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RECEIVED BY GIFT FROM

Pacific Coast Entomological Society
October 19, 1951

Memoirs of the Pacific Coast Entomological Society

Volume 1

THE SUCKING LICE

G. F. FERRIS

Professor o t Entomology
Stanford University, California

With the Collaboration of

Chester J. Stojanovich

Communicable Disease Center
United States Public Health Service
Atlanta , Georgia

SAN FRANCISCO. CALIFORNIA

October, 1951

Published and for sale by

THE PACIFIC COAST ENTOMOLOGICAL SOCIETY
California Academy of Sciences, Golden Gate Park, San Francisco 18, California

Printed by

New York Lithographing Corporation, New York

"In the progress of this work, however, the author has had to contend
with repeated rebukes from hts friends for entering upon the illustration
of a tribe of insects whose very name was sufficient to create feelings of
disgust. " — Henry Denny, 1842.

PREFACE

Tli is volume represents an attempt to summarize existing knowledge on the
systematics of the sucking lice, which are here considered to be the Order
Anoplura of the Insecta. Since sound taxonomy should rest upon morphology
and anatomy, an extended consideration will be given to these subjects.
Included also will be a review of the historical development of our knowl¬
edge of the group; a consideration of the classification here adopted; keys
to the families, subfamilies, genera, and species and a review of all these
categories; illustrations of all the available types of genera and of the
species occurring on domestic animals; an extended discussion of the spe¬
cies occurring on man and other Primates; a list of the mammalian hosts;
and finally, a consideration of the problems of geographical distribution.

I wish here to express my appreciation of the assistance given me by my
collaborator, Mr. Chester Stojanovich, whose unusual skill in minute dis¬
section and in illustration has made possible the sections on the mouth-
parts and internal anatomy. Mr. Stojanovich began this work while he was a
graduate student at Stanford University and later was granted leave from
his duties with the Communicable Disease Center of the United States Public
Health Service, which enabled him to return to my laboratory during the
summer of 1950 and complete it. I wish here to express my thanks to Dr. R.
A. Yonderlehr, Medical Director in charge of the Communicable Disease Cen¬
ter, by whom this leave was officially granted and to Dr. G. H. Bradley, by
whom it was recommended. My part in the preparation of the sections in
question has been that of supplying the textual material. Mr. Stojanovich
assisted also in the preparation and testing of various of the longer keys
to species.

The purely taxonomic portion of the work, all names therein proposed,
and all opinions herein expressed are to be ascribed strictly to myself.
This work has its basis in a series of studies published by me over a peri¬
od of years from 1919 to 1935> in eight papers entitled "Contributions
Toward a Monograph of the Sucking Lice." These appeared as Volume II of
what was in part entitled Leland Stanford .Junior Publications, University
Series, and in part Stanford University Publications, University Series,
Biological Sciences. In them will be found illustrations of almost all of
the species known up to 1935> together with a statement of the sources of
the material upon which the work was based. Owing to the cumbersome nature
of the full citation, this work will herein be cited merely as "Contribu¬
tions Toward a Monograph of the Sucking Lice."

This volume is closed as of July 1, 1951*

Stanford University, California

vii

TABLE OF CONTENTS

CHAPTER I

The Ectoparasites of birds and Mammals . 1

CHAPTER II

The Morphology and Anatomy of the Auoplura .

CHAPTER III

Growth and Development . 50

CHAPTER IV

The Taxonomic Status of the Sucking Lice . 53

CHAPTER V

The Classification (Vithin the Order Anoplura . 63

CHAPTER VI

Review of the Families, Subfamilies, Genera, and Species of the Anoplura 71
Errata . 285

CHAPTER VII

Host List . 286

304
311

CHAPTER VIII

The Distribution of the Anoplura .

INDEXES .

