.—F¥ort Collins, Colo.

Range.—Western part of the United States and Mexico.

This squirrel-infesting species is well illustrated by Baker (1904).
It is medium sized and dark brown and can be identified by the ~
characters of the terminalia as given in the description of the sub-
genus.

The Subgenus OPISOCROSTIS Jordan
Opisocrostis Jordan, 1933, Novitates Zool. 39% 73. Type, Pulex hirsutus Baker
(by original designation).

Frontal tubercle variable in shape and size, but never large and never
acutely angulate apically but frequently narrowly rounded or broadly rounded

THE FLEAS OF NORTH AMERICA 51

at apex. In some species frontal tubercle not distinctly differentiated but
merged into frontal margin of head. Bristlies on antennal segment II long,
in female extending beyond apex of club. In male, sternal plate VIII much
reduced, slender, with 2 long, ventral, subapical bristles and a pair of setalike,
semihiyaline, filamentous, apical appendages; sternal plate IX with a pair
of conspicuous, flattened, posterior processes, each of which is divided into
a basal and a distal lobe by a deep ventral emargination; movable finger of
clasper large, broad, flat, with upper portion curved or bent somewhat forward.
In female, 2 antepygidial bristles on each side, the lower being the shorter.
Receptaculum seminis with head broader than long and tail sausage shaped.

Except for the characters of the eighth sternal plate in the male
and the antepygidial bristles in the female, Opisocrostis is il] defined,
hence is here reduced to a subgeneric status. That it constitutes a
natural group of Nearctic fleas parasitizing hosts of the rodent
family Sciuridae appears very probable.

OropsyLLa (Optsocrostis) BRUNERI (Baker)

Pulex bruneri Baker, 1895, Canad. Ent. 27: 131.

Ceratophyllus bruneri Baker, 1904, U. S. Natl. Mus. Proce. 27: 4138, pl. 25,
figs. 1-5.

Oropsylla bruneri Wagner, 1929, Konowia 8: 313.

Opisocrostis bruneri Jordan, 1933, Novitates Zool. 39: 73; I. Fox, 1940, Fleas of
eastern United States, p. 43, pl. 11, figs. 50-51, 53.

Cotype hosts —Citellus tridecemlineatus (Mitch.) (thirteen-striped
ground squirrel) and (. franklinii (Sab.) (Franklin’s ground squir-
rel.)

Cotype localities —Lincoln, Nebr., and Fort Collins, Colo.

Range.—Middlewestern part of the United States (Idaho eastward
to Illinois).

Baker (1904) published detail drawings of this species.

OropsyLta (Opisocrostis) HIRsuTA (Baker), new combination

Pulex hirsutus Baker, 1895, Canad. Ent. 27: 130.

Ceratophyllus hirsutus Baker, 1904, U. S. Natl. Mus. Proc. 27: 392, pl. 17,
figs. 1-4.

Opisocrostis hirsutus Jordan, 1933, Novitates Zool. 39: 73; Jellison, 1939,
U. S. Pub. Health Serv. Rpts. 54: 840, 3 figs.

Type host—Cynomys ludovicianus (Ord) (prairie dog).

Type locality.—Stove Prairie, Larimer County, Colo.

Range—Western part of the United States.

This species has been known as the prairie dog flea. Baker (1904)
redescribed and figured the female, and Jellison (1939) for the first
time described and figured the male.

OropsyLia (Optsocrostis) Lasis (Jordan and Rothschild)

eed labis Jordan and Rothschild, 1922, Ectoparasites 1: 275, figs.
Oropsylla labis Stewart, 1930, Canad. Ent. 62: 152.
Opisocrostis labis Jerdan, 1933, Novitates Zool. 39: 73; Jellison, 19389, U. S.
Pub. Health Serv. Rpts. 54: 841, 3 figs.

Type host—Mustela longicauda Bonap. (weasel).

Type locality—Calgary, Alberta.

Range—Western part of the United States and Canada.
_ The female originally described for this species did not belong to
it, as was pointed out by Jordan (40, p. 32) in 1929. It was in fact
the female of Oropsylla (O.) rupestris (Jordan) and has been de-
scribed, along with other material, under the name of Ceratophyllus

5? MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

rupestris by Jordan (40, p. 32). Drawings of part of the terminalia

of both sexes of /abis were given by Jellison in 1939.
OropsyLLa (OpisocrosTis) OREGONENSIS (Good and Prince), new combination

Opisocrostis oregonensis Good and Prince, 1939, U. 8. Pub. Health Serv. Rpts.
54: 1867, figs. 1-2.

Type host—Citellus oregonus (Merr.) (ground squirrel).

Type locality —Baker County, Oreg.

Range.—Oregon and Idaho.

The original description of oregonensis 1s accompanied by a draw-
ing of the posterior part of the abdomen of each sex. The species
is especially characterized by the triangular manubrium of the
clasper in the male and the very deep V-shaped emargination on
each lateral posterior margin of sternal plate VII of the female.

OropsyLia (Opisocrostis) sAUNDERSI (Jordan), new combination

Ceratophyllus saundersi Jordan, 1933, Entomologist 66: 16, figs. 6-7.
Opisocrostis saundersi Jordan, 1933, Novitates Zool. 39: 73.

Type host—Citellus richardsonii (Sab.) (ground squirrel).

Type locality — Saskatchewan, Canada.

Range.-—Known only from Canada.

The original description of swundersi is accompanied by drawings
of a part of the terminalia of each sex. The species is especially char-
acterized by the broad, straight, movable finger of the clasper in the
male and the absence of the notch usually observed in the posterior,
lateral margin of sternal plate VII of the female.

OropsyLta (OpisocrosTIs) TUBERCULATA CYNOMURIS (Jellison), new combination

Opisocrostis tuberculatus cynomuris Jellison, 1939, U. S. Pub. Health Serv.
Rpts. 54: 848, 3 figs.

Type host—Cynomys ludovicianus (Ord) (prairie dog).

Type locality—Jefferson Canyon, Broadwater County, Mont.

Range—Montana, Colorado, and Wyoming. ,

Drawings of part of the terminalia of each sex accompany the orig-
inal description of this variety. It differs from the type variety in
that the apex of the movable finger in the male is rounded, instead
of being acute as in tuberculata tuberculata, and in that the upper
lobe on the posterior margin of sternal plate VII of the female is
broad and truncate instead of being narrow and acutely angulate as
in t. tuberculata.

OropsyLLa (OPprsocrosTIs) TUBERCULATA ORNATA (J. Fox), new combination
Opisocrostis ornatus I. Fox, 1940, Wash. Ent. Soe. Proc. 42: 67, pl. 10, fig. 8.

Type host—Cynomys gunnisoni (Baird) (prairie dog).

Type locality—Saguache County, Colo.

Range—Known only from the type locality.

The original description of ornata was based on the male and is
accompanied by a drawing of the clasper. This variety is near
cynomuris, but the movable finger of the male is distinctly broadened
opposite the apex of the dorsal process, and it terminates apically
in a more acute angle. It also is near the type variety of tuberculata,
but the movable finger is stouter.

THE FLEAS OF NORTH AMERICA 53

OropsyLLA (OpisocrosTis) TUBERCULATA TUBERCULATA (Baker), new
combination

Ceratophyllus tuberculatus Baker, 1904, U. S. Natl. Mus. Proc. 27: 398, pl. 18,
gs. 7-9.
er. tuberculatus Jordan, 1933, Novitates Zool. 39: 73.

Type host—Citellus. columbianus (Ord) (ground squirrel).

Type locality—Moscow, Idaho.

Range—Western part of the United States and Canada for the
species and western part of the United States for the variety.

The original description of this variable species and its type
variety was based on a single male and female. It is accompanied
by three detail drawings. Jellison (55, p. S42) in 1939 gave draw-
ings of part of the terminalia of each sex.

OropsyLia (OPIsocRosTIS) WASHINGTONENSIs (Good and Prince), new
combination

Opisocrostis washingtonensis Good and Prince, 1939, U. S. Pub. Health Serv.
Rots. 54: 1691, figs. 3-4.

Type hosi—Citellus washingtoni How. (ground squirrel).

Type locality—Adams County, Wash.

Range.——Washington and Oregon.

This species is closely related to oregonensis but differs from the
latter in having the manubrium of the clasper of the male longer and
less triangular and in having a small, instead of :a large and deep,
notch in each posterior, lateral margin of sternal plate VII of the
female. The original description of washingtonensis is accompanied
by drawings of the terminalia of both sexes.

The Genus AmpHatius Jordan

Amphalius Jordan, 1933, Novitates Zool. 39: 74. Type, Ceratophyllus rumatus
Jordan and Rothschild (by original designation).

Frontal tubercle angulate, partly sunk into frontal notch. Eyes well devel-
oped. Labial palpus long, extending to apex of trochanter I. Setae on antennal
segment If of female long. Pronotal comb with more than 24 spines. Femur I
with several lateral setae; coxa III without a row or patch of spinelets
on inner surface; first pair of plantar bristles of segment V of each tarsus
shifted somewhat ventrally. Basal sternal plate of abdomen without lateral
bristles. Antepygidial bristles on each side of abdomen in female 38. Stylet
of female cylindrical, apex rounded, bearing several bristles. In male, sternal
plate VIII narrow, fringed on upper surface, with large, fringed, apical flap;
sternal plate IX with vertical inner process bearing a tooth above middle of
posterior margin; process of clasper long, narrow, truncate or rounded api-
cally ; movable finger broad, flat, curved forward at apex, with large, black, sub-
apical bristle and a posteroventral process which may be as long as the finger
proper.

Species of this genus are rather remarkable for the very conspicuous
posteroventral process of the movable finger of the male claspers. In
the Asiatic species clarus Jordan and Rothschild this process is
actually longer than the finger itself and is dilated apically into a
disklike lobe. Besides this Asiatic species, Amphalius contains a
Nearctic species and another Asiatic-species.

54 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

AMPHALIUS NECOPINUS (Jordan)

Ceratophyllus necopinus Jordan, 1925, Novitates Zool. 32: 110, fig. 37.
Amphalius necopinus Jordan, 1933, Novitates Zool. 39: 74.

Type host.—Ochotona schisticeps muri (Grinn. and Stor.) (cony).

Type locality.—Pine City, Mono County, Calif.

Range.—Known only from California.

The very striking characters of the claspers of the male at once
distinguish this species from all other American fleas. These char-
acters are represented in a figure which accompanies the original
description of necopinus.

The Genus Nosoprsytius Jordan
(Fig. 3)

Nosopsyllus Jordan, 1933, Novitates Zool. 39: 76. Type, Pulex fasciatus Bose
(by original designation).

Frontal tubercle small, angulate apically. Forehead with a single complete
row of setae, the ocular row. Eyes black, not reduced. Labial palpus of
medium length, scarcely reaching apex of trochanter I. Postantennal region of
head with a single row of setae, the posterior marginal row. Pronotal comb
with the usual long, dark-brown spines. Femur I with several lateral setae;
coxa III without a row or patch of spinelets on inner surface; segment I of
hind tarsus a little shorter than II and III taken together; apical bristles of
segment II of tarsus III not extending beyond apex of segment IV; tarsal
segment V of tarsus III with 5 pairs of lateral plantar bristles, none being
shifted ventrally. Abdomen with few apical spines, all of which are dorsa!
or nearly so. Tergal plate VII without posterior median process. In female,
anal sternal plate angulate in middle; a single receptaculum seminis present,
the head of which is more strongly rounded above than below. In male, sternal
plate VIII very small, without setae, and entirely contained inside of sternal
plate VII; sternal plate IX with almost straight inner vertical processes, a
conspicuous, anterior, median apophysis, and a pair of long, flat, setae-bearing,
posterior lobes, each of which is divided into proximal and distal parts by a
deep, ventral emargination; clasper with rather small manubrium, 2 large,
acetabular bristles, and a broad, flattened, distally out-curved, movable finger.

This genus is particularly noted for the extreme reduction in size
of the nonsetigerous sternal plate VIII of the male. All its con-

tained species were originally confined to the Old World. Two of
them have been introduced into America.

NosopsyLius FAsciATus (Bosc)

(Fig. 3)
Pulex fasciatus Bosc, 1801, Paris, Soc. Philomath., Bul. des Sci. 2: 156. 4
Ceratophyllus oculatus Baker, 1904, U. S. Natl. Mus. Proc. 27: 396, pl. 19, figs.
10-14. |
Ceratophyllus canadensis Baker, 1904, U. S. Natl. Mus. Proc. 27: 407, pl. 20, |
figs. 1-4.
Ceratophyllus californicus Baker, 1904, U. S. Natl. Mus. Proc. 27: 395, pl. 17,
figs. 5-8.

Nosopsyllus fasciatus Jordan, 1933, Novitates Zool. 39: 76.

Type host—‘Myoxus nitela.”

Type locality —Unknown.

FRange.—Found in all or nearly all temperate countries and in
many others.

The three species given as synonyms above were each described
from a single specimen, and in the case of canadensis the host was

THE FLEAS OF NORTH AMERICA 55

not known. Among the more recent figures of this species are those
by C. Fox (23, p. 188, fig. 65) and I. Fox (25 , pl. 20).

The rat flea, Vosopsyllus fasciatus, is of considerable importance
in the transmission of plague in certain countries. It is not a species
that bites man as readily as does Xenopsylla cheopis and is more
commonly found in temperate regions where plague is less likely to
be present.

Bacot (7), in his extended studies on the biology of some common
fleas, gave us in 1914 facts as follows for Nosopsyllus fasciatus:
Warmth (75° F.), combined with low humidity, favors the fertility
of the eggs. The newly hatched larvae were able to live in some
cases for over a month without food, provided the environment was
not otherwise unfavorable. According to Bacot there is a difference
in the kind of food necessary for fasciatus as compared with Pulex
trritans and Xenopsylla cheopis, the first mentioned apparently re-
quiring faeces of the adults as a necessity. The maximum duration
for the various stages of fasciatus was found to be “egg stage 10 days,
larval 114, cocoon 450, adult when fed 106, unfed 95.”

NosopsYLLUs LONDINIENSIS (Rothschild)

Ceratophyllus londiniensis Rothschild, 1908, Ent. Rec. and Jour. Variation 15:
64, pl. 3.

Ceratophyllus italicus Tiraboschi, 1904, Arch. de Parasitol. 8: 266, fig. 20.

Nosopsyllus londiniensis Jordan, 1933, Novitates Zool. 39: 77.

Type host—Mus musculus Li. (house mouse).

Type locality—South Kensington, London.

Range.—In North America known only from California.

The first record of the occurrence of this species in North America
was published by C. Fox (21, p. 11) in 1909.

The Genus TricHopsyLia Kolenati

TrichopsyWa Kolenati, 1863, Soc. Ent. Rossicae Horae 2: 32. Type, Pulexr
penicilliger Grube (by designation of Baker, 1904).

Malaraeus Jordan, 1933, Novitates Zool. 39: 76. Type, Ceratophyllus telchinum
Rothschild (by original designation). (New synonymy. )

Megabothris Jordan, 1933, Novitates Zool. 39: 77. Type, Ceratophyllus walkeri
Rothschild (by original designation). (New synonymy. )

Frontal tubercle small to vestigial, frequently partly contained in frontal
notch. Forehead usually with 2 or 3 rows of lateral setae, the upper or third
row, when present, usually not complete. Eyes present, pigmented, but fre-
quently reduced. labial palpus short to medium in length, not extending
beyond apex of trochanter II. Pronotal comb with usual, long, dark spines
which are usually less than 24 in number. Femur I with several lateral setae;
coxa III without a row or patch of spinelets on inner surface; inner surface
of coxae II and III with thin setae at most on distal half; last segment of
posterior tarsus with 5 pairs of lateral plantar bristles, the first of which may
be very slightly shifted or bent ventrally. Abdomen with a few apical spines,
all of which are dorsal or nearly dorsal in position. Tergal plate VII without
a posterior median process. Female with a single receptaculum seminis, the
head of which is variously shaped. Sternum VIII of male modified and fre-
quently bearing 1 or more setae; movable finger of clasper with 1 or more
black, spinelike setae.

Since the present writers recognize the first designation of the type
species of Trichopsylla by Baker (4) in 1904 as valid under the In-
ternational Rules, and this species, penicilliger Grube, is now in-

56 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

cluded among those placed in JZalaraeus Jordan, they believe the
genus Trichopsylla should displace Malaraeus.

Furthermore, the characters used to separate Malaraeus from
Megabothris can hardly be regarded as of generic importance. In
the key given by Jordan (43, p. 79) Megabothris is separated from
Malaraeus and closely related genera by the size of the stigma of
tergum VIII. Although this stigma is much enlarged in the type
species of Megabothris, M. walkeri (Rothschild), it is not so en-
larged in some of the others, and hardly makes a good distinguish-
ing character. Hence Megabothris, as well as Malar aeus, should fall
into the synonymy of 7richopsylla.

The genus J/onopsyllus Kolenati requires consideration in connec-
tion with the synonymy of Megabothris. ‘To this genus, which has
as its type species Pulex se age Schrank, 1803, Jordan (43, p. 78),
in 1933, referred a number of Nearctic species. The species are
not reearded by the authors as congeneric with the Palaearctic type
of the genus, hence the genus J/ onopsyllus Kolenati is believed not
to occur in North America. The species listed by Jordan in it from
the Nearctic Region are here given under 7'richopsylla. In 1936
Toff (33, pp. 96-98) divided Monopsyllus into three groups of species,
including in the third group the American species which had been
referred to it and one other from Siberia. To this group he gave
the name Amonopsyllus but did not designate a type species, ‘and
further stated that he did not desire to “introduce any change in
nomenclature. The species Toff included in his Amonopsyllus are so
very closely related to those in Megabothris that 1f Amonopsyllus
had a valid nomenclatorial standing there would be some justifica-
tion in regarding it as a synonym of Trichopsylla Wolenati.

The Subgenus TRICHOPSYLLA Kolenati

This subgenus has the same synonyms as given for the genus
Trichopsylla Kolenati. It is here regarded as a subgenus for the
first time.

Eyes well pigmented, frequently reduced. Bristles on segment II of male
antenna short, never reaching to more than slightly beyond middle of club.
Stigmata of abdominal segment VIII sometimes much enlarged. In male,
sternal plate VIII either much reduced and without setae or long, narrow, and
with 1 or 2 apical bristles and a membranous flap on each side; sternal plate
IX with vertical process apically rounded on posterior side and sometimes
dilated apically. In female, stylet with or without lateral bristles; head of
receptaculum seminis much longer than broad and sometimes longer than
tail.

Type species.—Same as for the genus Trichopsylla.

The type subgenus of T’richopsylla is hard to differentiate from
Pleochaetis on any one character because there is a tendency for the
characters of the two subgenera to overlap. It appears, however,
that these two subgenera represent natural groups.

The subgenus Trichopsylla, as here recognized, could be further
divided into two groups or subgenera on the basis of the male
terminalia. In one group, including penicilliger Grube, would be
placed those species in which the eighth sternal plate of the male
is long and narrow and bears apically on each side one or two bris-
tles and a membranous flap. In the other group, including such

THE FLEAS OF NORTH AMERICA 57

species as sinoma (Jordan) and telchinum (Rothschild), would be
placed those species in which the eighth sternal plate of the male is
greatly reduced, being very short, without setae, and contained in
the seventh sternal plate. This latter condition of the eighth sternal
plate of the male is the same as is found in the genus Wosopsyllus
Jordan.

The subgenus Trvichopsylla is a large one including both Palaearc-
tic and Nearctic species.

TrIcHopsyLLA (TRICHOPSYLLA) ABANTIs (Rothschild), new combination

Ceratophyllus abantis Rothschild, 1905, Novitates Zool. 12: 164, pl. 6,
fig. 10.

Cotype hosts—Mustela longicauda Bonap. (weasel) and Microtus
drummondii (Aud. and Bach.) (meadow mouse).

Cotype localities—Canadian National Park, Alberta, and Horse
Creek, Upper Columbia Valley, British Columbia.

Range.—Western Canada.

The original description of abantis, which is accompanied by a
figure of the eighth sternal plate of the male, was based on two male
specimens. The length of the male was given as 2.80 mm.

TricHopsyLLa (TRICHOPSYLLA) ACERBA (Jordan), new combination

‘Ceratophyllus acerbus Jordan, 1925, Novitates Zool. 32: 111, fig. 48.
Megabothris acerbus Jordan, 1933, Novitates Zool. 39: 77; I. Fox, 1940, Fleas
of Eastern United States, p. 69, pl. 18, figs. 91-93.

Type host—Tamias striatus L.. (eastern chipmunk).

Type locality—Canada.

Range.—Eastern part of the United States and Canada.

The original description of acerba is accompanied by a figure of
part of the female terminalia and was based on a single specimen of
that sex. This species was redescribed by I. Fox (1940), drawings
being given of parts of both sexes.

TricHopsyLLA (TRICHOPSYLLA) ADVERSA (Wagner), new combination

Megabothris adversus Wagner, 1936, Ztschr. f. Parasitenk. 8: 656, fig 4.

Type host—Peromyscus maniculatus (Wag.) (white-footed
mouse).

Type locality—Vancouver, British Columbia.

Range-—Known only from the type locality.

Only a single female specimen is known. According to the author
of this species, it stands very near to abantis Rothschild, but the duct
of the receptaculum seminis is not “chitinized.”

TrICHOPSYLLA (TRICHOPSYLLA) AsIo Asto (Baker), new combination

Ceratophyllus asio Baker, 1904, U. S. Natl. Mus. Proc. 27: 406.
Megabothris asio Jordan, 1933, Novitates Zool. 39: 77; I. Fox, 1940, Fleas of
Eastern United States, p. 67, pl. 17. figs. 85-86, 88-89.
Type host—Otus asio (1.) (screech owl).
Type locality —W ellesley, Mass.
Range.—This variety is widely distributed in the eastern part of
the United States and may occur in British Columbia, since Wagner

58 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

(84, p. 201) reported asto from that Province in 1936 without indicat-
ing the subspecies or variety.

_ The original description of asio was based on a single female. A
recent description, based on both sexes and accompanied by detail
figures of the same, was given by I. Fox (1940). The species, like
some of the others of its genus, is characterized by the peculiar nature
of the movable finger in the male. This structure is large and in the
shape of an inverted L with two toothlike, black spines at the apex.
Although aszo was originally described from an owl, birds do not con-
stitute natural hosts, which are wild mice.

TrICHOPSYLLA (TRICHOPSYLLA) ASIO ORECTA (Jordan), new combination

Megabothris asio orectus Jordan, 1938, Novitates Zool. 41: 122, figs. 72-73.

Type host —“Nest of a mouse.”

Type locality—Ravalli County, Mont.

fange.—Known only from Montana.

According to the original description of orecta, it differs from the
type subspecies in that the distal part of the movable finger in the
male is produced farther backward and the upper lobe of sternal
plate VII of the female is slightly longer.

TRICHOPSYLLA (TRICHOPSYLLA) ATROX (Jordan), new combination

Ceratophyllus atroxg Jordan, 1925, Novitates Zool. 32: 112, figs. 45-46.
Megabothris atroxr Jordan, 1933, Novitates Zool. 39: 77.

Type host—Mustela sp.

Type locality —Blackfalls, Alberta.

Range.—Known only from Alberta.

This species was supposed by Jordan to be a bird flea when he wrote
the original description. Later, in 1933, when he erected Megabothris,
he included atrox with this statement, “J/. atrox with its comb of more
than 25 spines recalls the Bird-fleas (Ceratophyllus s. str.).” In addi-
tion to having more than 25 spines in the pronotal comb, atrow is
especially characterized by the very broad and deep sinus in sternal
plate VIT of the female.

TricHopsyLia (TRicHopsyLLA) BAKERI (Wagner), new combination

Ceratophyllus bakeri Wagner, 1933, Berlin Zool. Mus. Mitt. 18: 352, fig. 13.

Type host—‘Hausmaus.”

Type locality.—Labrador.

Range.—Known only from Labrador.

Wagner (84, p. 200), in 1936, stated that this species, which is based
upon a single female, possibly represents a race of wagneri (Baker).

TricHopsyLia (TRICHOPSYLLA) BITTERROOTENSIs (Dunn), new combination
Ceratophyllus bitterrootensis Dunn, 1923, U. S. Pub. Health Serv. Rpts. 38: 2771.
Ceratophyllus isus Jordan, 1925, Novitates Zool. 32: 110, figs. 39-40.

Type host.—Neotoma cinerea (Ord) (pack rat).

Type locality—Spoon Creek, southwest of Darby, Mont.
_ Range—Western part of the United States and Canada.

THE FLEAS OF NORTH AMERICA 59

The original description of bitterrootensis was based on two male
specimens. In this species sternal plate VIII of the male is long and
slender and bears four subapical bristles.

\

- TricHopsyLLa (TRICHOPSYLLA) CILIATA CILIATA (Baker), new combination

Ceratophyllus ciliatus Baker, 1904, U. S. Natl. Mus. Proc. 27: 397, pl. 16, figs. 1-6.

Type host —‘Chipmunk.”

Type locality—Mountain View, Calif.

Range.—For the species as a whole, western part of the United
States and Canada. This particular subspecies is known from the
Western States.

The original description of this subspecies is accompanied by
detail figures of both sexes. The subspecies occurs chiefly upon chip-
munks, which apparently are the natural hosts. °

TricHopsyLia (TRICHOPSYLLA) CILIATA MONONIS (Jordan), new combination

Ceratophyllus ciliatus mononis Jordan, 1929, Novitates Zool. 35: 35, pl. 1, figs.
17-18.

Cotype hosts—Mustela arizonensis (Mearns) (weasel) and Hutam-
zas speciosus frater (Allen) (western chipmunk).

Type locality—Pine City, Mono County, Calif.

Range.—Known only from California.

In this subspecies the process of the clasper is narrower than in
either of the other subspecies and the upper lobe of sternal plate
VII of the female is broadly rounded and very much broader than
the lower lobe. The original description of mononis is accompanied
by a drawing of the clasper of the male and sternal plate VII of
the female.

TrICHOPSYLLA (TRICHOPSYLLA) CILIATA PROTINA (Jordan), new combination

oe ciliatus protinus Jordan, 1929, Novitates Zool. 35: 34, pl. 1, figs.
15-16.

Type host.—Eutamias townsendii (Bach.) (western chipmunk).

Type locality.—Sumas, British Columbia.

Range.—Known only from British Columbia and Alaska.

In the subspecies protina the process of the clasper is broader than
in mononis but narrower than in ciliata. The upper lobe of sternal
plate VII of the female is triangular with an acute apex. The
original description of protina is accompanied by a drawing of the
clasper and another of sternal plate VII of the female.

TricHopsyLLa (TRICHOPSYLLA) EREMICA (Baker), new combination

Ceratophyllus eremicus Baker, 1904, U. S. Natl. Mus. Proc. 27: 417.
Malaraeus eremicus Jordan, 1933, Novitates Zool. 39: 76.

Type host.—‘Nest of Peromyscus eremicus” (white-footed mouse).

Type locality—Santa Rita Mountains, Ariz.

Range.—Known only from Arizona.

The original description of eremica was based upon two females.
The species is especially characterized by the length of the first
segment of the hind tarsus, which exceeds that of the second, third,

60 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

and fourth combined. A note by Jordan (1933) states that the
male genitalia of eremica are remarkably different from those of its
congeners (comprising his genus Ma/araeus).

TricHopsyLia (TRICHOPSYLLA) EUMOLPI EUMOLPI (Rothschild), new combination

Ceratophyllus eumolpi Rothschild, 1905, Novitates Zool. 12: 161, pl. 6, figs. 2-4.

Cotype hosts—Eutamias minimus borealis (Allen) and E'utamias
amoenus affinis (Allen) (western chipmunks).

Cotype localities —A number of localities in Alberta and British
Columbia.

Range.—The species is known from the western part of the United
States and Canada. The range of this particular subspecies is also
the western part of the United States and Canada.

Th®é original description of ewmolpi is accompanied by drawings of
the terminalia of both sexes. This subspecies parasitizes chipmunks
by preference.

TricHopsyLLa (TRICHOPSYLLA) EUMOLPI cyrTURA (Jordan), new combination

Ceratophyllus eumolpi cyrturus Jordan, 1929, Novitates Zool. 85: 34, pl. 1, fig. 14.

Type host—Mephitis sp. (skunk).

Type locality.—Paradise, Ariz.

Range.—Known only from Arizona.

According to the original description, cyrtura differs from the
type subspecies in that the process of the clasper and the movable
finger are broader, and the second pigmented spine is much nearer
to the long one than to the short subapical pigmented spine. In the
female of cyrtura the lobe of sternal plate VII is said to be less
rounded than in the type subspecies.

TricHopsyLia (TricHopsyLLa) EUPHORBI (Rothschild), new combination

Ceratophyllus euphorbi Rothschild, 1905, Novitates Zool. 12: 165, pl. 6, fig. 11.

Type host—‘Peromyscus canadensis” (white-footed mouse).

Type locality—Horse Creek, Upper Columbia Valley, British
Columbia.

Range—Known only from British Columbia.

The original description of ewphorbi, which was based on a single
male, is accompanied by a figure of part of the terminalia. The
species appears to be rare.

TricHopsyLia (TRICHOPSYLLA) EXILIS ExILis (Jordan), new combination

Megabothris exilis Jordan, 1937, Novitates Zool. 40: 264, fig. 46.

Type host—Onychomys leucogaster (Wied.) (grasshopper
mouse).

Type locality—Powderville, Powder River County, Mont.

Range.—The species as a whole occurs in the western part of the
United States. The type subspecies occurs in Montana.

The original description of this species, based on two males, is
accompanied by a figure of the male terminalia.

THE FLEAS OF NORTH AMERICA 61

TricHopsyLLa (TRICHOPSYLLA) EXILIs OPADA (Jordan), new combination

Monopsyllus erilis opadus Jordan, 1938, Novitates Zool. 41: 121, fig. 70.

Type host—Onychomys leucogaster capitulatus Holl. (grass-
hopper mouse).

Type locality. Yavapai, Ariz.

Range—Known only from Arizona.