CHAPTER I

The Ectoparasites of Birds and Mammals

When in the course ol' their evoLution the birds achieved leathers and
the inanunaLs achieved hair, a new world to conquer was offered to the Arth-
ropoda which must have swarmed upon the earth from almost the earliest a^es
of terrestrial life. Some of the arthropods moved into this new world and
became adjusted to the peculiar conditions of life that it presented. From
these pioneers came the ectoparasites that today infest in some measure
probably every species of bird and evex'y species of mammal except the Ce¬
tacea and the Sirenia.

These ectoparasites, as we see them today, have been derived from many
sources. One may surmise that the earliest to enter this world of hair and
feathers were mites, for the mites are a very ancient group and they may
very' well have fed upon the blood of hairless amphibians and reptiles, even
as they do today, before the birds and mammals had evolved. Today the
mites constitute probably the major ectoparasitic fauna ol' the vertebrates.
They have penetrated into the lungs of their hosts, into the ears, into the
skin, and even at times into the intestine and bladder and they swarm in
the hair and the feathers. They induce pathological conditions of their
hosts by their own activities and they act as transmitting agents of numer¬
ous diseases that are already' known and undoubtedly many more that are not
yet known. They are the greatest of all the groups of ectoparasites and
scarcely more than a beginning of their study has yet been achieved.

It seems probable that only after hair and feathers had become well de¬
veloped did the insects move in. These insect ectoparasites have come from
various Orders. There are the blood-sucking flies of the family Hippobos-
cidae, which occur on both birds and mammals, and the families Streblidae
and Nycteribi idae , which occur only on bats. There are the "bedbugs,"
which are llemiptera, of the family Cimicidae, and which occur chiefly on
birds and bats. There is the more or less closely related Hemipterous fam-
i Ly Polyctenidae, which occurs only on bats and which numbers scarcely two
dozen known species. There is at least one speciesofthe Order Dermaptera,
which occurs on an African rodent. There are a few beetles, including that
singular parasite, Platypsylla castoris Ritsema, which occurs only on
beavers. There is even a moth which lives in its larval stages in the hair
of a sloth and which may justifiably be included in the list. There is the
whole Order Siphonaptera, the fleas, which in their adult stage live upon
both birds and mammals and feed upon their blood.

There are two groups of what are commonly called lice. One of these
groups comprises what are frequently called "bird lice," although a consi¬
derable number of them live upon mammals. They are more appropriately-
called "biting lice" and constitute what is here considered to be the Order
Mallophaga, which includes something more than 2,000 known species. And
finally, there is the little group which is the subject of this volume, the
sucking lice, comprising what is here called the Order Anoplura. At pres¬
ent scarcely more than 225 species of Anoplura are known. They live exclu¬
sively upon mammals and feed upon the blood oi their hosts.

The ectoparasites of birds and mammals present many features of general
biological interest. Some of them are zoological curiosities because of
their unusual modit ications ot structure or habit. Thus, the paiasitic bat
flies of the families Streblidae and Nycteribiidae are strange forms be¬
cause of their structural modifications alone. But combined with this
strangeness of form, the members of these two families and of the more or

less related family Hippoboscidae have the habit, rare among insects, of
retaining the larva within the body of the female where it is nourished by
glandular secretions until it is ready to pupate. The members oi the little
family Polyctenidae likewise retain the immature insect within the tody of
the female until it is partly grown, but in this case it is nourished
through what is functionally a placenta that is developed from the body of
the young itself. The extraordinary developments of both structure and
habit which occur among the mites offer an endless variety, the exploration
of which has scarcely more than begun.

Throughout the entire series of these ectoparasites there are phenomena
of special interest from the point of view of evolution. There are beau¬
tiful examples of what appears to be convergent evolution which has re¬
sulted in the development of similar habits and similar structures in quite
unrelated forms. There is the problem of perennial interest which has to
do with the occurrence of related parasites upon related hosts and the con¬
sequent suggestion that the relationships of the parasites may perhaps
throw some light upon the phylogeny of the hosts and, conversely, that the
relationships of the hosts may throw some light on those of the lice. This
will be discussed in detail in a section of this work.

Above all there is the circumstance that certain of these ectoparasites
are concerned with the transmission of disease. Most of what is known con¬
cerning this has to do with the transmission of disease in man, but it
seems clear that the same thing must occur in the case of many other ani¬
mals. Since numerous diseases of man have unquestionably come over to him
from other animals the whole picture of the epidemiology of these diseases
cannot be grasped until the background in the animals from which a particu¬
lar disease has come is understood. In many instances the occurrence of a
particular disease in man is to be regarded as purely an accident. This,
for example, seems to be true of "murine typhus, " which arises each time
independently from the background formed by certain rodents and their para¬
sites. One of the most illuminating instances of this sort is the recogni¬
tion — as late as 1947 — of the disease called "rickettsial pox, " a disease
that was previously unknown and was shown to have been transmitted to man
through the agency of mites from the ordinary house mouse, an animal that
had previously not been known to be connected with any disease of man.

We are not here attempting to recite the story of the relation of ecto-
parasitic insects to the transmission of disease. The intention is merely
to emphasize the principle that no blood-sucking arthropod, even if it
occurs only on a bat on some remote island, may be eliminated entirely from
the range of our interest. The time is at hand for the building up of
greater knowledge of blood-sucking ectoparasitic insects in all groups.
Almost none of these groups, except perhaps the fleas, has received the
attention which it deserves. And, as will appear in subsequent sections of
this volume, our knowledge of the little Order of the sucking lice is still
barely past its earliest stages.

CHAPTER II

The Morphology and Anatomy of the Anoplura

As has already been noted, the sucking lice are very peculiar forms.
They are commonly described as degenerate, but this is a word which has no
proper place in a biological vocabulary, since it carries with it certain
purely anthropomorphic connotations that involve a flavor of reproach. It
would be better to say merely that they are highly specialized for life in
the peculiar environment which they occupy. The evolutionary processes
which have been at work upon them have produced a large amount ot speciali¬
zation by loss and reduction of parts. There is not the slightest vestige
of wings. The eyes have been very greatly reduced, being scarcely or not
at all recognizable in the majority of species and consisting of a single
facet even when best developed. The tarsi have been reduced lrom live to
one or at the most two segments. The antennae have been reduced to at the
most five segments. In many species the sclerotization of the abdomen has
been almost completely lost. There has been a reduction of the number of
spiracles, no known species having more than one pair on the thorax and six
pairs on the abdomen. The structures which comprise the ovipositor of the
more generalized insects are reduced to mere vestiges.

Along with these reductions and losses there have gone some specializa¬
tions of a quite different type. Thus, the mouthparts of the sucking lice
are perhaps as highly specialized as are to be found anywhere among the in¬
sects and the claws have become highly developed in correlation with the
habit of clinging to hair.

The great degree of reduction and modification found in these insects
has led^to much misunderstanding concerning their morphology and this mis¬
understanding has centered especially about the mouthparts. Here, even
yet, we cannot give a final answer to all the questions involved although
we are undoubtedly somewhere near that answer.

In the preparation of the following section on morphology and anatomy,
Mr. Stojanovich has done all the laborious work of dissection and of pre¬
paring the illustrations. Parts of this material have already been pub¬
lished by him in a paper entitled "The Head and Mouthparts ol the_ Sucking
Lice," which appeared in "Microentomology , " Volume 10, Part 1 (194b).

There are certain minor points concerning the anatomy of the lice which
have not yet been adequately explored, but the lollowing account will cover
the major features, especially those which may at some time have a bearing
upon problems of taxonony.

The Head

Preliminary Statement

The most difficult problems connected with the morphology of the suck¬
ing lice are those which concern the mouthparts. The head itself is not
difficult to understand, but before entering into a discussion of its mor-
pholon- it may be well to review certain developments which have occurred
within" recent years in the study of the comparative morphology of the in¬
sects. These developments have resulted chiefly from a series of papers
which have appeared in the journal 'Microentomology," which is published from
the Natural History- Museum of Stanford University. A briel summary ol the
conclusions which have been derived from this work and which are pertinent
to an understanding of the Anopluran head follows.