According to the original description of opada it differs from the
type subspecies in that the lower apical “spinform” of the movable
finger is narrower and much shorter than the upper.

TRICHOPSYLLA (TRICHOPSYLLA) EXILIs TRIPTA (Jordan), new combination

Monopsyllus exilis triptus Jordan, 1938, Novitates Zool. 41: 122, fig. 71.

Type host —-Dipodomys or Onychomys” (mice).

Type locality —Rugger, Colo. |

Range.—Known only from Colorado.

The original description of tvipta is based on two females. This
subspecies can be distinguished from the other two in that the sinus
of sternal plate VII of the female “is deeper and narrower” and
the upper lobe projects “farther distad.”

TricHopsyLia (TRICHOPSYLLA) FORNACIS (Jordan), new combination

Monopsyllus fornacis Jordan, 1957, Novitates Zool. 40: 263, fig. 45; Jellison,
1940, U. S. Pub. Heaith Serv. Rpts. 55: 491, fig. 2.

Type host —Sciurus griseus Ord (squirrel).

Type locality —Sevenoaks, Calif.

Range.——Known only trom California.

From the original description of this species, which is based on
two females and accompanied by a drawing of the female ter-
minalia, the authors quote as follows: “Closely related to W&M.
eumolpi Roths. 1905, apparently differmg only in the shape of
sternum VII and in bursa copulatrix.” The male of fornacis was
described by Jellison (1940). He states, “The male, like the female
shows very close relationship to I. eumolpi ...” In the male of
jornacis there are only two large setae on sternal plate VIII, instead
of five or six as in ewmolpi, and the lowest spinelike bristle of the
movable finger is but slightly longer than the middle instead of being
over twice as long.

TricHopsyLita (TRICHOPSYLLA) GROENLANDICA (Wahlgren), new combination

Sega rd groenlandica Wahlgren, 1903, Arkiv. fiir Zool. 1: 183, pl. 7, figs.
Type hest—‘*Myodes torquatus.”
Type locality—Kaiser Franz Josephs Fjord, East Greenland.
Range—Known only from East Greenland. The record from the
type locality for this species probably represents the farthest north
for any flea in the New World.
The original description of groenlandica was based on a single
male and is accompanied by two detail figures. In Wagner’s cata-
logue of Palaearctic Siphonaptera this species is listed (SV, p. 9) only

62 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

from East Greenland and the host is’ questioned. According to
Wahlgren’s figure of the clasper, its dorsal process is peculiar in
that it is bent or hooked backward at its apex.

TrIcHOPsyLLA (TRICHOPSYLLA) IMMITIs (Jordan), new combination

Ceratophyllus immitis Jordan, 1929, Novitates Zool. 35: 33, -pl. 1, fig. 18.

Type host—‘Lemming.”

Type locality.—Canada.

Range.—Known only from Canada.

The original description of ¢mmztis was based on the female. The
species is said to be nearest acerba (Jordan), but: “Sinus of VII
st. much shallower, the lower lobe hardly at all projecting.”

TricHopsyLia (TRICHOPSYLLA) JORDANI (I. Fox), new combination

Malaraeus jordani I. Fox, 1939, Iowa State Col. Jour. Sci. 18: 335, pl. 1,
figs, 2:

Type host.—‘Peromyscus” (white-footed mouse).

Type locality —Cerro Potosi, Nuevo Leon, Mexico.

Range.—Known only from the type locality.

The original description of jordani was based on a single male and
is accompanied by a drawing of the clasper. The species is separated
from its congeners by the details of the terminalia.

TricHopsyLLa (TRICHOPSYLLA) LucIFER (Rothschild), new combination

Ceratophyllus lucifer Rothschild, 1905, Novitates Zool. 12: 170, pl. 6, fig. 12.

Type host—Microtus drummondii (Aud. and Bach.) (meadow
mouse). ;

Type locality—Red Deer, Alberta.

Range.—Western Canada.

The original description of lwetfer was based on two female speci-
mens. According to this description and the figure which accom-
panies it, the eighth tergal plate bears two stout, spinelike setae near
apex.

TRICHOPSYLLA (TRICHOPSYLLA) MEGACOLPA (Jordan), new combination

Ceratophyllus megacolpus Jordan, 1929, Novitates Zool. 35: 33. pl. 1, fig. 12.

Type host—Microtus drummondii (Aud. and Bach.) (meadow
mouse).

Type locality—Okanagan Landing, British Columbia.

Range.—Known only from British Columbia.

The original description of this species was based on a single
female in the United Siates National Museum. The species is iden-
tified by the shape and size of the emargination or sinus of sternal
plate VII of the female. This sinus is very large and makes an
almost perfect semicircle, which in diameter is equal to about twice
the width of the lobe above it.

THE FLEAS OF NORTH AMERICA 63

TrIcHOpsYLLa (TRICHOPSYLLA) PENICILLIGER DISSIMILIs (Jordan),
new combination

Malaraeus penicilliger dissimilis Jordan, 1938, Novitates Zool. 41: 119, figs.
65-66.

Cotype hosts—Evotomys sp. (red-backed mouse) and Mzcrotus sp.
(meadow mouse).

Type locality —Rapids, Alaska.

Range—Known from Alaska and British Columbia.

The original description of dissimilis is accompanied by a drawing
of the male terminalia made by Jordan from a specimen in the United
States National Museum. The name dissimilis was suggested by
W. L. Jellison, who has supplied the same museum with a female
paratype. It applies to the American representatives of penicilliger,
which differ from the Old World subspecies particularly in that the
process of the clasper is longer and the acetabular bristles are
smaller.

TricHopsyLia (TRIcHoPsyLLA) QuirINi (Rothschild), new combination

Ceratophyllus quirini Rothschild, 1905, Novitates Zool. 12: 163, pl. 6, fig. 1.
Megabothris quirint Jordan, 1933, Novitates Zool. 39: 77; I. Fox, 1940, Fleas of
Eastern United States, p. 70, pl. 18, figs. 90, 94.

Cotype hosts—Evotomys gappert (Vigors) (red-backed mouse)
and “Evotomys saturatus.”

Type locality —Red Deer, Alberta.

Range.—United States, Canada, and Alaska.

Rothschild based his description of this species on five males and
gave a drawing of the male terminalia. I. Fox (1940) gave a re-
description of the species (accompanied by figures), which was
based on both sexes. Rothschild considered guirini as being closely
related to euwmolpi, stating that “the first midtarsal segment is long
and hairy as in euwmopli.” The peculiar shape of the movable finger,
with its broad and deep posterior emargination, identifies the species.

TricHoPsyLLa (TRICHOPSYLLA) sINoMA (Jordan), new combination

Ceratophyllus sinomus Jordan, 1925, Novitates Zool. 32: 110. fig. 41.

Type host —*Mus.”

Type locality.—Paradise, Ariz.

Range.—Western part of the United States.

This is a medium-sized, dark-brown species, identified by the
characters of the movable finger of the male clasper, which is very
broad and somewhat triangular. Its ventral margin is about
straight, posterior margin broadly outcurved, and anterior margin
broadly incurved. Near the posterior margin is a row of submar-
ginal, enlarged. dark setae, the lowest of which is longest, the upper-
most next in length. and the two others much shorter and subequal.

TricHopsyLLa ( TRICHOPSYLLA) TELCHINUM (Rothschild), new combination

Ceratophyilus telchinum Rothschild, 1905, Novitates Zool. 12: 153, pl. 8. fig. 21.

Cotype hosts—Evotomys gapperi (Vigors) (red-backed mouse)
and Sorex richardsonii Bach. (shrew).

64 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

Type locality—Kicking Horse Canyon, British Columbia.

Range.—Western part of the United States and Canada.

The original description of telehinum was based on two males and
is accompanied by a figure of the male terminalia. The movable
finger of the male bears five dark, enlarged setae near the posterior
border, the upper and lower of which are long and setiform, the
other three short, subequal, and spinform. Female specimens from
California are in the United States National Museum.

TricHopsyLia (TRICHOPSYLLA) THAMBA (Jordan), new combination

Ceratophyllus thambus Jordan, 1929, Novitates Zool. 35: 36, pl. 2, fig. 21.

Type host —Lynx sp.

Type locality —Red Deer, Alberta.

Range.—Known only from Alberta.

The original description of thamba, based on a single male, is ac-
companied by a drawing of part of the male genitalia. Jordan has
suggested that the type host is not the true host of this species. The
true host is probably a mouse.

TricHopsyLLa (TRICHOPSYLLA) vISON (BAKER), new combination

Ceratophyllus vison Baker, 1904, U. S. Natl. Mus. Proc. 27: 408.
Ceratophyllus lucidus Baker, 1904, U. S. Natl. Mus. Proce. 27: 410, pl. 20, -figs.

Megabothris vison I. Fox, 1940, Fleas of Eastern United States, p. 72, pl. 19,
figs. 95, 98-99.

Type host—Mustela vison Schreber (mink).

Type locality—Peterboro, N. Y.

Range.—United States, Canada, and Alaska.

The type host of vison is not a true host. The true hosts are squir-
rels. This flea has been recently redescribed and figured by I. Fox
(1940). The movable finger of the male has the posterior margin
incurved and bears two enlarged, darkened, subequal, submarginal
setae.

TrICHOPSYLLA (TRICHOPSYLLA) WAGNERI OPHIDIUs (Jordan), new combination
Ceratophyllus wagneri ophidius Jordan, 1929, Novitates Zool. 35: 36, pl. 2, fig. 20.

Type host —Mustela xanthogenys Gray (weasel).

Type locality—sSan Francisco, Calif.

Range.—Known only from California.

The name ophidius of this subspecies was suggested to Jordan by
the shape of the receptaculum seminis, “the organ recalling a snake.”
The same author differentiates ophidius from the other subspecies
stating that the movable finger is ‘ton the whole broader than in the
two previous subspecies [wagner and systaltus|,” and reaches “well
above process P of the clasper.” The type host probably is not a true
host of this flea.

TRICHOPSYLLA (TRICHOPSYLLA) WAGNERI sysTALTA (Jordan), new combination
Ceratophyllus wagneri systaltus Jordan, 1929, Novitates Zool. 35: 35, pl. 2,
He:

Type host—*Mouse.”
Type locality—Blackfalls, Alberta.

THE FLEAS OF NORTH AMERICA 65

Range.—Known only from Alberta.

According to J ordan’ s original description of systa/ta, the movable
finger of the male is “somewhat narrower than in the previous sub-
species [wagneri|, and its anterior margin more incurved.” A figure
of the female genitalia accompanies the original description.

°

TrIcHOPSYLLA (TRICHOPSYLLA) WAGNERI WAGNERI (Baker), new combination |

Ceratophyllus wagneri Baker, 1904, U. S. Natl. Mus. Proc. 27: 405, pl. 15,
figs. 4-7.

Ceratophyllus peromysci Stewart, 1928, Canad. Ent. 60: 148, pl. 3, figs. 1-2

Megabothris wagneri I. Fox, 1940, Fleas of Eastern United States, p. 71, pl. 19.
figs. 96-97.

Type host.—Peromyscus Wea daeits (Raf.) (white-footed mouse).

Type locality—Moscow, Idaho.

Range.—tThe species occurs in the western part of the United States
and Canada. This particular subspecies has been indicated by Jordan
(40) in 1929 as occurring in British Columbia, Idaho, Wyoming, and
western Montana. I. Fox (1940) reported it from Lowa.

The original description of wagneri was based on two males, one
of which came from the type host and one from the house mouse.
A description based on both sexes, and accompanied by figures, has
been given by I. Fox (1940). In the type subspecies the movable
finger of the clasper is intermediate in width between that of the
other two subspecies, and the head of the receptaculum seminis is
less differentiated from the tail.

The Subgenus PLEOCHAETIS Jordan

Pleochactis Jordan, 19383, Novitates Zool. 39: 77. Type, Ceratophyllus mundus
Jordan and Rothschild (by original designation).

Eyes well pigmented. Bristles on segment II of antenna short, in female not
reaching apex of club. Forehead with 2 or 3 rows of lateral setae; postan-
tennal region with 2 or more bristles behind base of antennal groove and 3
or more near middle of region. In male, sternal plate VIII slender, of vary-
ing length, with 1 or more subapical bristles and sometimes with membranous
flap. In female, stylet with 1 or 2 lateral bristles and head of receptaculum
seminis subglobular to oblong.

This subgenus is best differentiated from the type subgenus by the
chaetotaxy ‘of the head, the middle and upper rows of lateral bristles
on the forehead being well represented in some species of Pleochaetis
and poorly represented or not represented in 7'richopsylla. The male
terminalia in Pleochaetis tend to unite the two types found in the
type subgenus. Pleochaetis is essentially a Neotropical group, but
extends northward into Arizona.

TRICHOPSYLLA (PLEOCHAETIS) DOLENS (JoRDAN AND RoTHscHILD),
new combination

Ceratophyllus dolens Jordan and Rothschild, 1914, Novitates Zool. 21: 257,
figs. 1-2.
Type host—Sciurus hoffmanni (Peters) (squirrel).
Type locality—tIrazu, Costa Rica.
Range—Known only from Costa Rica,

467459—43——_5

66 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

The original description of this species was based on a single
male and female. It is accompanied by figures of the male and female
terminalia.

TRICHOPSYLLA (PLEOCHAETIS) GraPHis (Rothschild), new combination
e

Ceratophyllus graphis Rothschild,. 1909, Novitates Zool. 16: 62, pl. 10, figs. 3-4.

Type host-—Sciurus deppei Peters (squirrel).

Type locality —Nicaragua.

Range.—Known only from Nicaragua.

The original description of graphis is accompanied by a drawing of
the terminalia of each sex. The movable finger of the clasper bears
three large, subequal, dark bristles.

TRICHOPSYLLA (PLEOCHAETIS) MUNDUs (Jordan and Rothschild),
new combination

Ceratophyllus mundus Jordan and Rothschild, 1922, Ectoparasites 1: 272, fig. 266.

Type host—“Field mouse (vato de campo).”

Type locality—‘Facubaya,’ Mexico.

Range.—Known only from Mexico.

This very small flea (length of male “1.9 mm.”) was described from
a single male. The original description is accompanied by a drawing
of the male terminalia.

TRICHOPSYLLA (PLEOCHAETIS) sIByNUs (Jordan), new combination

Ceratophyllus sibynus Jordan, 1925, Novitates Zool. 32: 110, fig. 42.

Type host—Skunk.”

Type locality.—Paradise, Ariz.

Range.—Southwestern part of the United States and northern
Mexico.

This is a small fiea. The movable finger of the clasper bears a row
of five stout, darkened, submarginal setae, the uppermost of which
is the longest, the lowermost the shortest and stoutest; while the
other three are almost subequal in length, each being slightly longer
than the lowermost. A specimen of sibynus in the United States Na-
tional Museum was taken in the Mogollon Mountains, N. Mex.

The Genus CERATOPHYLLUS Curtis
(Fig. 7, E)

Ceratophyllus Curtis, 1829, A Guide to an Arrangement of British Insects,
Column 201, No. 1186; 1832, British Entomology, v. 9, No. 417; Jordan,
1933, Novitates Zool. 39: 75. Type, Ceratophyllus hirundinis Curtis (by
original designation).

The generic name Ceratophyllus was first used by Curtis in 1829,
but the genus itself was not described until 1832. For almost a cen-
tury after its establishment it included a large percentage of all the
species of fleas. Ceratophyllus is here used as it was redefined in a
restricted sense by Jordan in 1933.

Frontal tubercle small to minute, angulate at apex. Forehead frequently with
2 complete rows of lateral setae; third row never complete. Eyes black, not
reduced. Labial palpus not extending beyond tip of coxa I. Setae on antennal
segment II of female extending beyond apex of club. Postantennal region of

*

THE FLEAS OF NORTH AMERICA 67

head with 2 lateral median bristles. Pronotal comb with 24 or more long, black
spines. Femur I with several lateral setae; coxa III without a row or patch of
spinelets on inner surface; last segment of posterior tarsus with 5 pairs of
lateral plantar bristles, the first of which is not shifted ventrally. Abdomen
with a small number of apical spines which never extend down to level of
spiracles. Tergal plate VII without posterior median process. Antepygidial
bristles, 1 long and 2 minute in male, 1 long and 1 or 2 much shorter but
not minute in female. Female with 1 receptaculum seminis, the head of which
is cylindrical, concave dorsally, much longer than broad, and much longer
than tail. In male, sternal plate VIII long, slender, with stout apical bristles
and apical membranous flap; sternal plate IX with a pair of prominent, bent,
internal vertical processes, an anterior median apophysis, and a pair of distal
lobes each of which is divided by a deep ventral incision; movable finger of
clasper broad, flattened, and with 1 or more enlarged bristles or pigmented
spines on or near posterior margin.

This genus is probably fully justified as here used because of the
host relationships of its species, all of which are probably true bird
parasites. Morphologically the generic group has unity and as a
unified group can easily be described. The chief trouble in using the
genus as restricted to bird-parasitizing species is that its characters
almost completely dovetail with those of other genera. It is particu-
larly to be noted that the male terminalia of species of Ceratophullus
are of the same type as those found in some of the species of 7richop-
sylla._ This is true even when the shape and chaetotaxy of the mov-
able finger are considered. In such a species as C. riparius Jordan
and Rothschild the movable finger is broad, flattened, almost vertical,
and possesses a row of 5 stout bristles along the posterior margin.
In Trichopsylla the same condition exists in such species as 7’.
(Pleochaetis) sibynus (Jordan). The number of spines in the pro-
notal comb in Ceratophyllus is 24 or more, yet in T'richopsylla (T.)
atroz (Jordan) there are more than 24 spines in the pronotal comb.
One species of Ceratophyllus, C. adustus Jordan, may not have a
bird as a true host.

The genus Ceratophyllus, even in its restricted sense, is a large one.
It is confined entirely to the Holarctic Region.

CERATOPHYLLUs ADUsTUs Jordan
Ceratophyllus adustus Jordan, 1932, Novitates Zool. 38: 253, figs. 10-11.

Type host.—Erethizon epizanthum Brandt (American porcupine).

Type locality—Atlin, British Columbia.

Range.—Known only from British Columbia.

This species is known only from a single female which may have
been taken from an unnatural host. It is a typical bird flea except
for the reduction in the number of spinelike setae on the tibiae and
the loss of the upper one of the three antepygidial bristles. Jordan
gives two detail drawings of adustus accompanying the original de-
scription and adds:

The Porcupine probably is the true host of this modified bird-flea, the

peculiarities of the new species presumably being due to a change of host from
Bird to Mammal.

CERATOPHYLLUS BOREALIS Rothschild
Ceratophyllus borealis Rothschild, 1907, Ent. Monthly Mag. 18: 11, 1 fig.

(sus Bose * * probably came from the nest of a gannet
ula bassana).

68 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

Type locality —Island of Saint Kilda, Scotland.

Range.—In North America known only from west Greenland.

The original description of borealis, based on a single female, is
accompanied by a drawing of the terminalia. The species is said
to be particularly characterized by the shape of the seventh sternal
plate of the female: “This is truncate, with the upper angle pro-
duced into a lobe.”

CERATOPHYLLUS CELSUs APRICUs Jordan

Ceratophyllus celsus apricus Jordan, 1929, Novitates Zool. 35: 37, pl. 2, fig. 24.

Type host.—Petrochelidon fulva cavicola Bar. and Brooks (cliff
swallow). | )

Type locality—Soledad, Cuba.

Range.—Known only from Cuba.

There are several evident differences between apricus and the
typical subspecies of ce/sus. The process of the clasper is not so
slender in apricus, and it is of a different shape and bears a large
bristle at the apex. Also in apricus there are three large bristles along
the posterior margin of the movable finger instead of two. Type
specimens (male and female) of this subspecies are in the United
States National Museum.

CERATOPHYLLUS CELSUS CELSUs Jordan

Ceratophyllus celsus Jordan, 1926, Novitates Zool. 33: 387, fig. 4; I. Fox, 1940,
Fleas of Eastern United States, p. 49, pl. 12, figs. 56, 57.

Type host.—Riparia riparia (.) (bank swallow).

Type locality —Okanagan Falls, British Columbia. .
Range.—The species is known from the United States, Canada,
and Cuba. ‘This subspecies occurs only in the United States and

Canada.

The original description of ce/sus was based on a single male and
is accompanied by a drawing of the clasper. I. Fox (1940) pub-
lished a redescription of the subspecies based on both sexes and
accompanied by figures.

CERATOPHYLLUS DIFFINIs Jordan

Ceratophyllus diffinis Jordan, 1925, Novitates Zool. 32: 111, fig. 44; I. Fox,
1940, Fleas of Eastern United States, p. 50, pl. 12, figs. 58, 59.

Ceratophyllus rileyi Liu, 19385, Ent. Soc. Amer. Ann, 28: 123, pl. 1, figs. 4-6.

Type host—Colymbus holboelli (Rein.) (grebe).

Type locality—Okanagan Falls, British Columbia.

Range.—United States and Canada.

The original description and detail figure of diffinis were based on
a single male. A redescription of the species, based on both sexes, is
given by I. Fox (1940). Closely related to Ceratophyllus diffinis is
C. garet Rothschild, but difinis differs from the latter in having a
broader paramere and in the chaetotaxy of the movable finger. —

THE FLEAS OF NORTH AMERICA 69

CERATOPHYLLUS GALLINAE (Schrank)

Pulez gallinae Schrank, 1803, Fauna Boiea, v. 3, p. 195.

Pulex avium Taschenberg, 1880, Die Fléhe, p. 70, pl. 2, figs. 14-14a.

Ceratophyllus gibsoni C. Fox, 1914, U. S. Pub. Health Serv. Hyg. Lab. Bul.
97: 15, pl. 5, figs. 45. (New synonymy.)

Cotype hosts—‘Hiihnern, Miiusen, Fledermiusen, Maulwiirfen.”

Type locality —Not indicated.

Range—In North America known from the eastern part of the
United States and southeastern Canada.

This species is at times a rather serious pest of chickens and is now
found in several of the Northern States east of the Great Plains and
in the southeastern part of Canada. It was introduced into North
America apparently early in the present century, for our first record
of it (under the name of Ceratophyllus gibsoni C. Fox) was based
on specimens taken in a hen house at, Ottawa, Canada, July 13, 1909.
A few years later the species appeared in hen houses in the northeast-
ern part of the United States.

For a number of years after gallinae was encountered in North
America it was not positively identified. In 1924 one of the present
writers (Ewing, 15, p. 342) made direct comparisons between speci-
mens taken in this country and those taken in Europe and positively
identified them as gallinae. In this same paper (p. 344) Cerato-
phyllus gibsoni C. Fox was listed as a distinct species, with this
statement:

* * * The receptaculum seminis is short and stout and not at all like that
of either gallinae or niger; however, the type specimens of gibsoni are con-
siderably distorted, as if they had been boiled too long in caustic. In view
of this condition of the specimens some allowance should be made for the
distortion of the receptaculum seminis. Until more records are obtained in
the eastern part of North America for chicken infesting Ceratophylli the
sinking of this species can hardly be justified.

Since this was written further records of gallinae, both from
chickens and from other hosts, have been made, but in no case have
specimens showing the characteristics attributed to gibsont been
taken. Under these circumstances it appears that the synonymizing
of gibsoni is justified.

Wagner (80, p. 13), in his catalogue of 1930, indicates several
synonyms of this species.

CERATOPHYLLUs GAREI Rothschild
(Fig. 7, £)
Ceratophyllus garei Rothschild, 1902, Ent. Monthly Mag. (2) 13: 225, pl. 4,
figs. 1-3.
Ceratophyllus utahensis Chapin, 1919, Brooklyn Ent. Soc. Bul. 14: 60.

Type host—‘Nest of a waterhen (@allinula chloropus).”

Type locality—Near Tring, England.

Range—In North America known from western part of the United
States, Canada, and Alaska.

The male of Ceratophyllus garei is very similar to the males of
gallinae and niger, but the manubrium of the clasper is broader.
The female of C. garez is very distinct in that the head of the recepta-
culum seminis is more nearly oblong than cylindrical and hence is
not typical for the genus Ceratophyllus.

70 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

Wagner (80, p. 14) lists two additional synonyms which have been

used by European authors.

CERATOPHYLLUS cGILvuUs Jordan and Rothschild

Ceratophyllus gilvus Jordan and Rothschild, 1922, Ectoparasites 1: 270,
figs. 264-265.

Type host. —— Swallow.”

Type locality—‘Facubaya,” Mexico.

Range.—Known only from Mexico.

The original description of gi/vus, based on a single specimen of
each sex, is accompanied by drawings of the terminalia. The species
isa small, pale one, said to be “similar to C. rusticus Wa on.”

CERATOPHYLLUs IDIUs Jordan and Rothschild

Ceratophyllus idius Jordan and Rothschild, 1920, Ectoparasites 1: 73, figs. 70-72;
I. Fox, 1940, Fleas of Eastern United States, p. 51, pl. 12, figs. 60-61.
Type host —Iridoprocne bicolor (Vieill.) (tree swallow).
Type locality.—Okanagan Landing, British Columbia.
Range.—United States and Canada.
The original description of this common swallow flea is accom-
panied by drawings of the terminalia of both sexes. An important

feature of the species is that the upper half of the movable finger -

is strongly rounded.

CERATOPHYLLUS NIGER C. Fox

Ceratophyllus niger C. Fox, 1908, Ent. News 19: 434.
ae rns niger inflexrus Jordan, 1929, Novitates Zool. 35: 37, pl. 2,
pao:

Cotype hosts—Man and Rattus norvegicus (Erx.) (house rat).

Type locality.—Not indicated.

Range.—Western and northern parts of the United States, Canada,
and Alaska.

Wagner (84, pp. 200-201), in 1936, stated that he found “typical @.
niger as well as C. nig. inflexus” from “one and the same hen house’
in British Columbia, hence concluded that énflewus has no valid status
as a distinct variety.

Ceratophyllus niger is a species of some economic importance, being
a serious pest of hen houses in parts of western North America.
Control measures for this flea (under the name of Pulex avium) as a
hen-house pest have been published by Ewing (14). It readily
attacks man.

Figures of the terminalia are given by I. Fox (25, p. 155).

CERATOPHYLLUS PETROCHELIDONI Wagner

po uhuNae petrochelidoni Wagner, 1936, Ztschr. f. Parasitenk. 8: 655,
re 24

Type host.—Petrochelidon albifrons albifrons (Raf.) (a chff
swallow).

Type locality —Chilcotin, British Columbia.

Range.—British Columbia.

THE FLEAS OF NORTH AMERICA rfl

A figure of the male genitalia accompanies the original description
of Ceratophyllus petrochelidon.

CERATOPHYLLUS QUEBECENSIS I. Fox

Ceratophyllus riparius Jordan and Rothschild, 1920, Ectoparasites 1: 71, figs.
67-69

Type host—Eider down.”

Type locality—Saint Mary’s Island, Quebec.

Range—Known only from Quebec.

This species is readily separated from the other North American
species of the genus by the characters of the male and female geni-
talia, drawings of which accompany the original description.

CERATOPHYLLUS RIPARIUs Jordan and Rothschild

Ceratophyllus riparius Jordan and Rothschild, 1920, Ectoparasites 1: Tl,
figs. 67-69.

Type host—Riparia riparia riparia (l.) (bank swallow).

Type locality —Bay View, Milwaukee, Wis.

Range.—United States and Canada.

The original description of riparius is accompanied by drawings
of the terminalia of both sexes. The type host of this species is a
bird common to North America, Europe, and north Africa. In the
Old World Riparia riparia riparia is parasitized by a flea, Cerato-
phyllus styx Rothschild, which is very closely related to C. riparius,
differing only in the details of the terminalia. It is rather unusual
to find two very closely related species of ectoparasites belonging
to the same genus parasitizing the same host species. A redescription
of C. riparius is given by I. Fox (24, p. 55).

CERATOPHYLLUS SWANSONI Liu

Ceratophyllus swansoni Liu, 1935, Ent. Soc. Amer. Ann. 28: 121, pl. 1, figs. 1-3.

Type host—Asio wilsonianus (Less.) (long-eared owl).

Type locality —Fertile, Minn.

Range.—Minnesota and Montana.

Detail drawings accompany the original description of swan-
sont. It is a species belonging to the gallinae group, differing from
gallinae in having a stouter movable finger, “the exopodite [movable
finger] is less than three times as long as wide, whereas in gallinae
it 1s four times as long as wide.”

CERATOPHYLLUS VAGABUNDUs (Boheman)

Pulez vagabunda Boheman, 1865, Ofvers. af Finska Vetensk. Soc. Férhandl.
22: 576, pl. 35, fig. 1.
Ceratophyllus vagabundus Baker, 1905, U. S. Natl. Mus. Proc. 29: 152.
Type host—None.
Type locality—“Cap Todsen in Isfjorden,” Spitsbergen.
Range—In North America known only from Alaska, as reported
by Jellison and Kohls (37) in 1939.
This is a large bird flea. In Alaska it was found on fledglings
and in the nest of the golden eagle, Aguila chrysaetos. According

72 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

to Wagner (80, p. 6), digitalis Wahlgren and monedulae Wahlgren
are synonyms of vagabundus. The present writers have seen no
specimens of Wahlgren’s species.

The Genus CrENopHYLLUs Wagner
(Fig. 6, A)

Ctenophyllus Wagner, 1927, Konowia 6: 108. Type, Ceratophyllus armatus
Wagner (by subsequent designation).

Frontal tubercle small to vestigial. Forehead with’ 2 distinct rows of lateral
setae, those of the upper row being thickened and thornlike. A third indistinct
row may be between these 2 rows. Eyes present, pigmented. Labial palpus
5-segmented, not extending beyond apex of trochanter J. Setae on end of
antennal segment II in female short. Postantennal region of head with 2 or
3 rows of lateral setae. Pronotal comb with less than 24 long, black spines.
Femur I with several lateral setae; tibia and tarsal segment I of each leg
not hirsute; coxa III without a row or patch of spinelets on inner surface;
segment I of posterior tarsus shorter than II, III, and IV taken together; last
segment of each tarsus with 5 pairs of lateral plantar bristles, the first pair
being only slightly, if at all, shifted ventrally; apical bristle on segment II
of tarsus III not extending beyond apex of segment V. Abdomen with a few
apical spines, all of which are dorsal or subdorsal. Tergal plate VII without
a posterior median process. Antepygidial bristles on each side of abdomen, 3
in female. Female with a single receptaculum seminis. In male, sternal plate
VIII large, broad.

This genus contains four species of rodent-infesting fleas, three of

which are Asiatic and one North American. Three of the four
species have been found on conies (Ochotona sp.).

CTENOPHYLLUS TERRIBILIS (Rothschild)
(Fig. 6, A)

Ceratophyllus terribilis Rothschild, 1903, Novitates Zool. 10: 317, pl. 9, figs. 1-3.
Ctenophyllus terribilis Wagner, 1936, Canad. Ent. 68: 195.

Type host—Ochotona princeps (Rich.) (cony).

Cotype localities —Canadian National Park and Banff, Alberta.

Range.—Western Canada.

The original description of terribilis is accompanied by three
qo drawings, one of which represents the head and pronotum of
the male.

The Genus STENISTOMERA Rothschild
(Fig. 6, D)

Stenistomera Rothschild, 1915, Novitates Zool. 22: 307. Monotypic. Type, Typh-
lopsylla alpina Baker.

Frontal tubercle small, almost completely hidden in frontal notch. Front
somewhat angulate. Forehead with 3 irregular rows of thickened and darkened
setae. Eyes absent. Labial palpus 4-segmented, extending beyond apex of coxa
I. Postantennal region of head with 3 rows of setae. Pronotal comb with less
than 24 long, black spines. Femur I without lateral setae; each tibia with a
posterior, marginal row of heavy, black setae; coxa III without a row or patch
of spinelets on inner surface; segment I of posterior tarsus longer than II, III,
and IV taken together ; last segment of each tarsus with 5 pairs of lateral plantar.
bristles, the first pair shifted ventrally and situated between the 2 setae of
second pair. Abdomen with a few apical spines which are dorsal or subdorsal
in position. Tergal plate VII without a posterior median process. Female with
3 long, subequal, antepygidial bristles on each side of abdomen; male with 1
long (central) and 2 short antepygidial bristles on each side. Female with a

THE FLEAS OF NORTH AMERICA 73

single receptaculum seminis. In male, sternal plate VIII much reduced in
width and bearing a pair of stout, pigmented, apical spines; sternal plate IX
reduced, anterior apophysis apparently absent, distal lobes reduced.

Only a single rare species is included in this genus. In the original
diagnosis of Stentstomera, Rothschild states, “Vestige of eye exceed-
ingly slight, without pigment.” The present writers have been un-
able to find even the “vestige.”

STENISTOMERA ALPINA (Baker)
(Fig. 6, D)

Typhlopsylla alpina Baker, 1895, Canad. Ent. 27: 189.
Stenistomera alpina Rothschild, 1915, Novitates Zool. 22: 307.

Type host—‘Mountain rat.” .

Type locality— Georgetown, Colo.

Range——Known only from Colorado.

The original description of this species apparently was based on
a single male and female, now in the United States National Museum
and marked “Types of 7yphlopsylla alpina’ in Baker’s own hand-
writing. Stenistomera alpina is clothed in a striking fashion with
peculiarly shaped setae or bristles. Each is heavily pigmented and
swolien toward the base but drawn out distally into a very fine, al-
most invisible filament. Setae of this type are commonly found on
sucking lice, particularly those of the genus Hoplopleura, but are
rare on fleas. |

The Subfamily UROPSYLLINAE Oudemans

Head with fronto-epicranial groove poorly developed. Frontal notch and frontal
tubercle absent. Eyes usually well developed. Labial palpus with a small
number of segments and seldom extending beyond tip of trochanter I. Pronotum
with or without comb, and with 1 or 2 transverse rows of setae. Antepygidial
bristles present except in the genus Uropsylla. Abdomen in female never greatly
distended..:

The subfamily includes at least 10 valid genera of which only 1,
Conorhinopsylla Stewart, is represented in North America.
Uropsyllidae was proposed as a new family by Oudemans (58, p.
26) mn 1909 for the single genus Uropsylla Rothschild. It was re-
duced to a subfamily in 1929 by Ewing (106, p. 166).

The Genus CoNoRHINOPSYLLA Stewart
(Fig. 4, C)

Conorhinopsylla Stewart, 1930, Canad. Ent. 62: 178. Monotypic. Type, Conor-
hinopsylla stanfordi Stewart.

Forehead with 2 rows of lateral setae, none of which is spiniform. Eyes
vestigial, each in the form of a triangular dark spot. Gena not produced ven-
trally to form an acuminate lobe. Labial palpus 6-segmented, extending beyond
apex of coxa I but not reaching to apex of trochanter I. Setae on end of
antennal segment II short, in male not reaching to middle to club. Postantennal
region of head with but 1 row of setae, the posterior, submarginal row, the
setae of which are much reduced except for the most ventral one, which is much
enlarged. Pronotum with a single row of setae; pronotal comb present, with
less than 14 long, black spines. Tarsal segments I and II of leg III densely
hirsute with long, hairlike setae on dorsal surface; coxa III without a row or
patch of setae on inner surface; last segment of each tarsus with 5 pairs of

74 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

lateral plantar bristles, the first pair of which is shifted to a submedian
position. Abdomen without combs of spines but with a few apical spines which
are dorsal or subdorsal in position. Antepygidial bristles present; 3 on each
side in female, and 1 large bristle flanked with 2 microsetae on each side in
male. Female with 1 receptaculum seminis, the head of which is large and
oblong. In male, sternal plate VIII extended posteriorly into a long, slender
process which is armed apically with several downcurved, hooklike spines and
a small, semihyaline hood, fringed on its upper margin with several setae;
sternal plate IX with a half-circle bend instead of the usual elbow bend at
junction of distal lobe with vertical process, median apophysis vestigial, distal
lobe very long, slender, upcurved distally, and bearing setae along central part
of ventral surface; body of clasper considerably lengthened, movable finger
flattened, foliaceous, angulate apically,

This genus and its only species are so peculiar in the shape of the
head, its chaetotaxy, and the terminalia of the male that there would
be some justification in erecting a subfamily for its exclusive recep-
tion. Previous descriptions of “the male terminalia are decidedly in
error in certain respects, and although the present writers have a
male specimen for study in which the terminalia are spread apart,
yet they are not sure that the present description of them is free from
error. In order to understand the male terminalia properly in this
genus, a ventral and a posterior view of them under high magnifica-
tion is desirable. More material, making possible a dissection of the
male genitalia, is much desired.

CoNORHINOPSYLLA STANFORDI Stewart
(Fig. 4, C)
Conorhinopsylla stanfordi Stewart, 1980, Canad. Ent. 62:178, pl. 15, figs. 3-5.

Type host—Sciurus hudsonicus (Erx.) (red squirrel).

Type locality.—Ithaca, N. Y.

Range.—“astern part of the United States and as far west as Utah.

The original description of this species is accompanied by three
detail drawings. One of these is of the head and represents the labial
palpus as being five-segmented. This apparently is an error. The
labial palpus is really six-segmented as stated in a redescription of
the species by I. Fox (25, p. i 1). The male terminalia in this species
need much further study and a new description.

The Family HystricnorsyLyumar Tiraboschi

Head usually divided by a fronto-epicranial groove into anterior and posterior
parts. Gena sometimes enlarged and extended dorsally in front of antennal
groove but not divided by a groove. Forehead without helmet, seldom elongated,
and never with ventral flaps. Eyes well developed, vestigial, or absent. Cephalic
combs present. Thorax not greatly reduced; thoracic terga taken together
longer than abdominal tergum I. ‘Typical abdominal tergal plates each with
more than 1 transverse row of setae. Abdominal apical spines frequently
numerous. Abdominal combs frequently present. Females never with number
of abdominal spiracles reduced, and, when gravid, abdomen not greatly dis-
tended.

This family contains four subfamilies as follows: Ctenophthal-
minae, Hystrichopsyllinae, Macropsyllinae, and Leptopsyllinae.

Hystrichopsyllidae as a family name dates from 1905, when Baker
(5, p. 136) raised Hystrichopsyllinae Tiraboschi, 1904, to family
rank. Oudemans in 1909 (59, p. 155) attributed the authorship of

THE FLEAS OF NORTH AMERICA 75

the family to Tiraboschi. Ctenopsyllidae Baker, 1905, and Typhlo-
ceratidae Oudemans, 1900, should be regarded as synonyms of Hystri-
chopsyllidae, as also Leptopsyllidae Rothschild, 1915.

The family Hystrichopsyllidae, as here recognized, includes the
subfamily Ctenophthalminae, which had been previously included (16,
p. 166) in the Dolichopsyllidae.

The Subfamily CTENOPHTHALMINAE Rothschild

Head either divided or not divided by development of fronto-epicranial groove.
Front not greatly reduced, forming at least one-half of anterior margin of
head, and not dorsal in position. Frontal tubercle or frontal notch present,
although either or both may be vestigial. Gena seldom much enlarged and
longest in a horizontal direction. Eyes usually vestigial or absent. Labial
palpus with less than 9 segments. Cephalic comb seldom situated along both
ventral margin of gena and anterior margin of antennal groove, and never
with more than 6 spines. Hind coxae with or without a row or patch of
spinelets on inner surface. Abdomen with or without apical spines.

Included in the subfamily are 21 genera, 11 of which are repre-
sented in the North American fauna.

Ctenophthalminae was established by Rothschild (66, p. 77) in
1915, but the subfamily name as here used includes the genus Neop-
sylla Wagner, upon which the family Neopsyllidae Oudemans, 1909,
is based. The better known subfamily name Ctenophthalminae is
here retained, although it is not the oldest.

The subfamily Ctenophthalminae is believed to represent a natural
group of genera and species, although according to the classification
of Oudemans (59, p. 155) some species would fall into the suborder
Integricipita, some into Fracticipita, while others could not with any
certainty be placed into either suborder. Largely because of the
difficulty in placing the genera and species of this subfamily properly
into the suborders of Oudemans, most recent workers have abandoned
his grouping of the families. However, the character he uses for
dividing the whole order into two suborders may be used to advantage
in defining certain families and subfamilies.

Key to NortH AMERICAN GENERA OF CTENOPHTHALMINAE

1. Coxa III with a row or patch of spinelets on inner surface____________ 2
Coxa III without a row or patch of spinelets on inner surface_____- 6
Ze Seomy WALI. GTN Compile sh 22 2 Se 3
Gert comh wath. OF more spines... os 1o te: 4A ees 4

3. Plantar bristles of segment V of tarsi I and II all lateral
Tamiophila Jordan
Plantar bristles of first pair on segment V of tarsi I and II shifted
ventrally ; segment V of tarsus III with only 4 pairs of plantar brisiles
Epitedia, Jordan.
4. Segment V of each tarsus with 6 pairs of lateral plantar bDristies.
Known only from) Californians. 200 eesti bo 5 9, Actenophthalmus Fox.
Segment V of each tarsus with 4 or 5 pairs of lateral plantar bristles 5
5. Upper sclerite of metepisternum fused with metanotum
Micropsylla Dunn.
Upper sclerite of metepisternum separated from metanotum by in-
ternal chitinous ridge; metanotum sometimes fused with metepi-
iS YU | Saat GS Ft ce saree am oe Bly 2; CS oat, ae Rectofrontia Wagner.
S Geurleomy never with. more than 4 spineg 2...) 7
Genal comb with 6 spines; segment V of tarsus III with 6 lateral
plantar bristles; labial palpus with 8 segments. Known from Ore-
Paratyphtoceras Ewing.

76 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

Gi \Genal. comb with) |4 spines £255 2 2050s ee ee eee eee ee Aye eee 8
Genal comb with:3: Spimes! 2442.20 2 Eee ee er ee eens 9
8. Genal process visible above uppermost genal spine; abdominal tergal
plate VII produced into a process between the 2 sets of antepygid- .
fab bristles’ oA. e ss we a eee Corrodopsylla Wagner.
Without these characters____-- Doratopsylla Jordan and Rothschild.
9. None of spines of genal comb overlapping; tip of each palpus usually
with a terminal hooklike bristle -_-_______-_ Ctenophthalmus Kolenati.
Ventral spine of genal comb overlapped by middle spine; tip of each
palpus without terminal hooklike bristle; bristles on posterior bor-
ders of tibiae in groups of 324 Carteretta C. Fox.

The Genus TamiopHiLa Jordan
(Fig. 10, B)

Tamiophila Jordan, 1938, Novitates Zool. 41: 124. Type, Typhlopsylla grandis

Rothschild (by original designation).

Frontal tubercle acuminate. Head with a fronto-epicranial groove. Genal
comb consisting of 2 spines, of which the first is short and broad and overlaps
the more slender and much larger second one. Eyes vestigial. Labial palpus
consisting of 6 segments (counting the somewhat shortened basal segment).
Pronotum with comb. Antepygidial bristles present, at least in female. Coxa
III without a patch of spinelets on inner surface. Fifth tarsal segment of legs
I and II with 5 pairs of lateral plantar bristles. Fifth tarsal segment of leg III
with 4 pairs of lateral plantar bristles.

Two species are placed in this genus, one dbabenally so.

TAMIOPHILA GRANDIs (Rothschild)
(Fig. 10, B)
Typhlopsylla grandis Rothschild, 1902, Ent. Ree. and Jour. Variation 14: 62,
fig. 3

Neopsylla striata Stewart, 1926, Insecutor Inscitiae Menstruus 14: 124.
Tamiophila grandis Jordan, 1988, Novitates Zool. 41: 124.

Type host—Tamias striatus (.) (eastern chipmunk).

Type locality—Branchtown, Ontario.

Range.—Eastern part of the United States and Canada.

This species was first misidentified as Pulew gigas Kirby by Baker
(2, p. 163) in 1895, and for many years thereafter that name was
applied to it. In 1929, however, Jordan (41, p. 172) pointed out that
Pulew gigas Baker is identical with grandis and that Pulea gigas
Kirby is an altogether different flea belonging in the genus Hys-
trichopsylla.

The original description of grandis was accompanied by an illus-
tration of the male genitalia. The female terminalia and the head
have been figured by I. Fox (25, p. 179).

This species seems to be exceedingly rare and not often encountered
in collections. The extensive collections of the United States National
Museum include but a single female specimen.

TAMIOPHILA (?) TEXANA (Stewart), new combination

Neopsylla texana Stewart, 1930, Canad. Ent. 62: 179, pl. 15, fig. 6.

Type host—Rattus norvegicus (Erx.) (house rat).

Type locality—Houston, Tex.

Range.—Known only from Texas.

This species is placed in Z'amiophila with hesitation because of the
-chaetotaxy of the fifth tarsal segment of the hind legs. In the orig-

THE FLEAS OF NORTH AMERICA ti

inal description Stewart says, “There are four pairs of lateral plantar
bristles and a proximal median pair on the fifth tarsal segments of
the hind legs.” The members of 7'amiophila, Epitedia, and Neopsylla
are characterized by the possession of four pairs of lateral plantar
bristles, without a proximal median pair, on the fifth tarsal segment
of the hind legs. In Epitedia there are four pairs of lateral plantar
bristles and a basal ventral pair on segment V of tarsi I and II; hence
this species, which is said to have five pairs of lateral plantar bristles
on these structures, would not go into E'pitedia. The presence of five
pairs of lateral plantar bristles on the fifth segments of tarsi I and IT
is diagnostic for Zaméiophila, to which this species is provisionally
referred. ‘

Tamiophila texana (Stewart) was based upon a single female, and
no further record of its occurrence has been reported since the orig-
inal description. |

The Genus Epirepia Jordan
(Fig. 8, B)
Epitedia Jordan, 1938, Novitates Zool. 41: 124. Type, Ctenophthalmus wenmanni
Rothschild (by original designation).

Frontal notch prominent, frontal tubercle small, acuminate. Genal comb
consisting of 2 spines, of which the first is short and broad and overlaps the
more slender and much longer second one. Eyes vestigial: eye region darkly
discolored. Labial palpus consisting of 5 segments. Coxa III with a row or
patch of spinelets on inner surface. Fifth segment of tarsi I and II with 4
pairs of lateral plantar bristles and a basal ventral pair located near median
line. Fifth segment of tarsus III with 4 pairs of plantar bristles, all of which
are lateral. Base of tail of receptaculum seminis projecting well into lumen
of head.

Epitedia was the name given to a group of Nearctic species with-
drawn from Neopsyl/a by Jordan. The genus Neopsylla, estab-
lished by Wagner (79, p. 138) in 1903, included several different
genera. The type of Neopsyllais Typhlopsylla bidentatiformis Wag-
ner (78, p. 351), described in 1893, but, unfortunately, because of the
unavailability of a specimen of this species, it has not been possible
to ascertain which, if any, of the various North American species
described in Neopsylla are congeneric with the type species. Hence
Neopsylla is regarded as not occurring in our fauna, and the North
American species referred to it are considered to be members of either
Epitedia or Tamiophila, which were split off from Neopsylla by
Jordan.

Epirepia FACETA (Rothschild)

Neopsylla faceta Rothschild, 1915, Ectoparasites 1: 32, figs. 34-35.
Epitedia faceta Jordan, 1938, Novitates Zool. 41: 124.

Type host—Sciurus hudsonicus (Erx.) (red squirrel).

Type locality—Wilbraham, Mass.

Range.—Known only from Massachusetts.

This species is known from the original pair only, according to the
literature. It has not been rediscovered since its description. It is
separated from Epitedia wenmanni, its closest ally, by having the
posterior lobe of the process of the clasper higher than the anterior
and by the movable finger, which is not longer than the distance
from its base to the tip of the lower lobe of the clasper.

78 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

EpiTepia INopINA (Rothschild), new combination

Neopsylla inopina Rothschild, 1915, Ectoparasites 1: 30, figs. 32-33.

Type host.—Citellus richardsonit (Sab.) (ground squirrel).

Type locality—Calgary, Alberta.

Range—wWestern part of the United States and Canada.

In this species the fifth segment of tarsi I and IT has the basal pair
of plantar bristles shifted ventrally but not so close to the median
line as in the other species of H’pitediu. Hence it is possible that
this species goes into Veopsylla s. str., but without a close study of the
type species of Veopsylla its true status cannot be ascertained.

EpirEDIA JORDANI Hubbard

Epitedia jordani Hubbard, 1940, Pacific Univ. Bul. 37 (2): 10, 5 figs

Type host.—Nest of “Trowbridge’s shrew.”

Type locality —Newberg, Oreg.

Range—Oregon and Washington.

This species and /'pitedia stewarti are distinguished from the other
members of the genus by having the process of the clasper un-
divided. ££. stewarti and E. jordani may be separated by the struc-
ture of the terminalia, which have been illustrated in the original
descriptions.

EPITEDIA SCAPANI (Wagner), new combination

Neopsylla scapani Wagner, 1936, Ztschr. f. Parasitenk. 8: 657.

Type host—Scapanus orarius True (mole).

Type locality—vVancouver, British Columbia.

Range.—Known only from British Columbia.

George P. Holland has indicated in personal correspondence that
this species belongs in #’pitedia rather than in Veopsylla. It is based
upon two male specimens. The female has not been described.

EpIreDIA STEWARTI Hubbard

Epitedia stewarti Hubbard, 1940, Pacific Univ. Bul. 37 (2): 11, 6 figs.

Cotype hosts —“Pacifie shrews.”

Type locality —Smith River, Calif.

Range.—Recorded only from California.

Hubbard, in his original description of this species, states that it
differs from E’pitedia yordani primarily in the shape of sternal plate
1X in the male and sternal plate VII in the female.

Epirepia TEsTorR (Rothschild)

Neopsylla testor Rothschild, 1915, Ectoparasites 1: 34, fig. 36.
Epitedia testor Jordan, 1938, Novitates Zool. 41: 124.

Type host—‘Nest (of a mouse).”

Type locality—Lansingburgh, N. Y.

Range.—Known only from the type locality.

The original description of this species is based upon a single fe-
male and is accompanied by a figure of the female terminalia. The
male is unknown.

THE FLEAS OF NORTH AMERICA 79

EpITEDIA WENMANNI (Rothschild)
(Fig. 8, B)

Ctenophthalmus wenmanni Rothschild, 1904, Novitates Zool. 11: 642, pl. 14,
figs. 75, T7, 79.

Neopsylla similis Chapin, 1919, Brooklyn Ent. Soc. Bul. 14: 58.
Epitedia wenmanni Jordan, 19388, Novitates Zool. 41: 124.

Cotype hosts—Peromyscus leucepus (Raf.) (white-footed mouse)
and Neotoma cinerea (Ord) (wood rat).

Type locality—British Columbia.

Range.—Eastern part of the United States and Canada.

This species has recently been redescribed and illustrated by I. Fox
(25, pp. 96 and 183). It is the most common species of its genus in the
Eastern States.

The Genus AcTENOPHTHALMUus C. Fox
(Fig. 10, D)
Actenophthalmus C. Fox, 1925, Ent. News 36: 121. Monotypic. Type, Cten-
ophthalmus heiseri McCoy.

Frontal notch small, not acuminate. Genal comb consisting of 5 spines, of
which the uppermost is separated from the rest of the spines. Eyes vestigial.
Labial palpus 5-segmented. Coxa III with a row or patch of spinelets on
inner surface. Fifth tarsal segment of all legs armed with 6 pairs of lateral
plantar bristles. Pronotal comb present. Females with 2 antepygidial bristles
on a Side.

The members of this genus superficially resemble those of Recto-
frontia but may be immediately distinguished from them by the
presence of six pairs of lateral plantar bristles on the fifth tarsal
segments of all the legs.

This genus includes but one species.

ACTENOPHTHALMUS HEISERI (McCoy)
(Fig. 10, D)

Ctenophthalmus heiseri McCoy, 1911, Ent. News 22: 445, 2 figs.
Actenophthalmus heiseri C. Fox, 1925, Ent. News 36: 121.

Type host—Unknown.

Type locality —Mojave, Calif.

fange—Known only from type locality.

This species is based upon a single female, and no further record
of its occurrence has been made known. ‘The original description
is thoroughly illustrated.

The Genus MicropsyLttA Dunn
(Fig. 9, E)

Micropsylla Dunn, 1923, U. S. Pub. Health Serv. Rpts. 38: 2767. Monotypic.
Type, (Micropsylla peromyscus Dunn) = M. sectilis (Jordan).

Frontal tubercle acuminate. Head without a fronto-epicranial greove. Genal
comb consisting of 4 spines, of which the first is short and broad while the
others are longer and more slender. Eyes absent. Labial palpus 5-segmented.
Pronotum with a comb. Upper sclerotized ridge separating metepisternum
from metanotum lacking, the former structure being fused with the latter.
Antepygidial bristles present, at least in female. Coxa III with a patch of
spinelets on inner surface. Fifth tarsal segment of each leg with 4 pairs of
lateral plantar bristles. Tail of receptaculum seminis short, not well separated
from head.

80 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

Micropsy.ia sEcTILis (Jordan and Rothschild)
(Fig. 9, E)

Rhadinopsylla_ sectilis Jordan and Rothschild, 1923, Ectoparasites 1: 314,
fig. 318.

Micropsylia peromyscus Dunn, 1923, U. S. Pub. Health Serv. Rpts. 38: 2767,
Micropsylla sectilis Jordan, 1937, Novitates Zool. 40: 270.

Cotype hosts—Peromyscus sp. and Mus sp.

Type locality —Kelowna, British Columbia.

Range.—Western part of the United States and Canada.

The original description of this species was based on females and
is accompanied by a figure of the female terminalia. In 1936
Wagner (84, p. 203) described and illustrated the male.

The Genus Recrorrontia Wagner and Argyropulo
(Fig. 9, D)

Rectofrontia Wagner and Argyropulo, 1934, Ztschr. f. Parasitenk. 7: 230.
Type. (Rectofrontia pentactenus Wagner and Argyropulo) =Typhlopsylla
pentacanthus Rothschild (by original designation).

Frontal tubercle acuminate. Head without a fronto-epicranial groove. Genal
comb consisting of 5 spines, of which 4 are long and 1, the uppermost, is short.
Eyes absent. Labial palpus 5-segmented, last segment with 2 apical setae of
which 1 is long and hooklike and the other very short. _ Pronotum with a comb.
Upper sclerite of metepisternum separated from metanotum by internal ridge.
Males without antepygidial bristles, females with 2 or 3 on a side. Coxa III
with a patch of spinelets on inner surface. Fifth tarsal segment of each leg
with 4 pairs of lateral plantar bristles.

Tn their description of this genus Wagner and Argyropulo state, “Als
Type der Rectofrontia ist pentactenus Rothsch. (1897) zu betrach-
ten.” This is obviously a /apsus calami, since pentactenus is a bat flea
attributed to Kolenati. The name Lectofrontia pentactenus should
be attributed to Wagner and Argyropulo.

The genus fectofrontia includes but a single North American
species.

RECTOFRONTIA FRATERNA (Baker)
(Fig. 9, D)

Typhlopsylla fraterna Baker, 1895, Canad. Ent. 27: 189.
Neopsylla hamiltoni Dunn, 1923, U. S. Pub. Health Serv. Rpts. 38: 2770.
Rectofrontia fraterna Jordan, 1937, Noyitates Zool. 40: 270.

Type host—Unknown.

Type locality —Brookings, 8. Dak.

Range——United States and Canada. Reported from British
Columbia and Alberta by Holland (29, p. 47) in 1940.

The female holotype of this species is in the United States Na-
tional Museum. ‘The male terminalia were figured by I. Fox (25, p.
149) in 1940. )

The Genus CorropopsyLta Wagner
(Fig. 8, C)
Corrodopsylla Wagner, 1929, Konowia 8: 317; 1936, Canad. Ent. 68: 205. Type,
Doratopsylla curvata Rothschild (by present designation).

Although originally proposed as a subgenus of Doratopsylla
Jordan and Rothschild, Corrodopsylla was given generic status by
Wagner in 1936.

THE FLEAS OF NORTH AMERICA 81

Frontal tubercle absent. Head divided by a fronto-epicranial groove. Genal
comb consisting of 4 spines. Genal process yisible above uppermost genal
spine. Eyes vestigial, eye area heavily pigmented. Labial palpus consisting
of 4 segments. Pronotum with a comb. Abdominal tergal plate VII produced
into a process between the 2 sets of 3 antepygidial bristles. Coxa III without
a row or patch of spinelets on the inner surface. Fifth tarsal segment of
each leg armed with 4 pairs of lateral plantar bristles and a basal, submedian,
ventral pair. ;

Corrodopsylla is represented in North America by two described
species.

CorRODOPSYLLA CURVATA cuRVATA (Rothschild)
(Fig. 8, C)

Doratopsylla curvata Rothschild, 1915, Ectoparasites 1: 25, figs. 28-29.
Corrodopsylla curvata Wagner, 1936, Canad. Ent. 68: 205.

Type host—Blarina brevicauda (Say) (short-tailed shrew).

Type locality.—tlowa City, Iowa.

Range.—United States and Canada.

The original description of this subspecies is accompanied by
illustrations of the male and female terminalia.

CorRODOPSYLLA CURVATA OBTUSATA (Wagner)

Doratopsyila curvata obtusata Wagner, 1929, Konowia 8: 318.
Corrodopsylla curvata obtusata Wagner, 1936, Canad. Ent. 68: 205.

Type host.—Not given.

Type locality —British Columbia.

Range.—Known only from British Columbia.

The original description of this subspecies is based upon two
females and is accompanied by an illustration of the female termi-
nalia. The male has not yet been described.

CorRODOPSYLLA JELLISONI (Hubbard), new combination

Doratopsylla jellisoni Hubbard, 1940, Pacific Univ. Bul. 87 (2): 8, 7 figs.

Type host—“Trowbridge shrew.”

Type locality.—¥ orest Grove, Oreg.

Range.—Known only from Oregon.

This species is closely allied to Corrodopsylla curvata (Roth-
schild), from which it may be separated by the genitalia of the
male. In @. curvata the anterior of the two processes of the clasper
is subequal to the posterior; in C@. jellisoni, however, the anterior
process is much shorter than the posterior. Male and female para-
types of this species deposited in the United States National Museum
have been studied by the authors.

The Genus DoratopsyLia Jordan and Rothschild
(Fig. 8, H)

Doratopsylla Jordan and Rothschild, 1912, Novitates Zool. 19: 62. Monotypic.
Type. Typhlopsylia dasycnemus Rothschild.

Stenopsylla Cunha, 1914, Inst. Oswaldo Cruz Mem. 6: 125. Monotypic. Type,
Stenopsylla cruzi Cunha.

Tritopsylia Cunha, 1929, Sci. Med. Rio de Janeiro 7: 349. (Proposed to replace
Stenopsylla Cunha, 1914, not Stenopsylla Kuwayama, 1909.)

467459—43—_—6

82 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

Frontal tubercle absent. Head with a fronto-epicranial groove. Genal comb
consisting of 4 spines, of which the most posterior is closely appressed to
the genal border and partially covers the genal process. Genal process extend-
ing farther distad than adjacent genal spine; rounded apically. Eye vestigial;
eye area pigmented. Labial palpus consisting of 4 segments. Pronotum with
a comb. Three antepygidial bristles present on a side. Coxa III without a
patch of spinelets on inner surface. Fifth tarsal segment of each leg ‘armed
with 4 pairs of lateral plantar bristles and a basal, submedian, ventral pair.

This genus is represented in North America by two species.
According to Dalla Torre (10, p. 2) Stenopsylla Cunha is a synonym

of Doratopsylla.

DorRATOPSYLLA BLARINAE C. Fox
(Fig. 8, H)

Doratopsylla blarinae C. Fox, 1914, U. S. Pub. Health Serv. Hyg. Lab. Bul.
97: 11, pl. 4, figs. 1-3.

Type host—Blarina brevicauda (Say) (short-tailed shrew).

Type locality—Washington, D. C.

Range.—Eastern part of the United States.

This species has recently been redescribed and figured by I. Fox
(25, pp. 9 and 181).