There has long been difference of opinion among morphologists as to the
segmentation of the insect head, opinion as to the number of segments in¬
volved having varied from as low as four segments to as many as nine. Fur¬
thermore, there has been much disagreement as to the arrangement of these
segments and certain concepts that we believe to be based upon grievous
errors have become deeply imbedded in the literature of the comparative
morphology of the insects. Not until those errors have been removed is it
possible to arrive at any understanding of the morphology of a specialized
head such as that of the Anoplura.

The work that has been done in this laboratory has involved studies of
external structure, of musculature, and of the segmental distribution of
nerves. The study- of the nervous system of the head region has been carried
through a long series of forms ranging from the Oligochaete and Polychaete
worms to the insects and has demonstrated that a unified system of segmen¬
tation of this region exists throughout the Annulata. The following con¬
clusions have been derived from these studies.

1. The head of insects involves a total of six segments, these — in mor¬
phological order — being the labral, clypeal, ocular-antennal, mandibular,
maxillary, and labial segments.

2. The labral segment is segment one of the body' and the anterior por¬
tion of the alimentary canal, which is called the stomodaeum, is an invagi¬
nation of this segment. The ventral portions of this segment are much re¬
duced and seem to involve only the "hypopharyngeal suspensor plates, " upon
which the mouth angle retractor muscles insert.

The clypeus is the dorsal element of segment two and the hy-popharynx
is basically the ventral portion of this segment.

4. The hypopharynx is primarily a ventral lobe of segment two, but it is
possible that in some insects elements derived from the mandibular segment
may be involved in it.

5- The ocular-antennal segment bears the antennae, the ocelli, and the
compound eyes. It is commonly divided transversely by a suture, usually
called the epicranial, suture, which usually forms a Y-shaped line across
the head, with the lateral arms terminating on each side between the com¬
pound eye and the corresponding antenna. This suture is definitely intra-
segmental and is something more than a mere line of weakness along which
the derm splits at the times ol ecdysis. It is a line of very ancient ori¬
gin, the antecedents of which can be seen even in the Polychaete worms.
The area anterior to this suture and posterior to the clypeofrontal suture
may properly be called the frons.

No ventral, element ascribable to the ocular-antennal segment can be
identified in any insect thus far examined, although morphologically some
such element, however much reduced, should theoretically be present. No
muscles ascribable to this segment, other than those of the antennae and
cei tain evanescent muscles ol the ptilinum in flies, are known to exist.
The anterior tentorial pits apparently belong to this segment.

6. In cases where the mandibles have been lost the mandibular segment
may entirely disappear, at least as lar as any muscles and anyr sclerotized
elements are concerned.

7. The maxillary- segment is usually much reduced and in many insects is
entirely membranous. In many insects it forms no part of the sclerotized
head capsule. The posterior tentorial pits seem to belong to this segment.

,lhe Lablai segment is always much reduced dorsally and no insect is
known in which the dorsal wall of this segment is sclerotized and enters
into the composition of the sclerotized head capsule. The area called the
submentum, however, is commonly sclerotized and forms a portion of the
ventral or posterior-wall of the head capsule, although at times this
area may- he entirely membranous. The submentum is merely a portion of the
ventral wall ol segment six. The salivary glands belong to segment six

and the point of opening of the salivary duct belongs to this sequent and
is a definite landmark of this segment. It does not belong to the hypo
pharynx although it may become involved with this structure.

9. The distribution of segmental nerves to sensory structures and mus¬
cles is a positive indication of segmental homo Louies when all uerves are
present. At times nerves may be lost, if the structures which they serve
are Lost, and at times the clarity of segmental relations may be disturbed
by the fusion of nerves.

1U. The insertions of muscles are constant, but the origins may shill,
even from one segment to another, if the ectodermal areas upon which they
primitively originated are lost or displaced.