DorATOPSYLLA INTERMEDIA COPHA (Jordan), new combination

Stenopsylla intermedia copha Jordan, 1926, Novitates Zool. 33: 391, figs. 15-16.

Type host—‘Didelphys azarae” (opossum).

Type locality.—Boquete, Panama.

Range.—Known only from Panama.

The original description of this subspecies is accompanied by
illustrations of the male and female terminalia.

DorATOPSYLLA INTERMEDIA VIDUA (Jordan), new combination

Stenopsylla intermedia vidua Jordan, 1926, Novitates Zool. 33: 392, fig. 17.

Type host—Didelphis sp. (opossum).

Type locality —Misantla, Mexico.

Range.—Known only from Mexico.

This subspecies was based upon a single female. The original
description is accompanied by an illustration of the female termin-
alia. It is exceedingly brief, reading as follows: “The sinus of ster-
nite VII (text fig. 17) very shallow, the incrassation weak, the
upper bristle far in front of the subapical pair and thin.”

The Genus CrENOPHTHALMUS Kolenati
(Fig. 9, B)

Ctenophthalmus Kolenati, 1856, Die Parasiten der Chiroptern, p. 33. Type,
Ctenophthalmus bisoctodentatus Kolenati (by subsequent designation).

Frontal tubercle large and prominent, acuminate. Head without a fronto-
epicranial groove or with a very slight one in male; head with a fronto-epicranial
groove in the female. Genal comb consisting of 3 spines, the first 2 being much
shorter than the third. Eyes vestigial. Labial palpus consisting of 5 segments,
the distal one armed with a curved apical bristle. Pronotum with a comb. Three
antepygidial bristles present on a side. Coxa III without a patch of spinelets on
inner surface. Fifth tarsal segments of legs I and II armed with 4 pairs of

THE FLEAS OF NORTH AMERICA 83

lateral plantar bristles and a basal, submedian, ventral pair. Fifth tarsal seg-
ment of leg III armed with 3 pairs of lateral plantar bristles and a basal, sub-
median, ventral pair.

This genus is represented in North American by but one species.

CTENOPHTHALMUS PSEUDAGYRTES Baker
(Fig. 9, B)

Ctenophthalmus pseudagyrtes Baker, 1904, U. S. Natl. Mus. Proc. 27: 421,
pl. 11, figs. 7-12.

Type host —Geomys bursarius (Shaw) (pocket gopher).

Type locality — Agricultural College, Mich.

Range.—FEastern part of the United States.

This species has been redescribed and figured by I. Fox (25, pp. 34
and 149). Itis one of the most common fleas occurring in the eastern
part of the United States.

The Genus CarTErETTA C. Fox
(Fig. 10, A)

Carteretta C. Fox, 1927, Amer. Ent. Soc. Trans. 53: 209. Monotypic. Type,
Carteretta carteri C. Fox.

Frontal tubercle acuminate. Head divided by a fronto-epicranial groove.
Genal comb consisting of 3 spines, of which the first is overlapped and almost
completely covered by the second. Eyes pigmented, each with a ventral emargi-
nation. Labial palpus with 5 segments. Pronotum with a comb. Three ante-
pygidial bristles present on a side. Coxa III without a row or patch of spine-
lets on inner surface. Fifth tarsal segment of each leg with 4 pairs of lateral
plantar bristles and a basal, Ssubmedian, ventral pair.

This genus includes but a single species.

CARTERETTA CARTERI C. Fox
(Fig. 10, A)

Carteretta carteri C. Fox, 1927, Amer. Ent. Soc. Trans. 53: 209.

Type host—Neotoma fuscipes Baird (wood rat).

Type locality —tLos Angeles, Calif.

Range.—Known only from California.

The original description of this species was based upon a single
male specimen which is now in the United States National Museum
collection. Both the male and female were illustrated by Jellison
(36, p. 490) in 1940.

The Genus ParatTypHLoceras Ewing
(Fig. 9, C)

Paratyphloceras Ewing 1940, Wash. Biol. Soc. Proce. 53: 35. Monotypic. Type,
Paratyphloceras oregonensis Ewing.

Frontal tubercle small, angulate apically. Gena broad, truncate apically, with
an oblique comb of 5 or 6 pointed spines. Eyes absent. Labial palpus with 8
segments. Pronotum with a comb of long spines. Coxa III without a row or
patch of:spinelets on inner surface. Fifth segment of tarsus II with 5 pairs of
lateral plantar bristles; fifth segment of tarsus III with 6 pairs of lateral
plantar bristles ; in each case first pair of plantar bristles not displaced. Abdom-
inal tergal plate VII of female produced into a short process between 2 groups
of antepygidial bristles.

84 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

This genus is nearest Typhloceras Wagner, from which it differs
in having the labial palpus composed of eight segments rather than
five. The genus includes but one species.

PARATYPHLOCERAS OREGONENSIS Ewing

(Fig. 9, C)

Paratyphloceras oregonensis Ewing, 1940, Wash. Biol. Soc. Proc. 53: 85,

Type host —Mink.” )

Type locality —Mercer Lake, Oreg.

Range.—Known only from the type locality.

This species was based upon a single female, and no further speci-
mens have been recorded. At first glance it seems to belong to one
of the genera of the Hystrichopsyllinae, but the presence of a frontal
tubercle and less than seven spines in the genal comb (fig. 9, C) .
immediately separate it from the members of that subfamily.

The Subfamily HYSTRICHOPSYLLINAE Tiraboschi

Fronto-epicranial groove distinct. Front not greatly reduced, forming at
least one-half of anterior margin of head and not dorsal in position. Gena
seldom much enlarged and nearly always longest in a horizontal direction.
Frontal tubercle and frontal notch absent. Labial palpus with small number
of segments. The genal comb may have more than 6 spines. Hind coxae with
or without a row or patch of spinelets on inner surface. The abdomen may
have 1 or more combs of spines, some of them being composed of short apical
spines.

Included in the subfamily are nine genera, four of which occur in
North America.

Tiraboschi (75, p. 296) im 1904 established the subfamily Hystri-
chopsyllinae. The next year Baker (4, p. 136) raised it to the rank
of a family. Since then many and diverse genera have been added
to the family, hence the necessity for dividing it into subfamilies.
Two of the included genera, ystrichopsylla Taschenberg and Steno-
ponia Jordan and Rothschild, contain the largest known fleas.

The interpretation of Wagner (86, p. SV) concerning the grouping
of genera about the genus Hystrichopsylla is very different from that
of the present writers, and, exclusive of Hystrichopsylla itself, no
genus is common to both concepts of the subfamily. This is due to a
difference of opinion in regard to the significance of the double
receptaculum seminis, which is the chief character in Wagner’s
diagnosis of the subfamily. Inasmuch as the double receptaculum
seminis is a sex character, the present writers are unwilling to regard
it as of more than generic significance. Even in this capacity it is
not always reliable, for it is sometimes abnormally present in species
which ordinarily have a single receptaculum seminis. For example,
in the United States National Museum collection there is a specimen
of Stenoponia americana (Baker) from Microtus pennsylvanicus at
Edgartown, Mass., which possesses a double receptaculum seminis. .

For the genus Stenoponia, Wagner has established a subfamily
which includes no other genera, and here again the writers are not
in agreement with him. The various characters used to diagnose his
subfamily Stenoponiinae are, in the opinion of the writers, of no more
than generic significance.

THE FLEAS OF NORTH AMERICA 85

Key to NortH AMERICAN GENERA OF HysTRICHOPSYLLINAE

1. Coxa III with a row or patch of spinelets on inner surface; genal
comb with only 2 spines which overlap each other__________--___-_____ 2

Coxa III without a row or patch of spinelets on inner surface________ 3

Labial palpus reaching to apex of coxa I; anterior abdominal tergal

plates with apical spines-2 27 3 Ye Phalacropsylla Rothschild.

Labial palpus not reaching beyond three-fourths the distance to apex

of coxa 1; anterior abdominal tergal plates without apical spines______
Meringis Jordan.

3. Abdomen with 1 complete comb of long spines in addition to small
apical spines; segment V of each tarsus with only 4 pairs of lateral
plantar bristles; labial palpus with only 2 segments; female with 1
receptaculum seminis__-___________ Stenoponia Jordan and Rothschild.

Abdomen without a comb of long spines, but with combs of short apical
spines; segment of V of each tarsus with 5 pairs of lateral plantar
bristles; labial palpus with 5 segments; female with 2 receptacula
SCBREE DR ee Ss eS er ee Hystrichopsylla Taschenberg.

ty

The Genus PHatacropsyLLa Rothschild

Phalacropsylla Rothschild, 1915, Ectoparasites 1: 39. Monotypic. Type, Phala-
cropsylla paradisea Rothschild.

No species belonging to this genus has been available to the authors
for study. The following redescription is compiled from the litera-
ture.

Genal comb consisting of 2 spines so arranged that 1 crosses the other. Eyes
vestigial. Labial palpus reaching to apex of coxa I. Pronotum with a comb.
Anterior abdominal tergal plates with apical spines. Three antepygidial bristles
present on a side. Coxa III with a patch of small bristles and spinelets on
the inner surface. Fifth tarsal segment of each leg with 4 pairs of lateral plantar

bristles and a basal, submedian, ventral pair. Ventral arm of sternal plate IX
with a row of several stout, spinelike setae, below which are several longer ones.

This genus includes but two species, both from North America.

PHALACROPSYLLA ALLos Wagner

Phalacropsylla allos Wagner, 1936, Ztschr. f. Parasitenk. 8: 657, fig. 6.

Type host-—Neotoma cinerea Ord (wood rat).

Type locality —Logan, Utah.

Range.——Known only from the type locality.

The original description of this species is accompanied by a figure
of the apical end of the ventral arm of sternal plate IX of the male.
In 1939 Wagner (86, p. 88) published a figure of the male head.

PHALACROPSYLLA PARADISEA Rothschild

Phalacropsylla paradisea Rothschild, 1915, Ectoparasites 1: 39, figs. 42-48.
Cotype hosts —E pimys sp. and Mus. sp. (mice).
Type locality—Paradise, Ariz.
Range—Known only from Arizona.

_ The original description of this species is accompanied by illustra-
tions of the male and female terminalia.

86 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

The Genus Merineis Jordan
(Fig. 10, C)

Meringis Jordan, 1937, Novitates Zool. 40: 268; Meringis Jordan, 1937, Novit-
ates Zool. 40: 332. Type, Meringis parkeri (Jordan) (by designation of
Jordan, 1937).

Jordan described Veringis in 1937 without type designation, but
later the same year validated the genus by designating a type.

Genal comb consisting of 2 spines, 1 of which overlaps the other. Eyes vesti-
gial. Labial palpus 5-segmented, reaching to three-fourths the length of the
fore coxa. Pronotum with comb. Anterior abdominal tergal plates without
apical spines. Three antepygidial bristles usuaily present on a side. Coxa III
with a row of spinelets on inner surface. Fifth tarsal segment of each leg
with 4 pairs of lateral plantar bristles and a basal, submedian, ventral pair.
Posterior process of sternal plate IX armed apically with 1 to several stout,
spinelike setae which are heavily pigmented.

This genus includes six species.
MERINGIS ARACHIS (Jordan)

Phalacropsylia arachis Jordan, 1929, Novitates Zool. 35: 38, pl. 2, fig. 27.
Meringis arachis Jordan, 1937, Novitates Zool. 40: 268.

Type host—Dipodomys spectabilis (Merr.) (kangaroo rat).

Type locality—McCleary’s Ranch, 30 miles southeast of Tucson,
Ariz.

Range.—Known only from Arizona.

The original description of this species is accompanied by an illus-
tration of the male terminalia.

MerINcIs cumMine! (C. Fox)

Phalacropsylla cummingi C. Fox, 1926, Pan-Pacific. Ent. 2: 182, figs. 1-2.
Meringis cummingi Jordan, 1937, Novitates Zool. 40: 268.

Type host —Dipodomys agilis (Gam.) (kangaroo rat).

Type locality—tLos Angeles, Calif.

Range.—Known only from California.

The original description of this species was based upon a single
male, now in the United States National Museum collection, and is
accompanied by illustrations of the male head and terminalia. The
female has not been described.

MeERINGIS pipopomys Kohls
(Fig. 10, C)

Meringis dipodomys Kohls, 1938, U. S. Pub. Health Serv. Rpts. 53: 1219, figs. 6-7.

Type host.—Dipodomys sp. (kangaroo rat).

Type locality —Imperial County, Calif.

Range-—Known only from California.

The original description of this species is accompanied by illustra-
tions of the male and female terminalia.

MERINGIS HUBBARDI Kohls

Meringis hubbardi Kohls, 1938, U. S. Pub. Health Sery. Rpts. 53: 1217, figs. 1-3.

Type host—Sylvilagus sp. (cottontail rabbit).
Type locality—Mayfield, Idaho.

THE FLEAS OF NORTH AMERICA 87

Range—Western part of the United States.
The original description of this species is accompanied by illustra-
tions of the male and female genitalia.

MeERINGIS PARKERI Jordan

Meringis parkeri Jordan, 1937, Novitates Zool. 40: 269, figs. 54-55.

Type host—Dipodomys sp. (kangaroo rat).

Type locality—Ravalli County, Mont.

Range—Known only from Montana.

The original description of this species is accompanied by illustra-
tions of the male and female terminalia.

MERINGIS SHANNONI (Jordan)

Phalacropsylia shannoni Jordan, 1929, Novitates Zool. 35: 38, pl. 2, figs. 28-29.
Meringis shannoni Jordan, 1937, Novitates Zool. 40: 3869-370.

Cotype hosts—Field mice.”

Type locality —Ritzville, Wash.

Range——Known only from Washington.

The original description of this species is accompanied by illus-
trations of the male terminalia and of the female receptaculum seminis.

The Genus Srenoponta Jordan and Rothschild
(Fig. 9, A) ’

Stenoponia Rothschild, 1911, [London] Zool. Soe. Proc., p. 391. Type, Hystricho-
psylla tripectinata Tiraboschi (by original designation).

Genal comb consisting of many long, slender spines. Eyes absent. Labial
palpus short, consisting of 2 segments, not extending much beyond apex of
maxilla. Pronotum with a comb consisting of numerous long, slender spines.
Abdominal tergal plate I with a comb of many long, slender spines. Abdominal
tergal plates II to V with rows of many apical spines. Antepygidial bristles
present. Females normally with only 1 receptaculum seminis. Fifth tarsal
segment of each leg with 4 pairs of lateral plantar bristles and a basal, ventral,
Submedian pair. Length, 4 or 5 mm.

The labial palpus in the members of this genus is usually described
as having but one segment. The writers, however, are of the opinion
that this structure consists of two segments, and present an illustra-
tion (fig. 9, A), which was drawn from a dissected specimen. Wagner
(86, p. 12) in 1939 published an illustration of the head of a Steno-
ponia in which the structure they believe to be the first segment of
the labial palpus is labeled the labium.

9
STENOPONIA AMERICANA (Baker)
(Fig. 9, A)
Hystrichopsylla americana Baker, 1899, Ent. News 10: 37.
Stenoponia americana Rothschild, 1915, Ectoparasites 1: 30.
Stenoponia wetmorei Chapin, 1919, Brookiyn Ent. Soc. Bul. 14: 52.

Type host.—Evotomys sp. (red-backed mouse).

Type locality—Orono, Maine.

Range.—Kastern part of the United States.

This species has been redescribed and figured by I. Fox (245, p. 81).

‘illustration is more like that of a Hystrichopsylla than anything else.

88 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

The Genus HystricHopsytia Taschenberg
(Fig. 10, F)

Hystrichopsylla Taschenberg, 1880, Die Flohe, p. 88. Type, Pulex talpae Curtis
(by original designation).

Genal comb consisting of at least 5 long, Slender spines. Eyes vestigial or
absent. Labial palpus consisting of 5 segments. Pronotum with a comb con-
sisting of many long, slender spines. Abdominal tergal plates II, III, and IV
with rows of apical spines. Antepygidial bristles present. Females with 2
receptacula seminis. Fifth tarsal segment of each leg armed with 5 pairs of
lateral plantar bristles, without a basal ventral pair. Length, 4 to 8 mm.

This genus is represented in North America by three species.

HystricHopsyLLa cicas cicas (Kirby)

Pulex gigas Kirby, 1837, Fauna Boreali-Americana 4: 318.
Hystrichopsylla gigas Jordan, 1929, Novitates Zool. 35: 174.
Type host—Unknown.
Type locality—Boreal America. ‘Two specimens taken in Lat.
fae

Range.—This subspecies is known only from the type locality.

The identity of this species has remained doubtful because of the
inadequate original descr iption, which consists of a paragraph on the
coloration and an illustration in colors of a large flea with a prom-
inent genal comb and conspicuous abdominal spines. Largely on the
basis of the great size indicated in the description, Baker (2, p. 163)
referred a flea now known to be Tamiophila grandis (Rothschild) to
this species, and the mistake was repeated by Baker (4, p. 421) in
1904 and Stewart (72, p. 869) in 1928. Jordan (1929), however, clari-
fied the situation by pointing out that the genal comb in Kirby’s

Jordan writes that—

It is, therefore, highly probable, that Pulex gigas is the same flea as Hystrichop-
sylla dippiei Roths. 1900 from Calgary, Alberta. However, before establishing
the synonymy, it would be advisable to wait for material from the North of |
Alberta and to treat H. dippiei and H. gigas for the present as subspecies of one i]
species, of which the name would be H. gigas. ;

HystTrICHOPSYLLA GIGAS DIPPIEI Rothschild |
(Fig. 10, F)
Hystrichopsylla dippiei Rothschild, 1902, Ent. Ree. and Jour. Variation 14: 63,
nes 2.
Cotype hosts—Mustela longicauda (Bonap.) (weasel) and “Lutre-
ola energamos” (mink). |
Cotype localities —Alberta; Chilliwack, British Columbia.
Range—Western part of the United States and Canada.

The original description of this subspecies is accompanied by a
figure of the male terminalia.

HystTricHOPsyLLA GIGAS TAHAVUANA Jordan

Hystrichopsylla gigas tahavuana Jordan, 1929, Novitates Zool. 35: Lie: fig. 8

Type host—Microtus pennsylvanicus (Ord) (meadow mouse).
Type locality —Adirondack Mountains, N. Y.

THE FLEAS OF NORTH AMERICA 89

Range.—Northeastern part of the United States.

The original description is accompanied by a figure of the ventral
arm of sternal plate LX of the male. The female has been redescribed
and figured by I. Fox (25, pp. 78 and 171).

HystrIicHoPpsyLLA MAMMOTH Chapin

Hystrichopsylla mammoth Chapin, 1921, Wash. Ent. Soc. Proc. 23: 25.

Type host—Aplodontia rufa californica (Peters) (mountain
beaver).

ie fee locality —Mammoth, Mono County, Calif.

Range—Western part of the United States.

The male sternal plate IX of this species was figured by Jordan
(44, p. 270). This species is characterized by its very large size, some
females being as much as 8 mm. in length.

HysTRICHOPSYLLA SCHEFFERI Chapin

Hystrichopsylla schefferi Chapin, 1919, Brooklyn Ent. Soc. Bul. 14: 50.

Type host— “Nest of Aplodontia rufa (mountain-beaver).”

Type locality—Puyallup, Wash.

Range——Known only from Washington.

This species was described from a single female specimen and
has not been rediscovered. It is distinguished from Hystichopsylla
gigas dippiet by the difference in the number of spines in the pronotal
comb, there being 46 in H. scheffert and 36 in HZ. g. dippiezt.

The Subfamily MACROPSYLLINAE Oudemans

Head divided into anterior and posterior parts by the development of
fronto-epicranial groove; postvertical tubercle present. Front not greatly re-
duced and not dorsal in position. Gena not greatly enlarged. Eyes reduced.
Labial palpus with many segments (over 10) and greater in length than head.
Cephalic comb composed of spines with rounded tips and situated both along
ventral margin of gena and anterior margin of antennal groove. Antenna
with short club, segments not anchylosed. Abdomen provided with combs.

Contains but a single genus, Macropsylla Rothschild, which is
restricted to Tasmania.

Oudemans (59, p. 155) in 1909 established the family Mac-
ropsyllidae for Macropsylla Rothschild and Stephanocircus Skuse.
The first of these is restricted to Tasmania, while the second occurs
in Australia and Tasmania. Oudemans even went further and
erected for this family the superfamily Posttuberata. This was sep-
arated from his superfamily Intuberata because of the presence of
the postvertical tubercle. More recent collections of fleas from the
Southern Hemisphere have shown that other genera possess this
tubercle and that the dividing of the genae and the presence of the
helmet in Stephanocircus ally it with certain South American genera.

The Subfamily LEPTOPSYLLINAE Rothschild

Fronto-epicranial groove usually deep, dividing head into anterior and pos-
terior parts. Front greatly reduced, anterodorsal or dorsal in position, and
forming less than one-ha!f of anterior margin of head, possessing not more
than 2 pairs of dermal pits and never more than a vestigial frontal tubercle.

90 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

Gena much enlarged and extended dorsally in front of antennal groove. Eyes
greatly reduced or absent. Cephalic comb may be situated along anterior
margin of antennal groove and composed of as few as 2 or as many as 15 or
more spines. Hind coxa with or without a row or patch of spinelets on inner

surface.

Nine genera are included in the subfamily, five being represented
in the North American fauna.

The subfamily Leptopsyllinae dates from 1915, when Rothschild
(66, p. SO) proposed for the group the family name Leptopsyllidae.
Ctenopsyllinae Wagner, 1927, based on (Ctenopsyllus Kolenati,
1863, not Kolenati, 1856)=Leptopsylla Jordan and Rothschild,
1911, is to be regarded as a synonym. Ctenopsyllidae Baker, 1905,
can hardly be regarded as a synonym, since the author included in it
the very unusual genus Stephanocircus Skuse.

Kry to NortH AMERICAN GENERA OF LEPTOPSYLLINAE

1. Cephalic comb with 2 or 4 spines ; abdominal terga without incrassations_ 2
Cephalic comb with 5 or more flattened spines arranged in a row
parallel or nearly so with anterior margin of antennal groove__ 3

2; Cephalic comb with 2)spines as eee Peromyscopsylla I. Fox

Cephalic comb with 4 spines arranged in a row at about a right
angle to anterior margin of antennal groove
Leptopsylla Jordan and Rothschild.
3. Five spines in cephalic comb; abdominal tergal plates without trans-
verse incrassation ; no sexual dimorphism in pronotal comb
Nearctopsylla Rothschild.
Six Spines in’ cephalic comb. 2 2 ee ee ee ee
4. Third spine from bottom in genal comb longer than the others and
acuminate; abdominal tergal plates with transverse incrassations
Corypsylla C. Fox.
Third spine from bottom in genal comb not long and not acuminate;
abdominal tergal plates without transverse incrassations
Corypsylloides Hubbard.

The Genus PeromyscopsyLia I. Fox

Peromyscopsylla I. Fox, 1939, Wash. Ent. Soe. Proce. 41: 47. Type, Ctenopsyllus
hesperomys Baker (by original designation).

Head subangulate in front; anterior margin with a series of bristles, some
of which may be heavily pigmented. Genal comb consisting of 2 spines varying
in size and shape according to species. Genal process usually conspicuous, vary-
ing in size and shape. Eyes vestigial. Labial palpus consisting of 5 segments.
Pronotum with a comb of many long, slender spines. Abdominal terga each with
1 to several apical spines. Coxa III without a patch or row of spinelets on inner
surface. Fifth tarsal segment of each leg with 4 pairs of lateral plantar bristles
and a baSal ventral pair.

This genus is represented in North America by eight species.

PEROMYSCOPSYLLA ADELPHA (Rothschild)

Leptopsylla adelpha Rothschild, 1915, Novitates Zool. 22: 304, fig. 4.
Peromyscopsylla adelpha Jellison and Good, 1942, U. S. Pub. Health Serv. Natl.
Inst. Health Bul. 178; 121.

Type host—Mus sp. (mouse).

Type locality—Paradise, Ariz.

Range.—Known only from Arizona.

The original description of this species is accompanied by a figure
of the female sternal plate VII. Jordan and Rothschild (49, p. 59)
published an illustration of the male terminalia in 1915.

THE FLEAS OF NORTH AMERICA O1

PEROMYSCOPSYLLA CATATINA (Jordan)

Leptopsylla catatina Jordan, 1928, Novitates Zool. 34: 186, fig. 10.
Peromyscopsylla catatina I. Fox, 1940, Fleas of Eastern United States, p. 87,
pl. 22, figs. 110, 114, and 115.
Type host—Didelphis virginiana Kerr (opossum).
Type locality—Rolling Rock Club, Ligonier, Pa.
Range.—Kastern part of the United States and Labrador.

PEROMYSCOPSYLLA EBRIGHTI (C. Fox)

Leptopsylla ebrighti C. Fox, 1926, Pan-Pacifie Ent. 2: 182, figs. 3-4.
Peromyscopsylla ebrighti Jellison and Good, 1942, U. S. Pub. Health Serv. Natl.
Inst. Health Bul. 178: 121.

Type host.—Neotoma fuscipes Baird (wood rat).

Type locality—tLos Angeles, Calif.

Fange.—Known only from California.

The original description of this species is accompanied by illustra-
tions of the male and female terminalia.

PEROMYSCOPSYLLA HAMIFER HAMIFER (Rothschild)

Ctenopsyllus hamifer Rothschild, 1906, Canad. Ent. 38: 324, fig. 44.
Peromyscopsylla hamifer hamifer Jellison and Good, 1942, U. S. Pub. Health
Serv. Nati. Inst. Health Bul. 178: 121.

Type host—Mustela sp. (weasel).

ype locality.—Blacktfalls, Alberta.

Range.—The species is known from the western part of the United
States and Canada, the typical subspecies only from Alberta.

The original description of this subspecies, which was based on a
single male, i is accompanied by an illustration of the male terminalia.
No records of its occurrence have been published since the original
description, and the female has not been described.

PEROMYSCOPSYLLA HAMIFER LONGILOBA (Jordan), new combination
Leptopsylla hamifer longiloba (Jordan), 1939, Novitates Zool. 41: 319.

Type host—Microtus sp. (meadow mouse).

Type locality.—V aldez Creek Trail, 3,500 ft., Alaska.

Range.—Known only from Alaska.

The male of this subspecies is unknown. The original description
is accompanied by an illustration of the female terminalia.

PEROMYSCOPYLLA HAMIFER VIGENS (Jordan)

Leptopsylla hamifer vigens Jordan, 1937, Novitates Zool. 40: 265, figs. 47-48.
Peromyscopsylla hamifer vigens Jellison and Good, 1942, U. S. Pub. Health Serv.
Natl. Inst. Health Bul. 178: 122.

Type host—Microtus sp. (meadow mouse).

Type locality.—Ravalli County, Mont.

Range.—Known only from Montana.

The original description of this subspecies is accompanied by illus-
trations of the male and female terminalia.

PEROMYSCOPSYLLA HESPEROMYs (Baker)
(Fig. 8, E)

Ctenopsyllus hesperomys Baker, 1904, U. S. Natl. Mus. Proc. 27: 428.

Peromyscopsylla hesperomys I. Fox, 1929, Wash. Ent. Soe. Proe. 41: 49.

se AN el Atay spinifrons I. Fox, 1989, Wash. Ent. Soc. Proc. 41: 48, pl. 6,
gs. 1-2.

92 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

Type host.—Peromyscus sp. (white-footed mouse).

Type locality—Franconia, N. H.

Range—FKastern part of the United States.

This species has been redescribed and illustrated by I. Fox (25,
pp. 84 and 175), and the above synonymy indicated.

PEROMYSCOPSYLLA RAVALLIENSIS (Dunn) »

Ctenopsyllus ravalliensis Dunn, 1923, U. S. Pub. Health Serv. Rpts. 38: 2768.
Peromyscopsylla ravalliensis Jellison and Good, 1942, U. S. Pub. Health Sery.,
Natl. Inst. Health Bul. 178: 122.

Type host-—Neotoma cinerea (Ord) (wood rat).

Type locality —Tin Cup Creek in the foothills of Bitterroot Moun-
tains southwest of Darby, Mont.

Range.—Western part of the United States and Canada.

The female terminalia of this species were figured by Wagner
(84, p. 204) in 1936.

PEROMYSCOPSYLLA scoTTI I. Fox

Peromyscopsylla scotti I. Fox, 1939, Wash. Ent. Soc. Proc. 41: 49, pl. 6, figs. 4-5.

Type host-—Peromyscus leucopus (Raf.) (white-fcoted mouse).

Type locality Dubuque, Iowa.

Range.—Kastern part of the United States.

This species has been redescribed and figured by I. Fox (25, pp. 86
and 172).

PEROMYSCOPSYLLA SELENIS (Rothschild)

Ctenopsyllus selenis Rothschild, 1906, Canad. Ent. 88: 322, fig. 43.
Peromyscopsylla selenis Jellison and Good, 1942, U. S. Pub. Health Sery., Natl.
Inst. Health Bul. 178: 123.

Cotype hosts —vVarious field mice.

Cotype localities—Various localities in British Columbia and
Alberta.

Range.—Canada.

The original description of this species is accompanied by an illus-
tration of the male terminalia.

The Genus LepropsyLia Jordan and Rothschild
(Fig. 8, G)

Leptopsylla Jordan and Rothschild, 1911, Novitates Zool. 18: 85. Type, (Pulex
musculi Dugées) =P. segnis Schonherr (by original designation).

This generic name was proposed to replace Ctenopsyllus Kolenati,
1863, not Kolenati, 1856.

Head subangulate in front; anterior margin with a series of bristles, of which
at least 2 are heavily pigmented. Genal comb consisting of 4 spines. Eyes
vestigial. Labial palpus consisting of 5 segments. Pronotum with a comb of
many long, slender spines. Antepygidial bristles present. Coxa III without a
patch or row of spinelets on the inner surface. Posterior margin of hind tibia
armed with a series of about 14 bristles, among which are 3 or 4 very long
ones. Fifth tarsal segment of each leg with 4 pairs of lateral plantar bristles
and a basal, submedian, ventral pair.

Jordan and Rothschild (48, p. 85) held that Ctenopsyllus Kolenati,
1862 [really 1863], was a homonym of Ctenopsyllus Kolenati, 1856.

THE FLEAS OF NORTH AMERICA 93

The latter name was proposed, according to the opinion of these
authors, as a more appropriate name than Ceratopsyllus Curtis for
bat fleas. This substitution of the name Leptopsylla for Ctenopsyllus
Kolenati, 1863, has not been accepted by Wagner, who has held that
Kolenati (55, p. 31) in 1856 did not really propose a new generic
name but only mentioned the name incidentally in his discussion of
the lack of appropriateness of the name Ceratopsyllus Curtis. It is
not exactly clear from the language used by Kolenati whether he
intended to propose Ctenopsyllus as a new name. In this connection
it is to be noted that in the text of his paper he used Ceratopsyllus
Curtis, while the name Ctenopsyllus appears in a footnote.

In 1932 Jordan (42, p: 291) upheld and justified the substitution
of the name Leptopsylla for Ctenopsyllus Kolenati, 1863, although
apparently admitting that Ctenopsyllus Kolenati, 1856, was proposed
conditionally. He states:

* * * A name published in this way is as valid from the date of publication
as if Kolenati had said: “I name the comb-bearing fleas Cienopsyllus.” A very
large number of names have been published conditionally. Phrases in meaning
like the following are quite frequent: “If the differences here mentioned should
turn out to be constant, the name X—ws would be appropriate.” * * * A name
published with a description or as an alternate name is valid whatever phrase-
ology is employed.

While not entirely agreeing with Jordan’s opinion, the writers
believe that the conditional language used by Kolenati is. not
sufficient to exclude this name from consideration under the rules.

Leptopsylla is represented in North America by a single species.

LEPToPsyLLA sEGNIs (Schoénherr)
(Fig. 8, G)
Pulez segnis Schénherr, 1811, Svenska Vetensk. Akad. Handl. (ser. 2)
32: 98, fig.
Pulez musculi Dugés, 1852, Ann. des. Sci. Nat. Zool. (ser. 1) 27: 160, pl. 4,
figs. 3 and 10.
Typhiopsylla mexicana Baker, 1896, Canad. Ent. 28: 85.
Leptopsylla musculi Jordan and Rothschild, 1911, Novitates Zool. 18: S85.

Type host—Mus musculus L. (house mouse).

Type locality —Sweden.

Range—Iin North America, the coastal areas of the United States
and Mexico.

This species may be immediately recognized by the character of
the genal comb (fig. 8, @). Illustrations of diagnostic structures in
both the male and female may be found in I. Fox’s work (25, p. 177).

Two additional synonyms are indicated in Wagner’s catalog (80,
p. 22).

Wagner (82) claims that samples of segnis taken from squirrels in
the Old World differ from those taken from other hosts and has
proposed for them the name of scivrobius. North American speci-
mens all appear to be typical representatives of segnis.

The Genus Nearcropsytia Rothschild
(Fig. 8, D)

Nearctopsylla Rothschild, 1915, Novitates Zool. 22: 807. Type, Ctenopsyllus
brooksi Rothschild (by original designation).

Head flattened in front. Genal comb vertical, consisting of 5 unequal,

94. MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

more or less spatulate spines. Eyes absent. Labial palpus composed of 5
segments. Pronotum with a comb of many long, slender, acuminate spines.
Anterior abdominal tergal plates, at least, with 2 or more apical spines.
Antepygidial bristles present. Coxa III with a patch of spinelets on inner
surface, Fifth tarsal segment of legs I and II with 5 pairs of lateral plantar
bristles; fifth tarsal segment of leg III with 4 pairs of lateral plantar bristles.

This genus contains four species.

NEearcTopsyLia Brooks! (Rothschild)

Ctenopsyllus brooksi Rothschild, 1904, Novitates Zool. 11: 649, pl. 15, figs. 86 and 88,
and pl. 16, fig. 89.
Nearctopsylla brooksi Rothschild, 1915, Novitates Zool. 22: 307.
Cotype hosts—Martes americana (Turt.) (marten), Mustela
longicauda Bonap. and I, cicognanii richardsonii (Bonap.)- (weasels).
otype localities—Various localities in Alberta and British Co-
lumbia.
Range.—Canada.
The original description of this species is accompanied by illus-
trations of the male head and terminalia and the female terminalia.

NEARCTOPSYLLA GENALIS GENALIS (Baker)
(Fig. 8, D)

Ctenophthalmus genalis Baker, February 1904, U. S. Natl. Mus. Proce. 27: 424.

Ctenopsyllus hygini Rothschild, September 1904, Novitates Zool, 11: 650, pl.
15, fig. 85, and pl. 16, figs. 93, 94.

Nearctopsylla hygini Rothschild, 1915, Novitates Zool. 22: 307.

Nearctopsylla genalis Jellison and Good, 1942, U. 8S. Pub. Health Serv., Natl. Inst.
Health Bul. 178: 96.

Type host—The original description indicates Geomys bursarius
as the type host, but the type slide is labeled, “On Scalops.” Hence,
it seems very likely that the common garden mole, Scalopus aquati-
cus, is the type host of this species, rather than the pocket gopher,
Geomys bursarius.

Type locality—Agricultural College, Mich.

Range.—Eastern part of the United States and Canada.

This species has been redescribed and illustrated by I. Fox (25,
DD IL, LO)

NEARCTOPSYLLA GENALIS LAURENTINA Jordan and Rothschild

Nearctopsylla hygini laurentina Jordan and Rothschild, 1923, Ectoparasites
Li Sib) figs), 319 a2:
Type host.—Mustela sp. (weasel).
Type locality—New Brunswick, Canada.
Range.—Known only from New Brunswick.
_ The original description of this species is accompanied by illustra-
tions of the male and female terminalia.

NEARCTOPSYLLA HyRTACI (Rothschild)

Ctenopsyllus hyrtaci Rothschild, 1904, Novitates Zool. 11: 652, pl. 16, figs. 92, 95.
Nearctopsylia hyrtaci Rothschild, 1915, Novitates Zool. 22: 307.

Cotype hosts—Mustela vison energumenos (Bangs) (mink) and
Sorex obscurus Merr. (shrew).

Cotype localities —Cariboo and another unnamed locality in British
Columbia.

THE FLEAS OF NORTH AMERICA 95

Range.—Known only from British Columbia.
The original description of this species is accompanied by illus-
trations of the male and female terminalia.

NEARCTOPSYLLA JORDANI Hubbard

Nearctopsylla jordani Hubbard, 1940, Pacific Univ. Bul. 37 (1): [5], 9 figs.

Type host—Scapanus townsendii (Bach.) (Townsend’s mole).

Type locality —Forest Grove, Oreg.

Range—Probably that of the Townsend mole, which is west of the
Cascade Mountains in Oregon and Washington.” (Hubbard, in the
original description.)

The original description of this species is accompanied by illustra-
tions of the head and terminalia of the male and female.

The Genus CorypsyLLaA C. Fox
(Fig. 8, A)

Corypsylla C. Fox, 1908, Ent. News 19: 452. Monotypic. Type, Corypsylla
ornata C. Fox.

Head rounded in front. Genal comb consisting of 6 spatulate spines, unequal
in length. Eyes absent. Labial palpus consisting of 5 segments, the terminal
segment tapering distally. Pronotum with a comb of spines, exhibiting marked
sexual dimorphism, several of the most ventral spines in the female being
pale and fused together for most of their length. Mesopleuron not divided
by a pleural ridge. Metanotum fused with metepisternum. Each abdominal
tergal plate, exeept VII, with a heavy, transverse incrassation which bears
several heavily pigmented spines. Antepygidial bristles present. Coxa III
with a patch of spinelets on inner surface. Fifth tarsal segment of each leg
with 4 pairs of lateral plantar bristles.

This genus includes but two species, both being Nearctic.

CoryPsYLLA JORDANI Hubbard

Corypsylla jordani Hubbard, 1940, Pacific Univ. Bul. 37 (1): [9], 7 figs.

Type host—Nest of Peromyscus maniculatus rubidus (Osg.)
(white-footed mouse).

Type locality Gaston, Oreg.

The original description of this species is accompanied by illustra-
tions of the male and female heads and terminalia which show the
points in which this species differs from Corypsylla ornata C. Fox,
to which it is closely allied.

CoryPsYLLA ORNATA C. Fox
(Fig. 8, A)
Corypsylla ornatus C. Fox, 1908, Ent. News 19: 452, 2 figs.
Type host—‘Scapanus californicus” (mole).
Type locality —San Francisco, Calif.
fange.—Western part of the United States.
The original description of this species is accompanied by illustra-

tions of the male head and terminalia. Hubbard (30) has published
further illustrations, which include a figure of the female terminalia.

96 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

The Genus CorypsyLttowes Hubbard

Corypsylloides Hubbard, 1940, Pacific Univ. Bul. 37 (2): 7. Monotypiec. Type,
Corypsylla kohlsi Hubbard.

Front reduced; frontal tubercle and notch absent. Gena much enlarged;
genal comb obliquely vertical, consisting of 6 spines of about the same length.
Eyes absent. Labial palpus 5-segmented, the terminal segment tapering dis-
tally. Pronotum rounded cephalad, well separated from head, bearing on posterior
margin a comb consisting of spines of various sizes. Mesopleuron not divided
by a vertical rodlike sclerotization into a mesepisternum and a mesepimeron.
Abdominal tergal plates armed with prominent apical spines but without
heavy transverse incrassations. Coxa III with a patch of spinelets on inner
surface. Last segment of each tarsus armed with 4 pairs of lateral plantar
bristles.

This genus is most closely allied to Corypsylla C. Fox, from which
it may be distinguished by the absence of heavy transverse incrassa-
tions on the abdominal tergal plates and by the character of the
genal comb. It is separated from Nearctopsylla Rothschild, which
it superficially resembles, by the structure of the thorax. The genus

is represented in North America by one described species.

CorypsyLLowEs KoHLs! (Hubbard)

Corypsylla kohlsi Hubbard, 1940, Pacific Univ. Bul. 87 (1): [10], 7 figs.
Corypsylloides spinata I. Fox, 1940, Wash. Acad. Sci. Jour. 30: 378, figs. 1, 2.

Type host—Sorex obscurus batrdi Merry. (Baird’s shrew)..

Type locality.—Cannon Beach, Oreg.

Range.—Oregon.

This species was well illustrated in the original description. The
movable finger, however, is shown to have put two setae on the
posterior margin. The specimens upon which I. Fox based the
name Corypsylloides spinata indicate three larger setae and two
minute ones, as is shown in his illustration.

/The Family IscunNopsyttipaE Wahlgren

Head divided into anterior and posterior parts by fronto-epicranial groove.
Gena neither greatly enlarged nor divided by a suture. Forehead without
helmet, usually elongated, and always with 2 or 3 ventral flaps on each side
adjacent to fronto-genal angle. Eyes reduced, vestigial, or absent. Cephalic
combs represented by the 2 or 8 flaps on each side of head. Postantennal
region with or witheut dorsal incrassations. Thorax not greatl- reduced;
thoracic terga taken together longer than abdominal tergum I. ‘Typical ab-
dominal terga each with more than 1 transverse row of setae. Antepygidial
bristles present or absent. Abdominal apical spines frequently present and
may form 1 or more combs. True or false combs of long spines may be present
on abdomen. Females without a reduced number of abdominal spiracles and,
when gravid, abdomen not greatly distended.

The family Ischnopsyllidae was established by Wahlgren (87, p.
89) in 1907. The following synonyms are recorded: Ceratopsyllidae ©
Baker (4, p. 157) and Ischnopsyllinae Wagner (SO, p. 29). Ceratop-
syllidae is based upon Ceratopsyllus Kolenati, 1857, which is a syno-
nym of Jschnopsyllus Westwood, 1883.

Wagner (S86, p. 75) has divided this family into two subfamilies.
The characters are: ‘The presence or absence of a wide clear area
on the frons, the presence or absence of antepygidial bristles, and
the presence or absence of a comb of thickened bristles on abdominal

THE FLEAS OF NORTH AMERICA 97

plate VII. The members of his subfamily Ischnopsyllinae are said
to possess the wide, clear area on the head and the antepygidial
bristles, while the members of the subfamily Nycteridopsyllinae are
characterized by the absence of the wide, clear area on the head and
by the presence of a comb of bristles on tergal plate VII which
takes the place of the absent antepygidial bristles. Accordingly, the
genera Myodopsylla, Sternopsylla, and Rhinolophopsylla are refer-
red to the Ischnopsyllinae by Wagner; but of these genera only
Myodopsylia has a distinct, clear area on the frons. Hence this
character does not apply generally and is not satisfactory as a basis
for the establishment of a subfamily. The other characters used,
i. e., the presence or absence of antepygidial bristles and the presence
or absence of a comb of spines, while of great generic value, are not
here regarded sufficiently important, when used by themselves, to dis-
tinguish subfamilies. In our fauna only the genus /ptescopsylla
would go into the subfamily Nycteridopsyllinae. No provision was
made by Wagner for the genus Pfilopsylla, which he believes is sepa-
rated from both subfamilies and cannot be placed until more is
known about it.

The present writers are of the opinion that the Ischnopsyllidae,
as the family is now constituted, does not break into subfamilies.
However, the discovery of more genera and species, which will un-
doubtedly come with time, may lead to a revision of this belief.

Key To THE NortrH AMERICAN GENERA OF ISCHNOPSYLLIDAE

1. Maxillae either acuminate or narrowly rounded apically___________-~-
Maxillae truncate or nearly truncate apically______________________

2. Antepygidial bristles present; abdominal tergal plate VII without
a comb of thickened bristles____Sternopsylia Jordan and Rothschild.
Antepygidial bristles absent; abdominal tergal plate VII. with a
ts) OF Ale kened .brintien. > 2 os Eptescopsylla I. Fox.

3. Abdomen without combs, but sometimes with rows of minute teeth
Rhinolophopsylla Oudemans.

Abdomen with false combs composed of enlarged setae in some of the
iraneverse ‘TOwse 2 ee Myodopsylla Jordan and Rothschild.

W bro

The Genus SrernopsyLia Jordan and Rothschild
(Figs. 8, F, and 11, D)

Sternopsylla Jordan and Rothschild, 1921, Ectoparasites 1: 158. Type, Isch-
nopsyllus texranus C. Fox (by original designation).

Aptilepsylla Ewing, 1940, Wash. Biol. Soc. Proc. 53: 36. Monotypic. Type,
Aptilopsylla carilsbadensis Ewing.

Head without dorsal incrassations. Frontal tubercle present or absent.
Ventral flaps long, slender, curved, tapering to the acuminate apex, with pos-
terior flap longer than anterior. Genal process acuminate. Eye vestiges large.
Ocular bristle present. Maxilla acuminate. Pronotum with a true comb.
Abdominal terga without combs. Antepygidial bristles present. Tarsi long,
first segment of mid and hind tarsi with numerous small, hairlike setae, par-
ticularly in male. Fifth tarsal segment of each leg with 5 pairs of plantar
bristles, of which the first pair is ventral, the other 4 lateral.

This genus is represented in North America by two species. Both
were described from females, but recently the males have been de-
scribed by Guimaraes (27), and the writers have received a large
consignment of Sternopsylla, including males, from G. M. Kohls,
taken in Carlsbad Cavern, N. Mex.

467459 —43——7

98 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

The genus is remarkable in that the females of the two species and
their varieties appear to be identical, except that in one situation,
Carlsbad Cavern, N. Mex., some of the females possess a very distinct
frontal tubercle and the others not so much as a vestige of such a
tubercle. The males corresponding to the two kinds of females in
this cavern are so nearly alike that they cannot be separated except
for the presence or absence of the frontal tubercle.

While the females from other localities appear to be alike, the
males show slight differences in the shape of the posterior process
of the clasper, in the shape of the movable finger, and especially in
the chaetotaxy of the latter.

Guimaraes (27) gives a drawing of the clasper of the male of what
he identifies as Sternopsylla texana (C. Fox), based on material from
Mexico. According to this drawing the Mexican male is different
from all those known to the writers from the United States in the
shape of the posterior process of the clasper, which is longer in pro-
portion to its width than in forms occurring in the United States.
Also the chaetotaxy and shape of the movable finger are different.

When the known males of Sternopsylla from the United States are
compared, they are observed to fall into three groups according to
the following differences in the claspers:

1. Those from Carlsbad Cave, N. Mex., in which the movable finger is
about as broad as long and the anterior margin is very broadly incurved, the
two lower, posterior, marginal, spinelike setae of the movable finger are
situated at a distance from each other, and the seta adjacent to the third
posterior, marginal, spinelike seta is large and longer than the third seta itself.

2. The one male from Georgia, in which the movable finger is distinctly
longer than broad and the third spinelike seta is very slender and unaccom-
panied externally by another seta, and in which there are two external discal
setae above the middle of the finger.

3. The one from Florida, in which the movable finger is about as broad as
long, with the anterior margin deeply incurved or notched below its middle,
the two lower, posterior, marginal, spinelike setae are situated approximate
to each other, and the seta adjacent to the third posterior, marginal, spinelike
seta is flagelliform and shorter than the third seta itself.

These differences in the genitalia of the males from the United
States together with the differences in the presence or absence of the
frontal tubercle suggest that we have at least four varieties. But
four varieties of Sternopsylla terana or S. carlsbadensis? In view of
the fact that Guimaraes has figured a male (rightly or wrongly
associated with the proper female) of S. texana, they are here re-
garded as varieties of S. carlsbadensis, but are not named.

STERNOPSYLLA CARLSBADENSIS (Ewing), new combination
(Fig. 8, F)

Aptilopsylla carlsbadensis Ewing, 1940, Wash. Biol. Soc. Proc. 53: 36.

Type host—Unknown. Type labeled “on bat guano.”

Type locality.—Carlsbad, N. Mex.

Range.—The typical form (with frontal tubercle) known only from
Carlsbad Cavern, N. Mex.

As already explained in the discussion following the formal descrip-

tion of the genus Sternopsylla, there may be as many as four varieties
of this species.

THE FLEAS OF NORTH AMERICA 99

STERNOPSYLLA TEXANA (C. Fox)
(Fig. 11, D)

Ischnopsyllus texranus C. Fox, 1914, U. S. Pub. Health Serv. Hyg. Lab. Bul. 97:
16, pl. 5, figs. 6-8.

Sternopsylla texana Jordan and Rothschild, 1921, Ectoparasites 1: 158.

Type host—Tadarida mexicana (Sauss.) (bat).

Type locality.—Pecos, Tex.

Range.—Southern Texas and (?) northern Mexico.

For a discussion of the taxonomy of this species read the discussion
following the formal description of Sternopsylla. This species was
redescribed and figured by I. Fox (25, pp. 108, 191) in 1940.

The Genus Eprescopsy.ua I. Fox
(Fig. 11, 4)

Eptescopsylla I. Fox, 1940, Fleas of Eastern United States, p. 107. Monotypic.
Type, Nycteridopsylla chapini Jordan.

Head without dorsal incrassations. Frontal tubercle absent. Ventral flaps
long, slender, nearly straight, tapering to the acuminate apex, with posterior
flap much longer than anterior. Genal process rounded posteriorly. Eye ves-
tiges large. Ocular bristle absent. Mavxilla acuminate. Pronotum with a true
comb. Metanotum and anterior abdominal tergal plates each armed with 2 or 3
dorsal teeth on a side. Abdominal tergal plate VII with a false comb. Ante-
pygidial bristles absent. Fifth tarsal segment of each leg with 5 pairs of plantar
bristles, of which the first pair is ventral, the other 4 lateral.

This genus may be separated from Vycteridopsylla Oudemans, to
which it is allied, by the absence of combs on the anterior abdominal
tergal plates Ito III. The genus Vycteridopsylla does not occur in
North America. L'ptescopsylla is represented by two species. .

Although the generic host name upon which the name E'ptescop-
sylla was based has the letter z in it, this letter was omitted in the for-
mation of the latter for the sake of euphony.

EPTESCOPSYLLA CHAPINI (Jordan)
(Fig. 11, 4)

Nycteridopsylla chapini Jordan, 1929, Novitates Zool. 35:39, pl. 2, figs. 30, 31.
Eptescopsylla chapini I. Fox, 1940, Fleas of Eastern United States, p. 107, pl. 30.

Type host —Eptesicus fuscus (Beauv.) (big brown bat).

Type locality—Glen Echo, Md.

Range.—Probably eastern part of the United States.

Sexual dimorphism is shown in the chaetotaxy of the head, the
female being armed with heavily pigmented, spinelike bristles near
the antennal groove, while such bristles are absent in the male.

EPTESCOPSYLLA VANCOUVERENSIS (Wagner)

Sa oes vancouverensis Wagner, 1936, Ztschr. f. Parasitenk. 8: 658,
gs. 7-8.

Type host.—Lasionycteris noctiwagans (LeC.) (silver-haired bat).

Type locality—Vancouver, British Columbia.

Range—Known only from British Columbia.

In the description of his species, Wagner states that Eptescopsylla
vancouverensis is distinguished from EZ’. chapini by the possession of

100 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

a much narrower head, which is acuminate anteriorly, and by the
structure of the movable finger of the male. Paratype male and female
specimens of this species have been studied through the courtesy of
G. J. Spencer and G. P. Holland.

The Genus RHINOLOPHOPSYLLA Oudemans

Rhinolophopsylla Oudemans, 1910, Ent. Ber. 3: 3. Monotypic. Type, Typhlopsylla
unipectinata Taschenberg.

Head without dorsal inecrassations. Frontal tubercle absent. Ventral flaps
short and broad, the posterior longer than the anterior. Genal process acumi-
nate. Eye vestiges indistinct. Ocular bristle present. Maxilla truncate. Pro-
notum with a true comb. Abdomen without combs and without dorsal apical
spines. Antepygidial bristles present. Fifth tarsal segment of each leg with 5
pairs of plantar bristles, of which the first pair is ventral, the other 4 lateral.

This genus is represented in North America by but one species.

RHINOLOPHOPSYLLA PALPOsA (Rothschild)

Ceratopsylla palposus Rothschild, 1904, Novitates Zool. 11: 652.
Rhinolophopsylla palposus Wagner, 1936, Canad. Ent. 68: 206.

Type host—‘Brown bat.”

Type locality —Cowicham Demeans, British Columbia.
Range.—Known only from British Columbia.

The male of this species has not yet been described.

The Genus Myopopsyiia Jordan and Rothschild
(Fig. 11, B, 0, B)

* Myodopsylla Jordan and Rothschild, 1911, Novitates Zool. 18:88. Monotypic.
Type, Ceratopsylla insignis Rothschild.

Frons a wide, clear area without sclerotization. Frontal tubercle absent.
Ventral flaps short and broad, the posterior slightly longer and much wider
than the anterior. Genal process acuminate. Hye vestiges large. Ocular bristle
present. Maxillae truncate. Pronotum with a true comb. Metanotum and
abdominal tergal plates I to VII each with a false comb formed by thickening
of dorsal bristles. Antepygidial bristles present. Fifth tarsal segment of each
leg with 5 pairs of plantar bristles, of which the first pair is ventral, the other 4
lateral.

This genus is represented in North America by four species.

MyopopsyLia coLutnst Kohls
(Fig. 11, B, £)
Myodopsylla collinsi Kohls, 1937, Jour. Parasitol. 23: 300, figs. 1, 2.

Type host.—‘Bats.” ,

Type locality—Madera Canyon, Santa Rita Mountains, Santa
Cruz County, Ariz.

Range.—Known only from Arizona.

The original description of this species is accompanied by four fig-
ures of the male and female terminalia. The species may be adele
distinguished by the structure of the male terminalia (fig. 11, B).

THE FLEAS OF NORTH AMERICA 101

MyopopsytLA DIASI Costa Lima

Myodopsylla diasi Costa Lima, 1938, Rev. Med. Cirurg. do Brasil 46 (2): 182,
figs.

Type host—Myotis velifer (Allen) (bat). _

Type locality—S. Cristobal, Ecatepec, Mexico.

Range.—Known only from Mexico.

This species is most closely allied to Myodopsylla collinsi Kohls,
with which it may possibly be identical. Since no specimens are at
hand for comparison with those of collinsi, it is best to regard it as
a separate species for the present.

MyopopsyLia GENTILIs Jordan and Rothschild

Myodopsylla gentilis Jordan and Rothschild, 1921, Ectoparasites 1: 152, fig. 131.

Type host—‘Bat.”

Type locality—Okanagan Landing, British Columbia.

Range.—Western part of the United States and Canada,

The male terminalia are well illustrated in the original description
of this species. The seventh and eighth abdominal segments of the
female, however, are indicated as being the same as in Myodopsylla
insignis.

Myopopsy1a Insients (Rothschild)
(Fig. 11, C)

Ceratopsylla insignis Rothschild, 1903, Novitates Zool. 10: 319, pl. 9, figs. 8-12.
Ceratopsyllus crosbyi Baker, 1905, U. S. Natl. Mus. Proc. 29: 187. (New
synonymy.)

Myodopsylla subulata Chapin, 1919, Brooklyn Ent. Soc. Bul. 14: 55.

Type host.—M yotis lucifugus (Lec.) (little brown bat).

Type locality —Waterloo, Ontario, Canada.

Range.—Eastern part of the United States and Canada.

Through the kindness of Robert Matheson, the writers have been
able to examine a single female specimen labeled “Jschnopsyllus
crosbyi Baker,” belonging to Cornell University. This specimen ap-
peared to be the type of Baker’s crosbyi. Our examination of it
revealed that crosbyi was identical with M. insignis (Rothschild).

Jordan and Rothschild (51, p. 161), in 1921, placed Myodopsylla
subulata Chapin as a synonym of J. insignis (Rothschild), stating:
“An incorrectness in our original figure of the exopodite of MW. in-
signis has unfortunately been the cause of the creation of a synonym.”
Pas species has been redescribed and figured by I. Fox (24, pp. 105,
187).

The Family Purtcrwae Stephens

Head not divided by a fronto-epicranial groove. Gena not greatly enlarged
and extended dorsally in front of antennal groove, and not divided by a
suture. Forehead without helmet, not elongated and never provided with ven-
tral flaps. Cephalic combs present or absent. Thorax not greatly reduced;
thoracic terga taken together longer than tergum I of abdomen. Typical ab-
dominal terga each wiih a single transverse row of setae. Abdominal combs
absent. Antepygidial bristles present. Females without a reduced number of
abdominal spiracles, and, when gravid, abdomen seldom greatly distended.

102 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

The family Pulicidae contains three subfamilies, as follows: Spilop-
syllinae, Chimaeropsyllinae, and Pulicinae.

The family name Pulicidae was first used in 1829, by Stephens
(71, p. 328), who applied it to all the fleas. Oudemans (49, p. 157),
in 1909, attributed the family to Taschenberg, and Wagner (80, p. 4)
did likewise. Dalla Torre (10, p. 22) attributed the family to Oude-
mans. In the restricted sense In which the name Pulicidae is here
used, it has no synonyms.

The Subfamily SPILOPSYLLINAE Oudemans

Fronto-epicranial suture poorly developed or absent. Eyes usually well de-
veloped and heavily pigmented but may be absent. Front not greatly reduced,
with rounded or angulate margin; frontal tubercle present or absent. Genal
comb present or absent. Pronotal comb present or absent. Mesepisternum sep-
arated from mesepimeron by an internal, vertical ridge. Episternum III usually
divided into upper and lower parts. Hind coxa either with or without a row
or patch of spinelets on inner surface.

Included in this subfamily are 25 genera, of which 8 are represented
in the North American fauna.

The subfamily Spilopsyllinae was established by Oudemans (59, p.
156), in 1909, for the single genus Spilopsyllus Baker. Dalla Torre
(10, p. 7), in 1924, added three more. The family Pulicidae and the
subfamily Spilopsyllinae are closely related to the family Dolichop-
syllidae through the genus Megarthroglossus Jordan and Rothschild.
There has never been any agreement among workers on the fleas as to
just where the family limits should be placed.

Key to NortH AMERICAN GENERA OF SPILOPSYLLINAE

1. Coxa III with a row or patch of spinelets on inner surface__-______~- 2
Coxa III without a row or patch of spinelets on inner surface____-___ 6
2. | Pronotal’-comb wabsent220 216 oe ese ee eee ph Rel See 3
Pronotal: :comb:;; present..= 2 och a nee ae +
3. Segment V of each tarsus with only 3 pairs of lateral plantar bristles.
INorthsAmerican: 22 tas Actenopsylla Jordan and Rothschild.

Segment V of each tarsus with 4 pairs of lateral plantar bristles;
episternum III divided into upper and lower parts; antennal club

not segmented on anterior surface____________ Xenopsylla Glinkiewicz.
4, Genal comb absent; club of antenna subcapitate_____ Hoplopsyllus Baker.
Genal ‘comb ‘present:..=2 5 3 ease. See Ue) ee eee ein eee 5

5. Genal comb about horizontal and extending forward to fronto-genal
angle of head; front not angulate
Ctenocephalides Stiles and Collins. _
Genal comb more nearly vertical than horizontal and composed of spines
that are rounded at apex; front somewhat angulate dorsally; labial

G:°>Pronotal’ comb) present 2-22 A ee ee ee eee 7
Pronotal comb absent; frontal tubercle present; eyes absent
Anomiopsyllus Baker.
7. Labial palpus 4-segmented ; femora without lateral setae
Callistopsyllus Jordan and Rothschild.
Labial palpus 5-segmented, the fifth segment equal to, or longer than,
II, III, and IV taken eogether; lobe on posterior margin of coxa III
UCU LCs hays Oe eae Megarthroglossus Jordan and Rothschild.

THE FLEAS OF NORTH AMERICA 103

The Genus AcTENopsyLLA Jordan and Rothschild

Actenopsylla Jordan and Rothschild, 1923, Ectoparasites 1: 309. Monotypic.
Type, Actenopsylla suavis Jordan and Rothschild.

This genus includes but a single rare species, representatives of
which are not available for study, hence the original description by
Jordan and Rothschild is here given in full.