With these general conclusions in mind it is possible to proceed to the
problem of the organization of the Anoplurau head.

Of ail the species of Anoplura now known, those of the genus Pediculus
seem to present the least modification of the head. Therefore it is appro¬
priate to begin with the head of Pediculus.

The Head Capsule of Pediculus humauus
Figures 1, 2

In this head the anterior apex is more or less membranous and fox-ms a
slightly eversible protuberance, commonly called the haustellum, which sur¬
rounds the apparent mouth opening and is beset with small hooks. In this
membrane, on the dorsal side, there is a small, sclerotized plate which is
by all criteria the labrum. Posterior to the labriun is a large plate which
extends across the head in front of the antennae and is continued somewhat
to the ventral side. This plate is clearly the clypeus, since it bears the
origins of the cibarial muscles and the place of origin of these muscles is
constant in all insects that have been examined. This plate is separated
by a broad suture from the next plate, which makes up the greater part of
the dorsal wall of the head. The entire posterior portion of the head cap¬
sule, dorsally, is composed of the ocular-antennal segment. The antennae
are attached to the head at the anterior-lateral angles of this plate and
the eyes are placed on each side at about the middle of the lateral border.
The plate is divided by a V-shaped suture, the lateral terminations of
which are between the eyes and the antennae. The base of the V is connect¬
ed with the median coronal suture. The portion of the ocular-antennal seg¬
ment anterior to this V is the frons, while the areas posterior to the su¬
ture constitute the ocular lobes or, as they are sometimes called, the pa-
rietals.

The dorsal sclerotization of the ocular-antennal segment extends around
somewhat to the ventral side of the head. There are no dorsal areas that
can be assigned definitely to the gnathal segments.

Except for the reduction of the dorsal elements there is nothing in any
way peculiar thus far about this head capsule.

On the ventral side of the head the areas not occupied by the ventral
extensions of the dorsal sclerotization are entirely membranous and show-
no indications of segmentation.

The tentorium, as in all Anoplura, is lacking and there are consequently

no tentorial pits. , . , , . . . ,

The mouthparts are entirely retracted within the head and will be con¬
sidered later.

The Head Capsule of Haematopinus suis
Figure 3

The head capsule of this species is considered because it presents cer¬
tain features which would not be readily understood on the basis ol what

<1)

"ro

O 3

"ii

■a

Head structures as shown in Pediculus hurnanus Linnaeus

Figure 1

clypeus^

grig in of labial compressor muscle

bauslelhim

labium

origins of
cibarial muscles

or if/ in of lateral
pharyngeal muscle

WV >

. N

\ \

origins of
antennal
muscles**'

origin of
mandibular

muscle - /

clypeo frontal sul are

origin of dorsal labral
retractor

prigins of
_ >/ pharyngeal
muscles

1st anlennal
segment

rye

origin of formal muscle

antennal

segment

coronal sutur /

postfrontal suture

ocular lobe

Origins of muscles in head of Pediculus humanus Linnaeus

Figure 2

Structures and origins of muscles in head of Haematopinus suis (Linnaeus)

Figure 3

is found in Pedlculus.

It will be noted that in this head there is no external indication of
eyes. However, the ocular nerves are present and end in a small spot, just
at the base of the areas designated in the illustration as the oculai
points, which are morphologically the eyes. Webb has demonstrated that in
some specimens, at least, definite but small lenses can be recognized.

In comparison with the head of Pedicul us i t wi 11 be noted that one transr-
verse suture is missing. On the basis ol all the available evidence, in¬
cluding the study of other species not here considered, we interpret the
conditions as follows. We assume that the clypeus has been produced pos¬
teriorly until it has crowded out the frous, and the frontoclypeal and
postfrontal sutures have become confluent. Therefore nothing is lelt ol
the Irons except the small areas at each side which bear the antennae. The
transverse, V- or U-shaped suture is therefore morphologically the post¬
frontal + clypeo frontal suture. This conclusion is supported by the posi¬
tions of the origins of the cibarial muscles.