Near Spilopsyllus Baker (1905). Without ctenidia. Mandibles normal, not
enlarged.

Frons strongly rounded, without tubercle or angle, with two long bristles, one
in front of the eye, the other near the ventral margin above the maxillary palpus;
internal projection above maxilla small. Occiput with many small bristles
above antennal groove. Club of antenna as in Spilopsyllus segmented all round,
longer in ¢ than in Q, bristles on upperside of segment II. long. Labial palpus
with four segments.

Pro- and mesothoracic tergites and abdominal tergites II. to VII. with one row
of bristles. Metepimerum with a bristle above stigma, the distance of the stigma
from the upper margin of the epimerum being much greater than in Spilopsylius.
Stigmata III. to LX. of equal size, not so large as in Spilopsyllus, but the diameter
of a stigma more than twice the diameter of the groove of a large bristle of the
posterior row of the respective segment; cavity of last stigma very large.
Hindcoxa on inner side with a row of (about) six to eight small spines; mid-
and hindfemora with a subventral row of bristles on the inside; segment I. of
midtarsus as long as II, V. of all the tarsi with only 3 pairs of plantar
bristles.

The authors of this genus place it nearest Spilopsyllus Baker. It
is also closely related to Parapulex Wagner.

ACTENOPSYLLA SUAVIs Jordan and Rothschild

Actenopsylla suavis Jordan and Rothschild, 1923, Ectoparasites 1: 309, figs.
312, 313.

Type host—“Nest of Ptychoramphus aleuticus” (Cassin’s auklet).

Type locality —Coronado Island, Mexico.

cange.—Known only from Mexico.

The original description of this species was based on one male and
seven females, and is accompanied by a drawing of the genitalia of
the male and the tip of the abdomen of the female.

‘1 The Genus XENopsytia Glinkiewicz
(Fig. 12, D and G)

Xenopsylla Glinkiewicz, 1907, Akad. der Wiss. Wien, Math.-Nat. K]. Sitzber. 116:
385. Monotypic. Type (Xenopsylla pachyuromydis Glinkiewicz) =X. cheopis
(Rothschild).

Loemopsylla Jordan and Rothschild, 1908, Parasitology 1:15. Type, Pulex cheopis
Rothschild (by original designation).

Front short, high, with anterior margin broadly rounded. Frontal tubercle
absent. Genal comb absent. Eye large, unreduced, black. Labial palpus 4-seg-
mented, both anterior and posterior margins well sclerotized. Antennal club
short; not segmented on anterior surface. Episternum III divided into upper
and lower parts by a deep suture. Pronotal comb absent; pronotum with a
Single transverse row of setae. Hind coxa with a row or patch of spinelets
on inner surface. Femora almost devoid of lateral setae. Segment V of each
tarsus with 4 pairs of lateral plantar bristles. Abdomen with long setae, but
without apical spines. Sternal plate VIII of male large, broad, unmodified.
Sternal plate IX of male with an anterior, whiplike apophysis, a pair of poorly
sclerotized, inner, vertical processes, and a pair of simple, more or less clavate,
posterior processes,

104 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

Xenopsylla includes a large number of closely related species, some
of which should probably be considered more appropriately as varie-
ties. Originally it apparently was confined to Africa, or to Africa
and western Asia. Through commercial activities some of the species
have been introduced into Europe and the Western Hemisphere as
well as into various parts of Asia.

XENOPSYLLA CHEOPIS (Rothschild)
(Fig. 12, D and G@)

Pulex ‘cheopis Rothschild, 1908, Ent. Monthly Mag. (2) 14: 85, pl. 1, figs. 3, 9;
pl. 2, figs. 12, 19.

Xenopsylla cheopis Rothschild, 1909, Novitates Zool. 16: 132.

Loemopsylla cheopis Jordan and Rothschild, 1908, Parasitology 1: 42.

Cotype hosts—Seven hosts were indicated by Rothschild in his
original description.

Type locality—Shendi, Egypt.

Range.—Although this species is not cosmopolitan in its distribution,
it has been very widely disseminated through the introduction of
its favored hosts into many countries. However, since much of its
life cycle is spent detached from the warm bodies of the hosts, this
more. or less tropical species apparently has not been able to follow
them into all the areas they have invaded.

The distribution of Xenopsylla cheopis, or the oriental rat flea, in
the United States is interesting. For many years it was reported
only from the larger port cities, where it frequently made up a con-
siderable percentage in the total rat flea population. In 1925, how-
ever, F. N. Wallace(S8) reported finding cheopzs on rats in Indian-
apolis, Ind., but noted that no infestation of houses was observed.
About this time W. A. Riley identified a specimen from St. Paul,
Minn., taken from a rat. Then, in 1934, Roudabush and Becker (68)
reported the finding of many specimens of cheopis on rats shot on the
dumping ground of the city of Ames in Iowa. They called attention
to the statement of C. Fox (23, p. 133) in 1925 that the oriental rat
flea was not known to occur in the interior of the United States. This
infestation at Ames has continued since the observations of Rouda-
bush and Becker and constitutes, as far as known, a permanent in-
festation. In 1939 Roudabush (67) also recorded the flea from Des
Moines, Iowa. Owen (60), in 1936, reported an infestation of a dairy
barn at the University of Minnesota Farm, St. Paul, Minn., and in
1938 the present writers, Ewing and I. Fox (19), added new records
from Illinois and Ohio. Trembley and Bishopp (76) reported the
species from Nashville, Tenn. The specimens came from rats
captured in a grain store in the city near the Cumberland River in
March 1938. These authorities have reviewed their United States
records, as well as those from the literature, and find that XYenopsylla
cheopis has now been reported from 19 States and the District of Co-
lumbia. Of these 19 States, 13 are coastal with ports for foreign
commerce. The scattered records of this flea from the interior States
are of recent date and indicate the establishment of “islands” of in-
festation, in a region the climate of which might be thought adverse
to their existence.

Xenopsylla cheopis has been much studied, but only a few refer-
ences to the literature dealing with it can be given here. Bacot (7), in

THE FLEAS OF NORTH AMERICA 105

1914, in a lengthy paper reports some exhaustive studies on the bio-
nomics of this and a few other species. He found that cold (40° F.)
was fatal to the eggs of cheopis, that conditions of drought did not
affect egg production, but that low temperatures either checked or
prevented oviposition. Bacot found that the larvae of cheopis were
able to live from several days to over a month without food and that
the time consumed by the active larval period ranged from 12 to 84
days. The duration of the cocoon period was found to range from 7 to
182 days. Adults kept in a box and fed on their natural host lived for
100 days.

The oriental rat flea is the chief agent in the transmission of bu-
bonic plague, a disease that formerly took a frightful toll of life in
the Old World. According to Waterston (90) over seven million
deaths resulted from this disease in India between 1896 and 1911.
This flea also is implicated in the transmission of certain types of
endemic typhus.

Two species, Xenopsylla brasiliensis and X. astia, as well as some
others, are frequently confused with XY. cheopis. As pointed out by
Rothschild (65, p. 84) in 1914, brasiliensis differs from the other two
by having the antepygidial bristle situated on “a long pedestal” in the
male sex; while asta differs from brasiliensis and cheopis by having,
in the male, a somewhat curved and ribbonlike posterior process or
lobe of the ninth sternal plate instead of a club-shaped one.

Wagner (80, p. 35), in his catalog, indicates four synonyms of this
species.

The Genus Hopiopsytius Baker
(Fig. 12, C)

Hoplosyllus Baker, 1905, U. S. Natl. Mus. Proc. 29: 128. Type, Pulex anomalus
Baker (by original designation).

Anterior margin of front rounded. Genal comb absent. Frontal tubercle ab-
sent. Eyes well developed, subcircular, black. Labial palpus 4segmented, an-
terior margin thickened and well sclerotized, posterior margin weak, but slightly
sclerotized. Antennal club capitate or strongly clavate. Episternum III di-
vided into upper and lower parts by a deep horizontal suture. Pronotal comb
present, composed of more than 8 black, apically acute spines, and not extend-
ing ventrally on either side to lower margin of pronotum. Hind coxa witha
row or paich of spinelets on inner surface. Segment V of each tarsus with 4
pairs of lateral plantar bristles. Abdomen without apical spines.

Hoplopsyllus is confined entirely to the Northern Hemisphere, and
largely to the Nearctic Region. Two subgenera are recognized.

The Subgenus HOPLOPSYLLUS Baker

Antennal club capitate, segments anchylosed anteriorly. Antennal groove
closed posteriorly in both sexes. Clasper of male with a rather small but
broad anterior process, but without posterior fingerlike process; movable finger
broad, crescentic.

Certain kinds of squirrels are natural hosts.

_The type species of the genus Hoplopsyllus Baker has been recog-
nized, since its description, as a species apart from its congeners. For
this reason the genus was recently divided into two subgenera. The
chief differences between these two subgenera lie in the male genital
armature.

106 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

Hor.opsytitus (HopiopsyLtius) ANomMALUus Baker
(Fig. 12, C)

Pulex anomalus Baker, 1904, U. S. Natl. Mus. Proc. 27: 381, pl. 10, figs. 1-6.
Hoplopsyllus anomalus Baker, 1905, U. S. Natl. Mus. Proce. 29: 128.

Type host—*“A large gray-brown spermophile.”

Type locality—Arboles, Colo.

Range—Western part of the United States.

The original description of anomalus, based on a single male and
female, is accompanied by six detail figures.

The Subgenus EUHOPLOPSYLLUS Ewing

Euhoplopsyllus Ewing, 1940, Wash. Biol. Soc. Proc. 53: 37. Type, Holopsyllus
afinis Baker (by original designation).

Antennal club clavate to strongly clavate, but not capitate, segments separated
anteriorly by grooves. Antennal groove partly or entirely open in the male,
closed in the female. Clasper of male with a broad, setigerous, anterior process
and a slender, fingerlike, posterior process (immovable finger), armed apically
with a stout, pigmented, spinelike seta; movable finger subclavate, almost
straight.

Rabbits and hares are the natural hosts.

This subgenus includes three species listed in this paper as valid,
one of which (glacialis Taschenberg) is circumpolar, the other two
being North American only. ‘The species show a high degree of
individual variation.

Hoptopsyititus (EuHopLopsyLLus) AFFINIs (Baker)

Pulex affinis Baker, 1904, U. S. Natl. Mus. Proce. 27: 382.
Hoplopsyllus affinis Baker, 1905, U. S. Natl. Mus. Proe. 29: 139.
Type host—Lepus sp. (hare).
Type locality—Near Grand Canyon, Ariz.
Range.—Western part of the United States eastward to Iowa.
This species has recently been figured by I. Fox (25, p. 137). Itisa
common flea upon different species of rabbits throughout its range.

HoptopsyLius (EvuHopLopsyiius) Exoticus Jordan and Rothschild

Hoplopsyllus exoticus Jordan and Rothschild, 1923, Ectoparasites 1: 311, fig.
314.

Type host.—Felis sp.

Type locality—Boquete, Panama.

Range—Known only from Panama.

This species was described from a single female. The male is
unknown. According to the original description it is similar to lynx
Baker, but sternal plate VII of female is “more widely excised be-
neath than in H. /ynz” and the stylet is “much longer than in H. lyna.”

HopiopsyLius (EvHOPLoPSYLLUS) GLACIALIS GLACIALIs (Taschenberg)
4

Pulex glacialis Taschenberg, 1880, Die Fléhe, p. 76, pl. 3, figs. 17-17a.
Hoplopsyllus glacialis Baker, 1905, U. S. Natl. Mus. Proce. 29: 180.

Type host.—Lepus arcticus Ross (polar hare).
Type locality—‘Nordpole.”

THE FLEAS OF NORTH AMERICA 107

Range——Northern North America, including Greenland.
Hoplopsyllus glacialis glacialis and H. glacialis lynx are very close
to each other, and according to Kohls (54, p. 2019)—

the only difference between the two appears to be in size, glacialis lynz being
notably smaller. However, on the grounds of host relationships and geographical
distribution, it is probably desirable for the present, at least, to retain the
subspecific distinction.

Hoptopsytitus (EUHOPLOPSYLLUS) GLACIALIS LyNx (Baker)

Puler lyna Baker, 1904, U. S. Natl. Mus. Proc. 27: 383, pl. 10, figs. 7-11, and
pl AL hes. 1
Hoplopsyllus glacialis lynx Wagner, 1986, Canad. Ent. 68: 194.
Hoplopsyllus lynz I. Fox, 1940, Fleas of Eastern United States, p. 15, pl. 4,
figs. 11, 13.
Type host—Lynzx canadensis Kerr (Canada lynx).
Type locality—Moscow, Idaho.
Range.—United States, Canada, and Alaska.
This variety is very close to Hoplopsyllus glacialis glacialis, but it
is considerably smaller than the latter. The type host undoubtedly
is not a natural host of H. g. lynx. Natural hosts are rabbits.

HoptopsyLius (EuHoPLOPSYLLUS) GLACIALIS FOxI Ewing

Hoplopsyllus fori Ewing, 1924, Parasitology 16: 350.
Hoplopsyllus powersi C. Fox, 1926, Pan-Pacific Ent. 2: 184, figs. 8-9.
Hoplopsyllus minutus C. Fox, 1926, Pan-Pacific Ent. 2: 185, figs. 6, 10.

Type host.—Sylvilagus bachmani (Waterh.) (cottontail rabbit).

Type locality—sSan Francisco, Calif.

Range.—Known only from California.

Hoplopsyllus glacialis foxi is distinguished from the other two
varieties of H. glacialis in having the posterior process of the clasper of
the male almost twice as long as the anterior process instead of being
only about one and a third times as long. The synonymy here given
under foxi is that of Kohls (54, p. 2020). Hoplopsyllus minutus
C. Fox appears to have been based on a dwarfed specimen of foaz.

The Genus CTENOCEPHALIDES Stiles and Collins
(Fig. 12, F)

Ctenocephalus Kolenati, 1859 (not Hawle and Corda, 1847), Fauna des Altvaters,
Briinn, p. 66. Type, Ctenocephalus novemdentatus Kolenati (by subsequent
designation).

Ctenocephalides Stiles and Collins, 1930, U. S. Pub. Health Serv. Rpts. 45: 1808.
Type, Puler canis Curtis (by original designation).

This generic name was proposed to take the place of Ctenocephalus
Kolenati, which is preoccupied by Ctenocephalus Hawle and Corda,
1847.

Front with anterior margin rounded. Frontal tubercle absent. Genal lobe
not extending downward so as to produce a ventral marginal angle. Genal
comb present. about horizontal and extending anteriorly along ventral margin
of gena to frontogenal angle, spines of genal comb heavily pigmented and
sharply pointed apically. Upper margin of antennal fossa in male with patch
of microsetae. Eyes large. black. Labial palpus 5-segmented; anterior margin
thickened, well sclerotized: fifth segment not equal to II, Ill, and IV taken
together. Episternum III divided into upper and lower parts by a horizontal
suture. Pronotal comb present, composed of heavily pigmented spines, sharply

108 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

pointed apically. Hind coxa with a row or patch of spinelets on inner surface.
Femora with very few or no lateral setae. Segment V of each tarsus with 4
pairs of lateral plantar bristles. Abdomen without apical spines. Sternal
plate VIII of male large, broad, setigerous, unmodified. Sternal plate IX of
male with an anterior whiplike apophysis, a poorly sclerotized pair of inner,
vertical processes, and a simple pair of distal lobes.

Members of this genus were probably originally confined to Africa.
Two species, canis Curtis and felts Bouché, have been introduced into

many parts of the world.

CTENOCEPHALIDES CANIS (Curtis)
(Fig. 12, F)

Pulex canis Curtis, 1826, British Ent., vol. 3, No. 114.
Ctenocephalides canis Stiles and Collins, 1930, U. 8. Pub. Health Serv. Rpts.
45: 1308.

Type host.—Not indicated.

Type locality—Not indicated.

Range.—Almost cosmopolitan, but being apparently absent in cer-
tain regions inhabited by man although occurring on certain wild
carnivores in areas not yet reached by man. 7

In their summary of the distribution of Ctenocephalides canis in
the United States Trembley and Bishopp (76) record this flea from
31 States and the District of Columbia. The Rocky Mountain and
Intermountain regions are not represented. Hosts for which two or
more records are given include dog, man, cat, and gray fox.

Typical forms of Ctenocephalides canis are distinguished from
those of C. felis in having a much shorter forehead and in having
spine I of the head comb very distinctly shorter than spine IT, while
in C. felis spine I is about equal in length to spine II. In @. canis,
as well as in C. felis, there are certain sexual differences in the head
characters. ‘These are shown in some figures by I. Fox (24, p. 141).

Ctenocephalides canis is an important intermediate host of the do
tapeworm, Dipylidium caninum, a species very common in dogs an
' eats and occasionally found in man. C. canis has also been reported
aS an intermediate'host of the following parasites: Hymenolepis
-diminuta, Dirofilaria immitis, Leishmannia donovani, and L. infan-
tum. ‘This flea may be of importance in the transmission of plague
during epidemics.

The complete life cycle of Ctenocephalides canis is passed in from
2 to 4 weeks, which means that there may be several generations in
a year in the warmer sections of the world. Bacot (7) found that
C’. canis would live as long as 234 days when fed exclusively on man,

Three synonyms of this species are listed in Wagner’s catalog (80).

CTENOCEPHALIDES FELIS (Bouché)
(Figs. 1 and 2)

Pulex felis Bouché, 1835, Nova Acta Phys.-Med. Acad. Caes. Leop. Carol.
17: 505, fig. 2.

Ctenocephalides felis Stiles and Collins, 1930, U. S. Pub. Health Serv. Rpts.
45: 1309.

Type host.—‘Housecat.”
Type locality—Not indicated.

THE FLEAS OF NORTH AMERICA 109

Range.—Almost cosmopolitan, yet absent in many regions not
eee by man. Reported from several of the islands of the South

acific.

Trembley and Bishopp (76), summarizing their United States
records, list Ctenocephalides felis from 28 States and the District of
Columbia. They state that this flea is abundant in the eastern part of
the country but relatively scarce in the West. They had no speci-
mens from the Great Plains and Rocky Mountain regions, and among
material from the far West only Idaho, Oregon, and California
were represented. Numerous hosts are listed by these authors, but
most of the records are from man, dog, cat, and opossum.

In Ctenocephalides felis spine I and spine II of the head comb are
of about equal length, which fact distinguishes it from C. canis.
It has been noted by the present writers that the differential charac-
ters for separating these two species tend to become intermediate
where both species occur at the same locality, thus indicating the
possibility of hybridization.

Ctenocephalides felis and C. canis are almost cosmopolitan, both
species having followed man into many parts of the world. How-
ever, in certain regions one species may occur to the exclusion of the
other.

Ctenocephalides felis has been recorded as an intermediate host for
the tapeworm Dipylidium caninum and the nematode Dirofilaria im-
mites. It also may be an important vector of plague during epidemics
of this disease.

The larva of Ctenocephalides felis has been made the object of a
special study by Pinto (63).

Wagner (80, p. 34) indicates three synonyms of this species in his
catalog.

The Genus CrpropsyLia Jordan

(Fig. 13, F)

Cediopsylla Jordan, 1925, Novitates Zool. 32: 103. Type, Pulex inaequalis var.
simplez Baker (by original designation).

Anterior margin of front somewhat angulate. Genal comb more nearly verti-
cal than horizontal, composed of heavily pigmented spines with rounded apices.
Eyes large, black. Labial palpus 4-segmented, rather poorly sclerotized. An-
tennal club subcapitate, segments anchylosed anteriorly. Episternum III divided
by a suture into upper and lower parts. Pronotal comb composed of heavily
pigmented spines with sharply pointed apices. Hind coxa with a row or patch of
spinelets on inner surface. Femora with few or no lateral setae. Segment V
of each tarsus with 4 nearly equal pairs of lateral plantar bristles. Abdomen
short and high, without apical spines. .

Jordan established Cediopsylla for the American species of Spilop-
syllus. These he found differed from the European type species in
having the labial palpus four-segmented instead of two-segmented.
The type species of Cediopsylla, simplew Baker, which occurs in the
eastern part of the United States, differs from the two forms occurring
in western States rather markedly in the size of the mandibles and
the nature of the modified sternal plates of the male. Hence the
genus is divided into two subgenera.

The Subgenus CEDIOPSYLLA Jordan

Mandibles but slightly enlarged, not reaching tip of coxa I. Genal comb
composed of more than 5 spines. Sternal plate VIII of male almost completely

110 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

overlapped by VII, with a pair of simple, slender, slightly curved, horizontal,
posterior processes. Sterna! plate IX of male with anterior whiplike apophysis
and a pair of conspicuous, lobed, setigerous, posterior processes.

Type species —The same as for the genus.

Besides the type, which is a common parasite of rabbits in the east-
ern part of the United States, this subgenus includes Cediopsylla
spillmanni Jordan, a South American species.

CEDIOPSYLLA (CEDIOPSYLLA) SIMPLEX (Baker)

Pulex inaequalis var. simpler Baker, 1895, Canad. Ent. 27: 164.
Cediopsylla simplex Jordan, 1925, Novitates Zool. 32: 103.

Type host—Lepus sp. (hare).

Type locality—Michigan.

Range.—KEastern part of the United States.

This species, the only one of its genus in the eastern part of the
United States, is easily identified by its angulate forehead and sub-
vertical genal comb composed of eight black spines. It has been fig-
ured recently by I. Fox (25, p. 135).

Larvae of this species were placed individually in vials which con-
tained a variety of materials thought to be suitable for their food or
environment, the object being to observe their habits and rear adults.
In such vials no cocoons were made by the larvae before their trans-
formation into the pupalinstar. In one vial, in which a flea larva was
left without debris from the rabbit nest, a pupa was found. This pupa
was entirely concealed in the husk of a weed seed. When the husk was
broken open, only a naked pupa was seen. There was no evidence of
any silk to be found. This pupa was translucent to whitish, with the
legs extending free from the body and suggesting in their shape, to
a certain degree, the legs of an adult flea.

Data on the total length of the life cycle of Cediopsylla (Cediop-
sylla) simplex were not obtained, but a larva that was isolated in a
breeding vial on April 11 had transformed into an adult flea on
April 29.

The Subgenus ACEDIOPSYLLA Ewing

Acediopsylla Ewing, 1940, Wash. Biol. Soc. Proce. 53:37. Type, Pulex inaequalis
Baker (by original designation). Bh

Mandibles much enlarged, reaching considerably beyond tip of coxa I. Genal
comb composed of 5 spines. Sternal plate VIII of male not greatly overlapped
by VII, large, with a deep, broad, seta-margined, posteroventral depression and a
pair of expanded, broadly rounded, posterodorsal lobes. Sternal plate IX of male
with an anterior whiplike apophysis, a pair of curved, vertical, internal lobes,
and a pair of conspicuous, lobed, setigerous, posterior processes.

This subgenus includes one species which has two varieties, both
parasitic on rabbits.

CrpiopsyLLa (ACEDIOPSYLLA) INAEQUALIS INAEQUALIS (Baker)
(Fig. 13, E)

Pulez inaequalis Baker, 1895, Canad. Ent. 27: 163 and 164.
Cediopsylla (Acediopsylla) inaequalis Ewing, 1940, Wash. Biol. Soc. Proc. 53: 37.

Cotype hosts —‘Cottontail and jack rabbits.”
Type locality—Near Grand Canyon, Ariz.

THE FLEAS OF NORTH AMERICA 111

Range.—Southwestern part of the United States.
The original description of this species, although based upon a
series of specimens, is very short and unaccompanied with drawings.

CEDIOPSYLLA (AcEDIopsyLLa ) INAEQUALIS INTERRUPTA Jordan

Cediopsylla inaequalis interrupta Jordan, 1925, Novitates Zool. 32: 103.

Cotype hosts—Sylvilagus bachmanni (Waterh.), and “fox.”
Cotype localities—Palo Alto, San Francisco, and Claremont,
Calif.

Range—Known only from California.

The short original description of this variety, which is based on
both sexes, follows:

Genal comb usually with 6 or 7 spines, sometimes 5. Row of long bristles at
ventral margin of ¢-clasper divided into a proximal group of,4 to 6 anda
distal group of 2 or 3.

The Genus Ca.LiistopsyLLus Jordan and Rothschild

Callistopsyllus Jordan and Rothschild, 1915, Ectoparasites 1: 46. AYDe, Cera-
tophyllus terinus Rothschild (by original designation).

Genal comb absent. Eye vestigial, halfmoon shaped. Labial palpus 4-seg-
mented. Antennal groove not extending upward across vertex; second antennal
segment widened on outside covering a third of club. Pronotal comb composed
of many spines. Hind coxa without a row or patch of spinelets on inner surface.
Femora without lateral setae. Segment V of each tarsus with 5 pairs of lateral
plantar bristles, the first being displaced ventrally. Abdomen with a few
apical spines which are confined to first 3 terga. Sternum VIII of female angulate
posteriorly and bearing a marginal, apical row of very short micro-setae

This genus includes but two species, one of which is known only by
the female sex. The presence of apical spines on the abdomen indicates

a relationship with Dolichopsyllidae and Hystrichopsyllidae.

CALLISTOPSYLLUS DEUTERUs Jordan

Callistopsyllus deuterus Jordan, 1937, Novitates Zool. 40: 266, fig. 49.

Type host—Peromyscus sp. (white-footed mouse).
Type locality —Big Bear Lake, Calif.
Range.—Known only from California.

The original description of this species was based upon a single
female and is accompanied by a drawing of the female genitalia.

CALLISTOPSYLLUs TERINUS (Rothschild)

Ceratophyllus terinus Rothschild, 1905, Novitates Zool. 12: 158, pl. 8, fig. 26 and
pl. 9, fig. 29.
Callistopsyllus terinus Jordan and Rothschild, 1915, Ectoparasites 1: 46.
Type host.—Citellus columbianus (Ord) (ground squarrel).
Type locality—Mabel Lake, British Columbia.
fRange.—Known only from British Columbia.
This species is known only from female specimens.

112 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

The Genus Mscarturoctossus Jordan and Rothschild
(Fig. 13, A)

Megarthroglossus Jordan and Rothschild, 1915, Ectoparasites 1:46. Type,
Megarthroglossus procus Jordan and Rothschild (by original designation).

Anterior margin of front rounded. Frontal tubercle small to vestigial.
Genal comb absent. Eyes vestigial. Labial palpus 5-segmented, fifth segment
being equal to or longer than II, III, and IV taken together. Episternum III
divided into an upper and a lower part by a horizontal suture. Pronotal comb
present, composed of heavily pigmented spines with angulate ‘apices. Hind
coxa without a row or patch of spinelets on inner surface and with acute lobe
on posterior margin. Femur I without or almost without lateral setae. Seg-
ment V of last tarsus with 5 pairs of lateral plantar setae, the first pair being
shifted ventrally. Abdomen much longer than high; with some apical spines.
Sternal plate VIII of male much shortened, practically without setae, over-
lapped for much of its length by sternal plate VII and closely encasing sternal
plate IX. Sternal plate IX with a much reduced, anterior, whiplike apophysis,
a pair of somewhat expanded, internal, vertical processes, and a somewhat
clavate, Setigerous, posterior process.

This genus includes seven species from Western North America.

They are all very closely related and make a very compact natural
group.

MEGARTHROGLOSSUS BISETIS Jordan and Rothschild

Megarthroglossus bisetis Jordan and Rothschild, 1915, Ectoparasites 1: 54.

Type host—Neotoma sp. (wood rat).

Type locality —Beulah, N. Mex.

Range—Known only from New Mexico.

The very short original description of this species is based on two
females. It is said to be—

Like M. longispinus [now divisus], but with only two antepygidial bristles on

each side, the upper bristle being a little (one-fifth or one-sixth) longer than the
lower one.

MEGARTHROGLOSSUS DIvisus Divisus (Baker)
(Fig. 13, A)

Ceratophyllus divisus Baker, 1904, U. S. Natl. Mus. Proc. 27: 416, pl. 21, figs. 7-10.

Megarthroglossus longispinus Jordan and Rothschild, 1915, Ectoparasites 1: 52,
figs. 55, 56.

Type host—‘Fremont’s chickaree.”

Type locality —Georgetown, Colo. |

Range.—Both the species and variety occur in the western part of
the United States and Canada.

The name divisus was proposed by Baker to replace longispinus
Baker, 1895, preoccupied by Pulex longispinus Wagner, 1898. Baker’s
Pulex longispinus is a primary homonym and as such cannot, accord-
ing to the International Code, be resurrected because of the transfer of
Pulex longispinus Wagner to Xenopsylla, as has been done by Jordan
and Rothschild (1915).

MEGARTHROGLOSSUS DIVISUS EXSECATUs Wagner

Megarthroglossus longispinus exsecatus Wagner, 1936, Canad. Ent. 68: 196.

Type host-—Sciurus douglasid (Bach.) (red squirrel).
Type locality—Avola, British Columbia.

THE FLEAS OF NORTH AMERICA 113

Range.—Known only from British Columbia.
The original description of this variety was based upon a single
female and is included in its entirety in the following sentence:

This sinus in the 2 [apical sinus of abdominal sternal plate VII] from Avola
surprises by its bigness and I propose to name this variation ersecatus var. nov.

MEGARTHROGLOssUs PROcUs Jordan and Rothschild

Megarthroglossus procus Jordan and Rothschild, 1915, Ectoparasites 1: 47, figs.
0-52.

Type host.—Spilogale sp. (spotted skunk).

Type locality —Chilliwack, British Columbia.

Range-——Known only from British Columbia.

The original description of this species is very complete and is ac-
companied by three detail drawings.

MEGARTHROGLOsSUs PYGMAEUs Wagner

Negarthroglossus pygmaeus Wagner, 1936, Canad. Ent. 68: 196, pl. 11, fig. 4.

Type host—Neotoma cinerea (Ord) (wood rat).

Type locality—Nicola, British Columbia.

Range.—Known only from British Columbia.

The original description of pygmaeus is based on a single male.
The species is very close to divisus and sicanvws, but is distinct from
these two species in the structure of the genitalia. The species also
may be, according to Wagner’s statement, based only on the male of
Megarthroglossus bisetis Jordan and Rothschild, which was taken
from a Neotoma.

MEGARTHROGLOsSUS sICAMUs Jordan and Rothschild

arb so il es sicamus Jordan and Rothschild, 1915, Ectoparasites 1: 50, figs.
Type host—Canis latrans Say (coyote).
Type locality —Eagle River, Sicamous, British Columbia.
ange.—Known only from British Columbia.
The type host of sicamus is probably not a natural one. Natural
hosts most probably are rodents. The original description of this
species is accompanied by three detail drawings. |

MEGARTHROGLOsSUS SIMILIs Wagner

Megarthrogiossus similis Wagner, 1936, Canad. Ent. 68: 196, pl. 11, fig. 1.

Type host —Neotoma cinerea occidentalis (Baird) (wood rat).

Type locality —Beaverdell, British Columbia.

Range.—Known only from British Columbia.

The original description of similis is very short and based on two
males. However, it is accompanied by a drawing of part of the male
terminalia. According to the description, the structure of the male
genitalia of szmilis is intermediate between that of procus and that of
(longispinus) =divisus.

467453—43——_8

114 MISC. PUBLICATION 500, U. S. DH=PT. OF AGRICULTURE

MEGARTHROGLOsSUS SPENCERI Wagner

Megarthroglossus spenceri Wagner, 1936, Canad. Ent. 68: 196, pl. 11, figs. 2, 3.

Type host—Ochotona princeps (Rich.) (cony).

Type locality—Nicola, British Columbia.

Range.—Known only from British Columbia. E

The original description of spenceri was based on a single female, but
is accompanied by a drawing of the forehead and one of the female
terminalia.

The Genus ANomiopsyLLus Baker
(Fig. 12, F)

Anomiopsyllus Baker, 1904, U. 8. Natl. Mus. Proce. 27: 425. Monotypic. Type,
Typhlopsylia nudata Baker.

Anterior margin of front rounded. Frontal tubercle present, angulate apically.
Genal comb absent. Eyes absent. Labial palpus 4-segmented, never extending to
tip of trochanter I, well and equally sclerotized along anterior and posterior
borders. Antennal club long, segments not anechylosed anteriorly. Antennal
groove open posteriorly. Metathorax reduced, being shorter than mesothorax;
upper part of metepisternum, metepimeron, and metanotum fused, without a de-
limiting suture or inner ridge between them. Pronotal comb absent. Coxa II and
coxa III each with a posteroventral, spinelike lobe separated from coxa proper
by a deep incision. Coxa III without a row or patch of spinelets on inner surface.
Femora each with not more than a Single lateral seta. Segment V of tarsi I and
II with 5 pairs of plantar bristles, the first pair being situated between setae of
second pair; segment V of tarsus III with only 4 pairs of plantar setae, all being
lateral and forming 2 parallel rows. Abdomen much longer than high, without
apical spines. Sternal plate VIII of male considerably shortened and without
posterior processes. Sternal plate IX of male without anterior apophysis, with
a pair of large, expanded, inner, vertical processes, and a pair of more or less
flattened, clavate, posterior, spinigerous processes.

Baker (4, p. 127), in 1905, established for this genus the subfamily
Anomiopsyllinae. Although in this work the genus is not recognized
as constituting a subfamily, it is regarded as a somewhat isolated one,
particularly in the degree to which the metathorax has been reduced
and in the manner in which most of its sclerites have been fused. The

included species are all Nearctic.
ANOMIOPSYLLUS AMPHIBOLUS Wagner

Anomiopsyllus amphibolus Wagner, 1936, Ztschr. f. Parasitenk. 8: 654, fig. 1.

Type host—Neotoma desertorum Merr. (wood rat).

Type locality—Salina, Utah.

Range.—Known only from type locality.

In this species the movable finger bears two posterior, submarginal,
stumplike spines, and sternal plate IX of the male possesses four
terminal spines.

ANOMIOPSYLLUS FALSICALIFORNICUS C. Fox
(Fig. 12, #)
Anomiopsyllus californicus C. Fox, 1926, Pan-Pacific Ent. 2: 183, figs. 5 and 9.
Anomiopsyllus falsicalifornicus C. Fox, 1929, Ent. News 40: 218.

Cotype hosts—Neotoma fuscipes Baird (wood rat) and Spilogale
phenaz Merr. (spotted skunk).

Type locality —Los Angeles, Calif.

Range.—Known only from California.

THE FLEAS OF NORTH AMERICA eS

C. Fox (1926) misidentified this flea and described it under the name
of Anomiopsyllus californicus Baker. In 1929, after Jordan had
pointed out that it was really a new species, C. Fox proposed for it the
new name falsicalifornicus.

ANOMIOPSYLLUS MONTANUS Collins

Anomiopsyllus montanus Collins, 1936, Ent. News 47: 129, figs. 1 and 2.

Type host—Bubo virginianus occidentalis Stone (Montana horned
owl).

x sa locality Canyon Creek, west of Hamilton, Mont.

Range.—Known only from Montana.

The original description of montanus is accompanied by two draw-
ings, one of a part of the male terminalia and the other a part of the
female terminalia. The type host of this species is probably not a true
host. True hosts are probably rodents.

This species was based upon material misidentified as Anomiopsyllus
nudatus (Baker) by Dunn and Parker (12, p. 2774) in 1923.

ANOMIOPSYLLUs NuDATUs (Baker)

Typhlopsylla nudata Baker, 1898, N. Y. Ent. Soc. Jour. 6: 56.
AnomiopsylWus californicus Baker, 1904, Invertebrata Pacifica 1: 39.

Type host—Neotoma albigula (Hart) (wood rat).

Type locality—Tucson, Ariz.

Range.—Western part of the United States.

The original description of this species was based on two females.
Collins (6, p. 129), in 1936, published a drawing of sternal plate VII
of the abdomen of the female. Dunn and Parker (12, p. 2774), in 1923,
misidentified as Anomiopsyllus nudatus material which was later de-
scribed as representing a new species (A. montanus Collins) by Collins
(6, p. 129), in 1936.

The Subfamily CHIMAEROPSYLLINAE, new subfamily

Head divided into 2 parts by the presence of a well developed fronto-epicranial
groove or suture; postantennal region of head dorsally incrassate. Eyes present,
pigmented. Front considerably reduced, anterodorsal; gena extending dorsally
to level of upper margin of eye. Labial palpus short, not reaching apex of
coxa I. Pronotal comb present. Mesepisternum separated from mesepimeron
by an internal, vertical ridge. Hind coxa with a row of spinelets on inner
surface. Abdomen without apical spines.

Contains a single African genus, Chimaeropsylla Rothschild.

The genus for which this subfamily Chimaeropsyllinae is pro-
posed was described by Rothschild (64, p. 269) in 1911, being based
on a single species. He pointed out certain peculiarities of the mouth
parts stating:

* * * The mandibles as well as the labrum are very broad, particularly the
former, and closely resemble those of Spilopsyllus cuniculi, Dale (1878), and the
Sarcopsyllidae. This peculiar parallelism also obtains in the labial palpi, which
show a reduction similar to that which occurs in the fleas mentioned, being but
feebly chitinized and consisting of only three distinct segments.

Rothschild further states that the single included species probably
fixes itself to the host by means of the enlarged labrum and mandibles.
Chimaeropsylia is restricted entirely to Africa.

116 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

The Subfamily PULICINAE Tiraboschi

Head not divided by fronto-epicranial groove; postantennal region not dorsally
incrassate. Eyes large, nearly circular, and heavily pigmented. Front not
greatly reduced; frontal tubercle absent. Labial palpus short, not reaching apex
of coxa I. Pronotal comb absent. Mesepisternum not separated from mes-
epimeron. Hind coxa with either 1 or 2 rows or a patch of spinelets on inner
surface. Abdomen without apical spines.

This subfamily contains but three genera, Pulex Linnaeus, Jux-
tapulex Wagner, and Moeopsylla Rothschild. Only the two first-
mentioned genera are represented in the North American fauna.

The subfamily Pulicinae was proposed as such by Tiraboschi (75,
p.243) in1904. Itrepresents a very clear-cut natural group of genera.
The subfamily Pulicinae, as here recognized, does not include the
same genera as were included in Jordan’s classification of the Pulicidae
published in 1926 (39, p. 600). He included in the subfamily the
genera Zunga Jarocki, Hectopsylla Frauenfeld, and E'chidnophaga
Olliff.

Key to Norrh AMERICAN GENERA OF PULICINAE

Mandibles but slightly enlarged; front margin of head never angulate; genal
comb never represented by more than 1 pale, toothlike process. Occurring
in both New and’ Gld “Worlds=22 023422) a eee Pule# Linnaeus.

Mandibles much enlarged, as long as, or longer than, maxillary palpi; front
either round or angulate; genal comb represented by 2 short, pale, tooth-
like processes. Occurring in tropical and subtropical America

Jurtapuler Wagner.

The Genus Putex Linnaeus
(Fig. 12, A)

Pulex Linnaeus, 1758, Systema Naturae, ed. 10, p. 614. Type, Pulex irritans
Linnaeus (by subsequent designation).

Frontal region of head much higher than long. Eyes large, black, subangulate
ventrally. Two or 3 setae in ocular row. Antennal groove closed. Gena with
very broad ventral lobe; genal comb absent or represented by a single tooth.
Mandible not enlarged, much shorter than maxillary palpus. Labial palpus
with 4 distinct segments; anterior margin sclerotized. Metepimeron very much
more extended vertically than longitudinally. Hind coxa with a patch of spine-
lets on inner surface. Sternal plate VIII of male large, broad, unmodified;
sternal plate IX with vertical internal processes.

Linnaeus included two species in Pulex, the type species and P.
penetrans (now the type species of Zwnga Jarocki). As the genus is
defined in the present work, it includes but two species, the type

species and P. conepati Cunha, 1914. ;

PuLeEx IRRITANS DuGEsII Baker
(Fig. 12, A)

Pulez irritans var. dugesii Baker, 1899, Ent. News 10: 37.

Pulez dugesii Baker, 1904, U. S. Natl. Mus. Proc. 27: 379.
Type host.—“Spermophilis macrourus” (ground squirrel).
Type locality —Guanajuato, Mexico.
Range.—Known only from Mexico.

THE FLEAS OF NORTH AMERICA 7

Baker originally described dugesii as a variety. Later (1904) he
raised it to the rank of species, stating:

First described as a variety of irritans, this form is now given the rank of a
species. The examination of a large series Shows the characters to be uniform
and thoroughly distinctive.

An examination of the type specimens of dugesz by the writers in-
dicates that, except for the longer labial palpus, it agrees fairly well
with irritans. Hence it is given only the status of a subspecies in
this paper.

PuULEX IRRITANS IRRITANS Linnaeus

Pulez irritans Linnaeus, 1758, Systema Naturae, ed. 10, p. 614.
Pulex simulans Baker, 1895, Canad. Ent. 27: 65.

Type host.—*Man.”

Type locality—Sweden (?).

Range.—Of all the species of fleas, Pulex irritans is probably the
most nearly cosmopolitan in distribution. Yet there are good reasons
for not regarding it as truly cosmopolitan. It is now established in
all the major zoogeographical regions, yet there is evidence that it
does not occur in many wild areas uninhabited by man, and in addition
it probably is absent or extremely rare in certain large cities.

Jordan and Rothschild (47, p. 9), in 1908, said regarding irritans:

* * * Our knowledge of the variation and the present and former distribu-
tion of this insect is extremely meagre.

They stated:

* * * The nearest allies of this species which are found on various animals
are all inhabitants of the Old World. The fleas found in America are only
distantly related to P. irritans. It may therefore be concluded that the present
species originated from the Old World stock of Siphonaptera.

This statement was made a long time ago. Since then Jwxtapulex
poreinus (Jordan and Rothschild) and J. echidnophagoides Wagner
have been described from native hosts in the New World, the former
being originally placed in Pwlex. In view of these discoveries the
present writers are of the opinion that Pulex, as well as Jwxtapulea,
was originally an American genus, and that it was in the New World
that P. irritans originated.

The apparent absence of Pulex irritans in certain regions of the
Old World was noted by Jordan and Rothschild (47, p. 9). They
state that—

The tropical countries of the eastern hemisphere do not appear to be inhabited
by P. irritans except where Huropean settlements exist, or, at least, where
intercourse with Europeans goes on. * * *

The absence of Pulex irritans from.the oases of the Sahara and from the
Haussa countries south of the Sahara appears to be a well-established fact.

The distribution of Pulex irritans in the United States has been
especially considered by Ewing (18, p. 363), who stated in 1931:

* * * The unusual thing about its distribution has to do rather with its
sparseness in certain large areas than its absence in them. . Yet certain areas
exist in which repeated search for this flea has failed to reveal its presence.

These areas are chiefly wooded regions not inhabited by man. They
probably are free from infestation because of the absence of true

118 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

hosts, at least two of which are known, the pig and the prairie dog.
In the city of Washington it is questionable whether there is a true
infestation, and C. Fox and Sullivan (24, p. 1917) found that of the
4,408 fleas taken in the rat-fiea survey of New York City, extending
from April 18, 1923, to February 28, 1925, not a single specimen was
Pulex irritans.

Concerning the occurrence of this flea in the United States, Trembley
and Bishop (76) state:

* * * The human flea causes severe annoyance in buildings, farmhouses,
stables, and surrounding premises, especially in the Middle West and South and
on the Pacific Coast. It is the predominant species affecting man on the Pacific
Coast.

Bacot (7) has done much work on the biology and ecology of irritans.
He found that cold (40.9° F.) was fatal to the eggs and that warmth
(75° F.) combined with low humidity favored the fertility of the
egos. According to Bacot the newly hatched larvae were able to sur-
vive for several days to over a month without food. The cocoon period
ranged from 7 to 239 days, and adults, at 45°-50° F. with nearly
saturated air, lived without food for 125 days.

Pulex irritans varies considerably in America. Besides the variety
dugesii Baker, already considered in this paper, Cunha (8, p. 134)
described in 1914, as a new American variety, bahiensis, from Brazil.

The Genus JuxTapuLex Wagner
(Fig. 12, B)

Juxctapuler Wagner, 1933, Berlin Zool. Mus. Mitt. 18: 341. Type, Ju«rtapulen
echidnophagoides-Waguer (by original designation),

Front with anterior margin either broadly rounded or angulate. Eyes sub-
angulate ventrally. Two setae in ocular row. Antennal groove closed. Gena
with broad ventral lobe; genal comb composed of 2 short, apically angulate,
brown, spinelike processes on posteroventral aspect of gena. Mandibles enlarged,
extending almost to apex of coxa I. Labial palpus indistinctly 4segmented.
Metepimeron very much more extended vertically than longitudinally. Hind coxa
with a row or patch of spinelets on inner surface. Sternal plate VIII of male
large, broad, unmodified. j

Juxtapulex Wagner was established for two North American species,
the type species and Pulex porcinus Jordan and Rothschild. As
pointed out by Wagner, the genus, through the characters of its type
species, shows a decided affinity with Hchidnophaga Olliff.

JUXxTAPULEX ECHIDNOPHAGOIDEs Wagner

Juztapulex echidnophagoides Wagner, 1933, Berlin Zool. Mus. Mitt. 18: 348,
fig. 1A and figs. 2-6.

Type host—‘Dasypus novemcinctus” (nine-banded armadillo).

Type locality —Kast of Cartago, Costa Rica.

Range.—Known only from Costa Rica.

The original description of this species is extensive and accom-
panied with six detail drawings. Juatapulex echidnophagoides is
easily distinguished from J. porcinus (Jordan and Rothschild) be-
cause of the angulate front margin of its head. This margin in
porcinus is broadly rounded as in most fleas.

THE FLEAS OF NORTH AMERICA 119

JuxTaPULEx porcinus (Jordan and Rothschild)
(Fig. 12, B)

Pulex porcinus Jordan and Rothschild, 1923, Ectoparasites 1: 312, figs. 315-317.
Jurtapuler porcinus Wagner, 1933, Berlin Zool. Mus. Mitt. 18: 341.

Type host—Tayassu sp. (white-lipped peccary).

Type locality.—Texas.

Range.—Known only from Texas.

This species is easily distinguished from its congener by the shape
of the front margin of the head, which is broadly rounded instead of
angulate. Juatapulex porcinus is of particular interest because of the
nature of the genal comb. This is made up of from two to three short,
pale, triangular spines which appear to be true morphological spines,
since none of them is provided with a seta pit or a trichopore. The
anterior spine of the genal comb, when present, is smaller than the
other two. It may be represented by a vestigial spur at the base of the
second spine.

The Family STeEPHANOcIRCIDAE Wagner
y g

Head divided by fronto-epicranial groove into an anterior and a posterior
part. Gena greatly enlarged, being extended dorsally in front of antennal
groove and divided by a vertical suture. Forehead with a helmet composed of
greatly reduced front and anterior parts of enlarged genae; not elongated and
without ventral flaps. Eyes reduced or vestigial. Postantennal region of head
with 1 or more transverse, dorsal incrassations. Thorax not greatly reduced,
thoracic terga taken together longer than tergum I of abdomen. Typical ab-
dominal terga with more than 1 transverse row of setae. Antepygidial bristles
present. Abdominal apical spines usually present. Abdominal combs present or
absent. Females without a reduced number of abdominal spiracles, and when
gravid not greatly distended.

This family, as here recognized, is composed of five genera, three
of which are Neotropical and two Australian. While members of
the family Stephanocircidae exhibit probably the greatest modifica-
tions of the head of any group of the fleas, they are in most respects
generalized.

The Family Hecropsytymar Baker

Head not divided into an anterior and a posterior part by fronto-epicranial
groove. Gena not greatly enlarged and not divided by transverse suture. Fore-
head without helmet, not elongated, and not bearing ventral flaps. Eyes pres-
ent, pigmented. Both cephalic and pronotal combs absent. Postantennal
region without dorsal incrassations. Thorax greatly reduced; thoracic terga
taken together not so long as tergum I of abdomen. Abdomen almost nude,
but some or all of its segments with a single transverse row of small setae.
Antepygidial bristles absent. Abdominal apical spines and combs absent. Fe-
males sometimes with a reduced number of abdominal spiracles. Gravid fe-
males greatly distended.

This family is not divided into subfamilies.

_ Hectopsyllidae was established by Baker (4, p. 375) in 1904 for the
single genus Hectopsylla Frauenfeld, 1860. As recognized in this
work it includes three other known genera, Echidnophaga Olliff,
Rhynchopsyllus Haller, and Tunga Jarocki. The following five
synonyms of Hectopsyllidae are known: Sarcopsyllidae Taschen-
berg, 1880, based on Sarcopsylla Westwood, 1840, = Zunga Jarocki,

120 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

1838; Rhynchoprionidae Baker, 1906, based on Rhynchoprion Oken,
1815, = Z’unga Jarocki, 1838; Hectoropsyllidae Oudemans, 1906,
based on Hectoropsylius Oudemans, 1906, = Hectopsylla Frauen-
feldt, 1860; Dermatophilidae Oudemans, 1906, based on Dermato-
philus Guérin, 1839, = Tunga Jarocki, 1888; and Tungidae, appar-
ently first used by C. Fox (23, p. 130) in 1925 and based on Tunga.

Key To tHe Norra AMERICAN GENERA OF HECTOPSYLLIDAE

Coxa IIT with a patch of spinelets on inner surface; abdominal segments II and
III each with a pair of spiracles. Occurring chiefly in tropical and sub-

tropical regions of the World
Echidnophaga Olliff.

Coxa III without a patch of spinelets; abdominal segments II and III of female
without spiracles. Restricted to the warmer regions of Asia and Africa and

to tropical America
Tunga Jarocki.

The Genus EcuipnopuHaca Olliff
(Fig. 18, C)

Echidnophaga Olliff, 1886, Linn. Soc. N. S. Wales, Proc. (2) 1: 171. Monotypic.
Type, Echidnophaga ambulans Oliiff.

Argopsylla Enderlein, 1903, Deutsches Tief—See Exped. 1898-99, v. 3, p. 263.
Monotypie. Type, Sarcopsylla gailinacea Westwood.

Xestopsylla Baker, 1904, U. S. Natl. Mus. Proc. 27:373. Monotypic. Type,
Sarcopsylla gallinacea Westwood.

Front angulate along anterior margin, but not produced into a tubercle.
Maxilla triangular, sharply pointed at apex, and largely concealed within head.
Labial palpus very poorly pigmented and sclerotized, indistinctly segmented, and
closely ensheathing the large mandibles. Eyes large, black, without anterior
marginal incision. Hind coxa with a patch of spinelets on inner surface. Femur
III without toothlike projection at its base. Plantar bristles on segment V of
each tarsus of the usual kind, being short, well pigmented, and composing 4
pairs. Abdominal segments II and III each with a pair of spiracles. Sternum
VIII of male enlarged, broad, without long setae and posterior lobes, and com-
pletely overlapping sternum IX. Sternum IX of male with conspicuous anterior
apophysis, a pair of inner vertical processes, and a pair of simple, slender,
curved, posterior processes.

Type species—A monotypical genus with E'chidnophaga ambulans
Olliff as type. |

EcHIDNOPHAGA GALLINACEA (Westwood)
(Fig. 13, C)

Sarcopsyllus gallinaceus Westwood, 1875, Ent. Monthly Mag. 11: 246.

Pulex pullulorum Johnson, 1890, Wash. Ent. Soe. Proce. 1: 204.

Echidnophaga gallinacea Jordan and Rothschild, 1906, Liverpool Univ., Thompson
Yates and Johnson Lab. Rpts. (n.s.) 7: 52.

Type host—‘Domestic fowl.”

Type locality—Columbo, Ceylon.

Range.—Widely distributed in the tropical and subtropical regions
of the world, including Australia. In North America found
throughout the southern part of the United States and permanently
established as far north as Virginia and Kansas. Occasionally in-
troduced into the Northern States but not known to be established
there. In North Carolina Shaftesbury (69) examined 38 collections —
of this flea taken from 39 counties and found that its distribution
approximated the extent of the Lower Austral Life Zone.

THE FLEAS OF NORTH AMERICA 121

Trembley and Bishopp (76) have summarized their United States
records for Echidnophaga gallinacea, listing this flea from 22 States
and citing numerous hosts.

E'chidnophaga gallinacea, known as the sticktight flea, occurs on
many wild hosts and undoubtedly is yearly carried in large numbers
outside of its normal range. Its dispersal because of this fact has been
discussed by Stewart (73).

This fiea, in the adult stage, attaches firmly to its host and en-
gorges with blood. The females, as a rule, do not change their
position of attachment, but according to Ilingworth (32) the
“males move about during the night, “being attached in different
positions each morning pete

Many kinds of warm- “leg “vertebrates are attacked by Echid-
nophaga gallinacea. It has been reported from a considerable num-
ber of our wild birds and is a serious pest of poultry in the South,
where chickens frequently have dark flea-covered patches about the
eyes, comb, or wattles. Dogs and cats, as well as man himself, are
attacked. Rats, as hosts to this pest, play an important part in its
distribution, although the fleas are not very abundant on these
rodents. According to C. Fox and Sullivan (24, p. 1923), out of
3,928 fleas taken in a rat-flea survey of New Orleans in 1922-23, only
12 specimens were £’. gallinacea.

The life history of this species has been worked out by Parman
(62). He found that the incubation period lasted from 4 to 14 days,
the larval period from 14 to 31 days, and the pupal period from 9
to 19 days. The females became engorged and oviposition began
in from 6 to 10 days after attachment. Adults were killed by freez-
ing temperature and could not endure an incubator temperature of
100° F.

The Genus Tunea Jarocki
(Fig. 13, B, D)

Tunga Jarocki, 1838, Zoology or General Description of Animals, p. 50.
Monotypic. Type, Pulez penetrans Linnaeus.

Front produced into an angular tubercle along anterior margin. Maxilla
greatly reduced, narrowly rounded at apex, and entirely concealed within
head. Labrum, as well as mandibles, large. - Labial palpus very weak, poorly
pigmented, with indistinct segmentation, and practically devoid of setae. Eye
large, black. Hind coxa without a patch of spinelets on inner surface. Femur
III without toothlike projection at its base. Plantar bristles on segment V
of each tarsus slender, not darker than other setae on tarsi. Abdominal seg-
ments II and III of female without spiracles. Sternum VIII of male enlarged,
very broad, without long setae. Clasper slender; process similar in size and
shape to movable finger and forming with the latter a pair of forceps. Penis
complicated, consisting of a basal or anterior part hinged to a distal or pos-
terior part; anterior part composed of a dorsal, pedicellate, basal plate, arising
from segments I and II of abdomen, and a ventral, elbowed part attached to
distal part by a strong ligament; posterior part of penis bearing distally a
pair of slender, forked, almost straight parameres and a somewhat longer,
unpaired, slender process. Female with a spherical receptaculum seminis,
situated in posteroventral region of abdomen.

Tunga has the following synonyms: Dermatophilus Guérin, 1839;
Dermatophylus Lucas, 1839; Rhynchoprion Oken, 1815 (not Her-
mann, 1804); Sarcophaga Westwood, 1836 (not Meigen, 1826) ;
Sarcopsylla Westwood, 1840 (proposed to replace Sarcophaga West-
wood); and Sarcopsyllus Kolenati, 1857.

122 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

Besides the type species, which is restricted to the tropical regions
of Africa and the New World, five other species of Z’wnga are known,
including Z’wnga caecatus Enderlein, which is found in Brazil, and
T. caecigena Jordan and Rothschild, which has been taken from the
ears of sewer rats in China.

TUNGA PENETRANS (Linnaeus)
(Fig. 18, B, D)
Pulex penetrans L., 1758, Systema Naturae, ed. 10, p. 614.

Type host—Man

Type locality. —‘America.”

Range.—Widely distributed in the tropical regions of America
and Africa. Although reported from Florida, Louisiana, and Texas,
it probably is not permanently established in the United States.

This species is known as the jigger, the sand flea, or the chigoe.
Also it has been termed the chigger, but incorrectly so, as this last
term is more appropriately applied to the larvae of mites of the sub-
family Trombiculinae which parasitize vertebrates.

L. C. Johnson (38, p. 203) sent a note to the Entomological Society
of Washington in 1889 reporting the occurrence of a flea in Florida
on chickens, turkeys, etc., with habits somewhat similar to those of
Tunga penetrans. Unfortunately he referred to this flea as the jigger
fiea, a name which should have been reserved for 7. penetrans. His
flea, which he named Pulex pullulorum, proved to be a synonym of
Echidnophaga gallinacea (Westwood).

Faust and Maxwell (20), in 1930, reported an unusual case of the
infestation, by the larvae of Tunga penetrans, of the skin of the
pubic and inguinal regions of a man in New Orleans. The source of
-infestation appeared to be sisal hemp from Yucatan, since the patient
gave a history of having sat on piles of this hemp as the cargo was
being checked.

The life history of Tunga penetrans is rather remarkable, since
the fertilized female actually burrows into the skin of the host. Here
her engorgement with blood and the development of many eggs
brings about a great enlargement of the abdomen. This distension
may increase until the abdomen is spherical in shape and many times
its original size. The dilation of the abdomen takes place entirely
through the stretching of the intersegmental membrane between the
second and third abdominal segments. According to Hicks (28) eggs
hatch in from 3 to 4 days, while the entire life cycle is about 17 days.

In the case of man the favorite points of attack by this flea are
between the toes and under the toe nails. Here the enlarging females
cause intense itching and inflammation and produce swellings and.
ulceration. Finally secondary infection may result in tetanus or
gangrene. In addition to man, many of his domestic animals are
attacked. One of the writers (Ewing) identified specimens of this
flea (including eggs) taken from an adult male gorilla at Djaposten,
Africa, by H. Raven, February 1920. He also extracted many eggs
of penetrans from a lump of skin of a baby gorilla taken in the
Belgian Congo by Benjamin Burbridge in 1923.

According to Dalla Torre (10, p. 21) this species has as a synonym |
Sarcopsyllus canis Westwood, described in 1840 (19, p. 203).

LIST OF SYNONYMS OF NORTH AMERICAN
SPECIES Ob SIPMONAP FERA

(Synonyms are in italics, valid names in Roman)

Amphipsylla hadweni Ewing, 1927=Mioctenopsylla arctica Rothschild, 1922.
Anomiopsyllus californicus Baker, 1904—=Anomiopsyllus nudatus (Baker), 1898.
Ceratophyllus acasti Rothschild, 1905=Opisodasys pseudarctomys (Baker), 1904.
Ceratophyllus acutus Baker, 1904=Oropsylla (Diamanus) montana Baker, 1895.
Ceratophyllus aeger Rothschild, 1905—Orchopeas leucopus (Baker), 1904.
Ceratophyllus bertholfi C. Fox, 1927=Oropsylla idahoensis (Baker), 1904.
Ceratophyllus caedens durus Jordan, 1929—Orchopeas caedens (Jordan), 1925.
Ceratophyllus californicus Baker, 1904—Nosopsyllus fasciatus (Bosc), 1801.
Ceratophyllus canadensis Baker, 1904=Nosopsyllus fasciatus (Bosc), 1801.
Ceratophyllus citelli Stewart, 1928=Thrassis fotus (Jordan), 1925.
Ceratophyllus gibsoni C. Fox, 1914—Ceratophyllus gallinae (Schrank), 1803.
Ceratophyllus isus Jordan, 1925—Malaraeus bitterrootensis (Dunn), 1923.
Ceratophyllus italicus Tiraboschi, 1904=Nosopsyllus londiniensis (Rothschild),
1903.
Ceratophyllus lucidus Baker, 1904—Trichopsylla vison (Baker), 1904.
Ceratophyllus niger inflerus Jordan, 1929—Ceratophyllus niger C, Fox, 1908.
Ceratophyllus oculatus Baker, 1904=Nosopsyllus fasciatus (Bose), 1801.
Ceratophyllus peromysci Stewart, 1928—=Trichopsylla wagneri wagneri (Baker),
1904.
Ceratophyllus ponerus Rothschild, 1909=Odontopsyllus dentatus (Baker), 1904.
Ceratophyllus poeantis Rothschild, 1905=—Oropsyla idahoensis (Baker), 1904.
Ceratophyllus proximus Baker, 1904—Thrassis arizonensis (Baker), 1898.
Ceratophyllus rileyi Liu, 19835—Ceratophyllus diffinis Jordan, 1925.
Ceratophylius utahensis Chapin, 1919=Ceratophyllus garei Rothschild, 1902.
Ceratopsylla crosbyi Baker, 1905—=Myodopsylla insignis (Rothschild), 1903.
Corypsylloides spinata I. Fox, 1940—Corypsylloides kohlsi (Hubbard), 1940.

Ctenopsyllus hygini Rothschild, 1904—Nearctopsylla genalis genalis (Baker), 1904..

Hoplopsyilus minutus C. Fox, 1926—Hoplopsyllus (Huhoplopsyllus) glacialis foxi
Ewing, 1924.

Hoplopsyllus powersi C. Fox, 1926—Hoplopsyllus (EHuhoplopsyllus) glacialis foxi
Ewing, 1924.

Micropsylla peromyscus Dunn, November 23, 1923—Micropsylla sectilis (Jordan
and Rothschild), November 10, 1923.

Neopsylla hamiltoni Dunn, 1923—Rectofrontia fraterna (Baker), 1895.

Neopsylla similis Chapin, 1919=Epitedia wenmanni (Rothschild), 1904.

Neopsylla striata Stewart, 1926—Tamiophila grandis (Rothschild), 1902.

Opisodasys spatiosus I. Fox, 1940—Opisodasys robustus (Jordan), 1925.

fee spinifrons I. Fox, 1939=Peromyscopsylla hesperomys (Baker),
1904.

Pulex gillettei Baker, 1895—Orchopeas howardii (Baker), 1895.

Pulez longispinus Baker, 1895=Megarthroglossus divisus divisus (Baker), 1904.
Pulez musculi Dugés, 1832—Leptopsylla segnis (Schonherr), 1811.

Pulez pullulorum Johnson, 1890—Echidnophaga gallinacea (Westwood), 1875.
Pulez serraticeps Gervais, 1844—Ctenocephalides canis (Curtis), 1826.

Pulegz simulans Baker, 1895—Pulex irritans irritans Linnaeus, 1758.

Pulez wickhami Baker, 1895=Orchopeas howardii (Baker), 1895.

Sarcopsyllus canis Westwood, 1840=Tunga penetrans (Linnaeus), 1758.
Stenoponia wetmorei Chapin, 1919=Stenoponia americana (Baker), 1899.
Typhlopsylia americana Baker, 1895=Dactylopsylla ignota ignota (Baker), 1895.
Typhlopsylla mexicana (Baker), 1896=Leptopsylla segnis (Schénherr), 1811.

123

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124

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THE FLEAS OF NORTH AMERICA 125

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1840. ON THE CHARACTERS OF THE CHIGOE OR JIGGER. Ent. Soc. London.
Trans, (1) 2: 199-208.

ILLUSTRATIONS

(Figs. 4-13)

Ficure 4.—Side view of heads: A, Trichopsylloides oregonensis, male; B, Rho-
palopsyllus gwyni, male; C, Conorhinopsylla stanfordi, male; D, Arctopsylla
ursi, male; H, Chaetopsylla lotoris, female.

467459—43——_9 129

130 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

IE

FicurE 5.—A, head of Dactylopsylla bluei, male; B, head of Mioctenopsylla
arctica, female; C, head of Catallagia charlottensis, female; D, fifth tarsal
segment of leg II of Dasypsyllus gallinulae perpinnatus; E, antepygidial
bristles of Dolichopsylla stylosa, male; F', antepygidial bristles of Dolichopsylla
stylosa, female.

THE .FLEAS OF NORTH AMERICA 131

FIcuRE 6.—A, head of Ctenophyllus terribilis, female; B, tarsus of leg III of
Tarsopsylla coloradensis; C, tibia of leg III of Amphipsylla ewingi; D, head
of Stenistomera alpina; E, receptacula seminis of Atyphloceras multidentatus.

132 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

Fieurp 7.—A, movable finger, process of claSper, and sternal plate VIII of
Thrassis acamantis, male; B, movable finger and process of clasper of Opiso-
dasys keeni; C, movable finger and process of clasper of Orchopeas sexdentatus
sexdentatus ; D, head and prothorax of Oropsylla idahoensis, male; H, movable
finger and process of clasper of Ceratophyllus garet.

THE FLEAS OF NORTH AMERICA 133

FieurE 8.—A, head of Corypsylla ornata, male; B, genal comb of Epitedia wen-
manni, male; C, genal comb of Corrodopsylla curvata, male; D, genal comb
of Nearctopsylla genalis, male; F, genal comb of Peromyscopsylla hesperomys ;
F, head of Sternopsylla carlsbadensis, female; G, genal comb of Leptopsylla
segnis ; H, genal comb of Doratopsylla blarinae, male.

134 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

Ficure 9.—A, labial palpus of Stenoponia americana, female; B, genal ctenidium
of Ctenophthalmus pseudagyrtes, male; C, head of Paratyphloceras oregonensis,
female; D, metathorax of Rectofrontia fraterna, female; H, metathorax of
Micropsylla sectilis.

THE FLEAS OF NORTH AMERICA 135

Fieure 10.—A, head of Carteretta carteri, male; B, fifth tarsal segment of leg
III of Tamiophila grandis; C, head of Meringis dipodomys; D, head of Acteno-
phthalmus heiseri, female; £, fifth tarsal segment of leg III of Actenophthal-
hus heiseri, female; F, head of Hystrichopsylla pigas dippiei, female.

136 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

4
FIguRE 11.—A, combs of Eptescopsylla chapini, female; B, genitalia of Myodop-
sylla collinsi, male; C, combs of Myodopsylla insignis, female; D, head of
Sternopsylla terana, male; H, head of Myodopsylla coilinsi, female.
?

THE FLEAS OF NORTH AMERICA 137

FIcuRE 12.—A, head of Pulegx irritans dugesii, male; B, head of Juxtapulex
porcinus; C, genitalia of Hoplopsyllus (Hoplopsyllus) anomalus, male; D,
fifth tarsal joint of leg III of Xenopsylia cheopis, male; H, coxa III of Anomio-
psyllus falsicalifornicus, female; F', head, prothorax, and mesothorax of Cteno-
cephatides canis, male; G, coxa III of Xenopsylla cheopis, male.

138 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

Ficure 13.—A, head of Megarthroglossus divisus diwisus, female; B, coxa III of
Tunga penetrans, female; C, coxa III of Echidnophaga gallinacea, male; D,
head of Tunga penetrans, male; E, head of Cediopsylla (Acediopsyila)
imaequalis, male.

TAXONOMIC INDEX

(Valid names are in Roman, synonyms are in italics)

abantis, 57.
acamantis, 44, 45, 47, 48, 132.
acasti, 37, 123

Acediopsylla, subgenus of Eos 110.
acerba, 57, 62.
Actenophthalmus, 5, 75, 79.
ActenopsyHa, 102, 103.

acutus, 50, 123.

adelpha, 90.

adustus, 67.

adversa, 57.

aeger, 34, 123.

affinis, 4, 106.

agilis, subspecies of sexdentatus, 35.
alaskensis, 49.

albertensis, variety of ignota, 41, 42.
allos, 85.

alpina, 72, 73, 131.

ambulans, 120.

americana, species of subgenus Foxella, 41.
americana, Stenoponia, 84, 87, 123, 134.
Amonopsyllus, 56

Amphalius, 24, 53.

amphibolus, 114.

Amphipsylla, 24, 30.

anomalus, 105, 106, 137.
Anomiopsyllinae, 23.
Anomiopsyllus, 102, 114.

apachina, variety of ignota, 41, 42.
Aphaniptera, 14,

apricus, subspecies of celsus, 68.
Aptilopsylla, 97.

arachis, 86.

Archiscatopse, 11.

arctica, 43, 44, 123, 130.

arctomys, 49, 5

Arctopsyila, 16, ra 18.

Argopsylla, 120.

arizonensis, 45, 46, 123.

arizonensis, subspecies of arizonensis, 45.
armatus, 72

artius, 38.

asio, 57, 58.

asio, subspecies of asio, 57.

astia, 105.

atrox, 58, 67.

Atyphloceras, a. as

australis, subspecies of australis, 21.
avium, 69, 70.

bacchi, 45.

bahiensis, variety of irritans, 118.
bakeri, 58.

bertholfi, 49, 123.

bidentatiformis, ie

bisetis, 112, 113.

bishopi. 38.

bisoctodentatus, 82.

bitterrootensis, 58, 123.

blarinae, 82, 133.

bluei, 39, 40, 130.

bohlsi, 22.

borealis, Catallagia, 27, 28.
borealis. Ceratophy Hus, 67, 68.
Boreopsyllus, 24, 43,

brasiliensis, 105.

brooksi, 93, 94.

bruneri, 51.

cacicus, 21.

caducus, subspecies of gladiolis, 46.
caecatus, 122.

caecigena, 122.

caedens, 33, 34, 123.

californicus, Anomiopsyllus, 114, 115, 123.
californicus, Nosopsyllus, 54, 123.

Callistopsyllus, 102, 111.

canadensis, 54, 12

caninum, Dipylidium, 108, 109.

canis, Ctenocephalides, 4. 6, 107, 108, 109,
123, 137.

canis, Tunga, a Pas ERY

carlshadensis, = 98, 133.

Carteretta, 76,

earteri, 83, 135.° if

eascadensis, subspecies of sexdentatus, 35.

ee are 26, 28.

eatatina,

Gelencctia: 102, 109.

Cediopsylla, subgenus of Cediopsylla, 109.

celsus, 68.

celsus, subspecies of celsus, 68.

Ceratophyllidae, 15.

Ceratophyllinae, 15.

Ceratophyllus, 3, 14, 15, 16, 24, 30, 66, 67.

Ceratopsyllidae, 96.

Ceratopsyllus, 92, 93, 96.

Chaetopsylla, 16, 17

chamberlini, 27.

chapini, 99, 136.

charlottensis, 25, 26, 27, 28, 130.

cheopis, 4, 55, 103, 104, 105, 137.

Chimaeropsylla, 8, 115.

Chimaeropsyllinae, 15, 102, 115.

ciliata, 59.

ciliata, subspecies of ciliata, 59.

citelli, 46, 123.

clarus, 53.

Coleoptera, 11, 12.

collinsi, 100, 101, 136.

coloradensis, 32, 131.

comis, 40.

conepati, 116.

Conorhinopsylla, 73.

copha, subspecies of intermedia, 82.

Corrodopsylla, 76, 80, 81

Corypsylla, 90, 95.

Corypsylloides, 90, 96.

Craneopsylla, 5.

Cratynius, 7.

crosbyi, 101, 123.

eruzi, 81

cryptoctenes, 20.

Ctenocephalides, 5-8. 102! 107;

Ctenocephalus, 107,

Ctenonotus, 31.

Ctenophthalminae, 15, 74, 75.

Ctenophthalmus, 5, 76, 82.

Ctenophyllus, 24, 72.

Ctenopsyllidae, 75, 90.

Ctenopsyllinae, 90.

Ctenopsyllus, 90, 92, 93.

cummingi, 86.

cuniculi, 115.

curvata, 80, 81, 133.

curvata, subspecies of curvata, 81.

eynomuris, variety of tuberculata, 52.

cyrtura, subspecies of eumolpi, 60.

Dactylopsylla, 24, 39, 40, 41, 43.

Dactylopsylla, subgenus ‘of Dactylopsylla, 40.

dasycnemus, 81,

Dasypsyllus, 24, 30.

decipiens, 27, 28.

Delotelis, 23. 26527:

dentatus, 25, 26, 123.

Dermatophilidae, 120.

Dermatophilus, 120, 121.

Dermatophylus, 121,

desertorum, 46.

deuterus, 111.

139

140

Diamanus, subgenus of Oropsylla, 24, 48, 50.

diasi, 101.

dieteri, 33.

diffinis, 68, 123.

digitalis, 72.

diminuta, Hyatenolepis, 108.
dipodomys, 86,

dippiei, 88, 89, 135.

Diptera, Tbe soy alah

dissimilis, subspecies of penicilliger, 63.
divisus, 112, 113.

divisus, variety of divisus, 112, 123, 138.
dolens, 65.

Dolichopsylla, 24.29, 39:

Dolichopsyllidae, 13, 14, 15, 16,.29, 75, 102.

Dolichopsyllinae, 14, 15, 23.

donovani, Leishmannia, 108,

Doratopsylla, 76, 80, 815 82°

Buccs, subspecies of irritans, 116, 117, 118,

dunni, 22.

durus, subspecies of caedens, 33, 123.

ebrighti, 91.

Echidnophaga, 12, 116, 118, 119, 120.

echidnophagoides, 117, 118.

echis, 38, 39.

enoplus, 36.

Epitedia, 75, 77, 78.

Eptescopsylla, 97, 99.

eremica, 59, 60.

Euhoplopsyllus,
106.

eumolpi, 60, 61, 63.

eumolpi, subspecies of eumolpi, 60.
euphorbi, 60.

ewingi, 31, 131.

exilis, 60.

exilis, subspecies of exilis, 60.
exoticus, 106.

exsecatus, variety of divisus, 112.
faceta, 77.

falaichit cendeee h4, 15, 137,
fasciatus, w) BITES By 54, 55, 123.
felis, 4, 5, a. 108, 109,

felix, 39.

floridensis, vie

fornacis, 61.

fotus, 46, 123.

Foxella, Subgenus of Dactylopsylla, 24, 40,

41, 43
foxi, variety of glacialis, 107, 123.
Hench oie, (03y,
franciscana, variety of ignota, 41, 42.
francisi, 46.
fraterna, 80, 123, 134.
gallinacea, 120, 121, 1238, 138.
gallinae, 69, (al 123.
gallinulae, 30.
garei, 68, 69, 123, 132,
genalis, 94, 133.
genalis, subspecies of genalis, 94, 123.
gentilis, 101.
gibsoni, 69, 123.
gigas Baker, syn. of grandis, 76.
gigas Kirby, Hystrichopsylla, 76, 88.
gigas, subspecies of gigas Kirby, "88.
gillettei, 353, 123.
gilvus, 70.
glacialis, 106.
glacialis, variety of glacialis, 106, 107.
gladiolis, 46.
gladiolis, subspecies of gladiolis, 46, 47.
globiceps, 17.
grandis, 76, 88, 123, 135.
graphis, 66.
groenlandica, 61.
gZwyni, 22, 23) 129.
hadweni, ‘43, ‘44, 1238},
hamifer, 91.
hamifer, eupspeces of hamifer, 91.
hamiltoni, 80, 123.
Hectopsylia, 416. 119,
Hectopsyllidae, 13, 14, 119.
Hectoropsyllidae, 120.
Hectoropsyllus, 120.
heiseri, 79, 135.
Hemiptera, 10.
hesperomys, 90, 91, 123, 133.

subgenus of Hoplopsyllus,

MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

hirsuta, 50, 51.

hirundinis, 66.

homoeus, 16, 17.

Hoplopleura, 73.

Hoplopsyllus, 5, 102, 105.

Hoplopsyllus, subgenus of Hoplopsyllus, 105.
howardii, 32, 33, 34, 123;

howelli, 47. s,
howelli, subspecies of howelli, 47.
hubbardi, species of Meringis, 86.
hubbardi, species of Trichopsylloides, 19.
hygini, 94, 128.

Hymenoptera, 12,

hyrtaci, 94.

Hystrichopsylla, 84, 85, 88.
Hystrichopsyllidae, 13, 14, he iy
Hystrichopsyllinae, 15, 74, 8

idahoensis, 49, 128, 132.

idius, 70.

ignota, 41, 42, 43.

ignota, variety of ignota, 41, 123.

immitis, species of Dirofilaria, 108, 109.
immitis, species of subgenus Trichopsylla,

inaequalis, 109, 110, 138.
inaequalis, variety of inaequalis, 110.
infantum, Leishmannia, 108.
inflecus, subspecies of niger, 70, 123.
inopina, 78.
insignis, 100, 101, 123, 136.
Integricipita, 75.
interrupta, variety of inaequalis, 111.
Intuberata, 89.
irritans, 4, 55, 1G. Ti bees
irritans, subspecies of irritans, 117, 123.
Ischnopsyllidae, 13, 14, 96,
Ischnopsyllinae, 96, 97.
Ischnopsyllus, 96.
isus, 58, 123.

123.

italicus, 55,
jellisoni, 81.

jordani, Corypsylla, 95.

jordani, Epitedia, 78.

jordani, Nearctopsylla, 95.

jordani, Trichopsylla, 62.
Juxtapulex, 116, 117, 118.

keeni, 37, 132.

klagesi, 32,

kohlsi, 96, 123.

labiatus, 33.

labis, 51, 52.

latens, 34.

laurentina, subspecies of genalis, 94.
Lepidoptera, al 74

Leptopsylla, 90, 92, 93.
Leptopsyllidae, "5.

Leptopsyllinae. 15, 74, 89, 90.
leucopus, 34, 123.

littoris, subspecies of arizonensis, 45.
Loemopsylla, 103.

londiniensis, 55, 123.

longiloba, supspecies of hamifer, 91.
longispinus, E12, 123.

lotoris, 17, 18,

lucidus, 64, 123.”

lucifer, 62.

lutzii, 50, Pall

lynx, variety of glacialis, 106, 107.
Macropsylla, 89

Macropsyllinae, 15, 74, 89.

Matlaraeus, 24, 55, 56, 60.
Mammoth, 89.

mandarinus, 50,

Mecoptera, 12.

Megabothris, 24, 55, 56, 58.

megacolpa, 62.
Megarthroglossus, 102, 112.
Meringis, 85,

mesus, 21, 39°

mexicana, in subgenus Foxella, 43.
mexicana, synonym of segnis, 93, 123.
Micropsylla, 75, 79.

minutus, 107, 123,

Mioctenopsylia, 24, 43.

Moeopsylla, 8, 116.

monedulae, 72,

moneris, 28.

mononis, subspecies of ciliata, 59.

THE FLEAS OF

Monopsylius, 56. :

moritatie, species of subgenus Diamanus, 5¥,
123:

montanus, Anomiopsyllus, 115.

motei, 28.

multidentatus, 38, 39, 131.

multispinosus, 25.

mundus, 65, 66.

musculi, 92, 93, 123.

Myodopsylla, 97, 100.

Nearctopsylla, 90, 93, 96.

necopinus, 54. s

Neopsylla, 5, T5, at, 15:

Neopsyllidae, 75.

neotomae, 31.

nepos, 33, 34, 55.

Neuroptera, 12. a

nevadensis, subspecies of sexdentatus, 35.

niger, 69, 70,123.

Nosopsyllus, 24, 54, 57.

novemdentatus, 107.

nudatus, 144, 115, 1238.

Nycteridopsylla, 99. _

Nycteriodopsyllinae, 97.

obtusata, subspecies of curvata, 81.

octodecimdentatus, 31.

oculatus, 54, 123.

Odontopsyllus, 23, 25, 39.

onaga, 27, 28.

Oncopsylla, 16. _

opada, subspecies of exilis, 61. _

ophidius, subspecies of wagneri, 64.

Opisocrostis, subgenus of Oropsylla, 8, 24,
49, 50, 51.

Opisodasys, 24, 36.

Orchopeas, 24, 32. (

erecta, subspecies of asio, 58.

oregonensis, Paratyphloceras, 83, 84, 134.

oregonensis, species of subgenus Opisocros-
tis, 52, 53.

oregonensis, Trichopsylloides, 19, 129.

ornata, Corypsylla, 95, 133.

ornata, variety of tuberculata, 52.

Oropsylla, 24, 48, 50.

Oropsylla, subgenus of Oropsylla, 48, 49.

Oxytelus, 11.

pechyuromydis, 103,

palposa, 100. ¢

Pancoleoptera, 12.

pandorae, 47.

Panmecoptera. 12.

Pansiphonaptera, 12.

pansus, 47.

paradisea, 85.

Parapsyllinae, 20.

Parapsyllus, 20.

Parapulex, 103.

Paratrichoptera, 12.

Paratyphloceras, 75, 83.

parkeri, 86, 87.

penetrans, 116, 121, 122, 123, 138.

penicilliger, 16, 17, 55, 56, 63.

pennsylvanicus, subspecies of sexdentatus,
a.

pentacanthus, 8&0.

pentactenus, 80.

Permochoristidae, 12.

peromysci, 65, 123.

Peromyscopsylla, 90.

peromyscus, 79, 80, 123.

perpinnatus, subspecies of gallinulae, 30, 130.

petiolatus, 48.

petrochelidoni, 70, 71.

Phalacropsylla, 85.

Phaneris, 19.

‘ cane, subgenus of Trichopsylla, 24, 56,

a.
poeantis, 49, 123.
pollionis, subspecies of sibirica, 31.
Polygenis, subgenus of Rhopalopsyllus, 22.
ponerus, 25, 123.
porcinus, 117, 118, 119, 137.
Posttuberata, 89.
powersi, 107, 123.
procus, 112, 113.
protina, subspecies of ciliata, 59.
Protohymenoptera, 12,
prorimus, 45, 123.

NORTH AMERICA 141

pseudagyrtes, 83, 154.

pseudarctomys, 37, 38, 125.

Ptilopsylla, 97.

Pulex, 8, 116.

Pulicidae, 13, 14, 16, 20, 23, 101.

Pulicinae, 15, 23, 102, 116.

pullulorum, 120, 122, 123.

pygmaeus, 113.

quebecensis, 71.

quirini, 63.

rara, 43.

rayalliensis, 92.

Rectofrontia, 8, 75. 79, 80.

recula, variety of ignota, 41, 42.

Rhadinopsylla, 80.

Rhineclophopsylia, 97, 100.

Rhopalopsyllinae, 14, 15, 20, 23.

Rhopalopsyllus, 20, 21.

Rhopalopsyllus, subgenus of Rhopalopsyllus,
21.

Rhynchoprion, 120, 121.
Rhynchoprionidae, 120.

Rhynchopsyllus, 119.

rileyi, 68, 123.

riparius, 67, 71.

roberti, 22.

robustus, 37, 123.

Rothschildella, 20, 21.

rothsehildi, 16.

runatus, 53.

rupestris, 50, 51, 52.

rusticus, 70.

saevus, subspecies of cacicus, 21.
samuelis, subspecies of klagesi, 22.
Sarcophaga, 121.

Sarcopsylla, 119, 121.

Sarcopsyllidae, 119.

Sarcopsyllus, 121.

saundersi, 52.

scapani, 78.

Seatopse, 11.

schefferi, 89.

schelkovnikovi, 30.

schisintus, subspecies of sexdentatus, 36.
sciurobius, 93.

sciurorum, 56.

scotti, 92.

sculleni, 28.

sectilis, 79, 80, 123, 134.

segnis, 92, 93, 123, 133.

selenis, 92.

serraticeps, 123.

setosa, 18.

sexdentatus, 34, 35, 132.

sexdentatus, subspecies of sexdentatus, 34.
shannoni, 87.

sibirica, 31.

sibynus, 66, 67.

sicamus, 113.

sigmodoni, 22, 23.

silantjewi, 48, 49.

similis, species of Epitedia, 79, 123.
similis, species of Megarthroglossus, 113.
simplex, 109, 110.

simulans, 117, 123.

sinoma, 57, 63.
Siphonaptera, 12, 13, 14, 16.

spatiosus, 37, 123.

spenceri, species of Megarthroglossus, 114.
spenceri, species of Odontopsyllus, 26.
spenceri, species of Thrassis, 48.

Spicata, subgenus of Dactylopsylla, 43.
spillmanni, 110.

Spilopsyllinae, 15, 102.

Spilopsyllus, 8, 102, 103, 109.

spinata, 96, 123.

Spinifrons, 91, 123.

sie anes species of Conorhinopsylla, 73, 74,

stanfordi, species of Thrassis, 48.
Staphylinoidea, 11.

Stenistomera, 7, 24, 72, 73.
Stenoponia, 84, 85, 87.
Stenoponiinae, 84.

Stenopsylla, 81, 82.
Stephanocircidae, 13, 14, 119.
Stephanocircus, 89, 90.
Sternopsylla, 5, 97, 98, 99.
stewarti, 78.

ee

142 MISC. PUBLICATION 500, U. S. DEPT. OF AGRICULTURE

stimsoni, 40.

Strepsiptera, 12.

striata, 76, 123;

stylosa, 29, 130.

suavis, 103.

subulata, 101.

Suctoria, 14.

swansoni, 71.

systalta, ‘subspecies of wagneri, 64, 65.
Tabanidae, 12.

tahavuana, subspecies of gigas, 88.
talpae, 88.

Tamiophila, 75, 76, 77.

tamoyus, subspecies of australis, 21.
Tarsopsylla, 24, 31, 32.

telchinum, 55, 57, 63, 64.

telegoni, 25, 26.

terinus, 111.

terribilis, iPpalsae

testor, 78.

Tetrapsyllus, 20.

texana, species of Sternopsylla, 97, 98, 99,

texana, species of Tamiophila, 76, 77.

thamba, 64.

Thrassis, 24, 44.

Tiamastus, subgenus of Bhove ope Hue. 21.
Trichopsylla, i eee 67.
Trichopsylla, subgenus fee "Trichopsylla, 56,

65.
Trichopsylloides, 8: 215; 2695
Trichoptera, 11, 12.
tripectinata, 87.

tripta, subspecies of exilis, 61.
Tritopsylla, 81.

tuberculata, 53.

tuberculata, variety of tuberculata, 52, 53.
tuberculaticeps, 18, 19.

Tunga, 14, 116, 119, 120, 121.
Tungidae, 120.

tupinus, subspecies of australis, 21.
Typhloceras, 84.

Typhloceratidae, 75.

unipectinata, 100.

Uropsylla, 73.

Uropsyliinae, 14, 15, 73.

ursi, 18, 19, 129.

utahensis, ‘Ceratophyllus, 69, 123.
utahensis, subspecies of howelli, 47.
utahensis, variety of ignota, 42.
vagabundus, Whe eas

vancouverensis, 99.

Vermipsylla, 13, 16.

Vermipsyllinae, 14, 15, 16.

vesperalis, 36, 37, 38.

vidua, subspecies of intermedia, 82.
vigens, subspecies of hamifer, 91.
vison, 64, 123.

vulpes, 1%

wagneri, 58, 65.

wagneri, subspecies of wagneri, 64, 65, 123.
walkeri, 55, 56

washingtonensis, 53.

wenmanni, 77, 79, 123, 133.
wetmorei, 87, 123.

wickhami, 32, 33, 123.

wolthaerey 5.

wymani, 29.

Xenopsylla, 102, 103, 104, 112.
Xestopsylla, 120. ‘

ORGANIZATION OF THE UNITED STATES DEPARTMENT OF AGRICUL-
TURE WHEN THIS PUBLICATION WAS EITHER FIRST PRINTED OR
LAST REVISED.

peeterary. of Agricalisire=- =!) ea CLAUDE R. WICKARD.
rE SCCTORITY 5 Es Oe ee ets ee Pavut H. APPLEBY.
Eee SOCTCUIT YS eS Se ee oe Se Ee GROVER B. HILL.

Chief, Bureau of Agricultural Economics_____--______ Howakgp R. TOLLEY.
Director of Agricultural Defense Relations____.________ M. CLIFFORD TOWNSEND.
Dérector of Fiztension Work __-> == = M. L. WILSON.

SU ee i Bo) | ee Oe ee en ape rede ee ae W. A. JUMP.

Director of Foreign Agricultural Relations____________ L. A. WHEELER.

p2 gee ae aes ir b 4), , eae a lee ay A ee Morse SALISBURY.

pi ge OS a eh) 2 eel Aen ie T. Roy Re.
Re RS a Sea MILTON S. EISENHOWER
DT a SR RE EA ee a Pa eco RALPH R. SHAw.
2 aS ST ae eee nee se MASTIN G. WHITE.
Chief, Office of Civilian Conservation Corps Activities_. FRED MORRELL.

Chief, Office of Plant and Operations______.-__-___--_ ARTHUR B. THATCHER.
Administrator of Agricultural Marketing___._.______-- Roy F. HENDRICKSON.
Administrator, Surplus Marketing Administration. E. W. GAUMNITz.
Chief, Commodity Exchange Administration____-- JOSEPH M. MEHL.

Chief, Agricultural Marketing Service_____-_____- CLARENCE W. KITCHEN.
Administrator of Agricultural Adjustment and Conser-

SOT 0 Ee ili ee eer yn R. M. Evans,

Administrator, Agricultural Adjustment Adminis-

UTED DE i i SS ee Sete 2 ae FRED S. WALLACE.

Chief, Soil Conservation Service__.______.____--~- HueH H. BENNETT.

Manager, Federal Crop Insurance Corporation_._._._ Leroy K. SMITH.

TE AP RE | a eee ee JOSHUA BERNHARDT.
Administrator of Agricultural Research___---_--------- E. C. AUCHTEE.

Chief, Bureau of Animal Industry_--------------- JOHN R. MOHLER.

Chief, Bureau of Agricultural Chemistry and

PCOS So) A Ee 2 es eee Henry G. KNIGHT.
Chief, Bureau of Dairy Industry__-_--_-_-_-------- OLLIE E. REED.
Chief, Bureau of Entomology and Plant Quaran-

ee ee 2 ee P. N. ANNAND.

Chief, Office of Experiment Stations_____-_----__-. JAMES T. JARDINE.

Chief, Bureau of Plant Industry___-_--_----__------- E. C. AUCHTER.

Chief, Bureau of Home Economics__-------------- _ LOUISE STANLEY.
President, Commodity Credit Corporation_____------~-- J. B. Hutson.
Administrator of Farm Security Administration___---- C. B. BALDWIN.
Governor of Farm Credit Administration___-__--_----- ALBERT G. BLACK.
CRICE TS presr Ber rere oe oe ee EARLE H. CuLapp, Acting.
Administrator, Rural Electrification Administration._ HARRY SLATTERY.

143

U.S. GOVERNMENT PRINTING OFFICE: 1943