One other feature of this head calls for mention.

At the posterior border of the head are two invagiuated sclerotizations,
the occipital apophyses. To these attach muscles which serve to retract
or elevate the head. Evidence derived from the innervation ol these mus¬
cles indicates that they and the structures upon which they insert belong
to the head. To just what segment of the head they belong remains to be

determined. „ .. , , .,

The ventral side of the head is so similar to that ol Pedlculus that it

is not here illustrated.

The Head Capsule of Other Anoplura

There are some modifications ol the head capsule in other species ol
Anoplura, but they can all be accounted for on the basis of the two species
which have been discussed. Consequently, we need belabor this subject no

further.

The Mouthparts and Internal Structures

As has already been noted, the mouthparts ol the sucking lice are ol a
very peculiar type which is not found elsewhere in the insects. Function¬
ally they are to be described as "piercing and sucking, but their mecha¬
nism is very different from anything that is to be lound in other piercing
and sucking forms such as the Hemiptera and the blood-sucking (lies, lhey
have been the subject of much controversy and there are still certain de
tails which remain doubtful and which will require emhiyological work lor

their final elucidation. ... ,, . , ,

Since, apart from the space occupied by the brain, the internal struc

tures of the head are almost exclusively associated with the mechanism ol
feeding we may consider all these structures together.

The°first necessary step is that of forming a clear picture ol the re l a
tionship of the piercing structures and the food channel. It must be
grasped at the very beginning that the piercing structures do not in any
way form the food channel. In order to understand this clearly, reference
maV be made to Figure 4. In Part A of this figure is shown a longitudinal
section through a normal insectan head, with the lateral parts such as man
dibles and maxillae omitted. The roof of the oral chamber or cibanum is
here formed by a continuation of the ventral wall of the labrum and the
floor of the chamber by the hypopharynx and the anterior wall ol the labium.
Leavincr aside any argument as to where the true mouth opening actually is
we ma\B for the present consider it to be the opening into the pharynx.
Note especially the opening of the salivary duct between the hypopharynx

Diagram of mouthparts

Figure 4

ami the labium.

In Part B oi' this figure is shown a similar diagram of the head of a
sucking louse. Note that the upper portion of the head remains unchanged
and the modifications involved are all in the lower — morphologically the
posterior — portion. It is evident that the parts involved in the piercing
structures have been pulled back into a sac that lies entii'ely beneath the
mouth. The three needles which form the piercing instrument originate from
the base of this sac. When the instrument is brought into action it is ex¬
truded through a small opening in the lloor ol the oral chambei jusi in
from the apex of the head. It is very' evident that the blood of the host
does not pass into this sac on its way into the alimentary canal. The ac¬
tual arrangement of the sac and its relation to other parts is shown in
less diagrammatic form iu Figure ?•

The questions at issue have to do mostly with the homologies ot the
three stylets which comprise the piercing instrument. In the diagrammatic
illustration they are greatly exaggerated in thickness. Actually, they are
exceedingly delicate and are closely appressed together. However, the
questions concerning these stylets are not all there is to the problem,
since there are certain other structures which do not appear in this dia
irram and which have to be explained.

For our explanation it is necessary to turn to the musculature and the

innervations of the muscles. .

Perhaps the most favorable subject for study ol these problems is the
hoj louse, Haematopinus suis. This is a rather large species and it shows
ail the structures. It may be noted that all the work on which the accom¬
panying illustrations were based was done by the method ol dissection em
ploying powers up to xll5 of the binocular dissecting microscope.

In Figure 6 are shown the structures which appear upon removal ol the
dorsal wall of the head. We may in passing note the muscles ol the anten¬
nae, which originate upon the dorsal wall. In insects which possess a ten¬
torium they would originate upon that structure, but since the tentorium is
here lacking the origins must of necessity shift to another position,
some Anoplura they originate in part upon the lateral wall ol the head ven

trad of the antennae. , „ ,

The "brain" is relatively large and is composed of the supraoesophageai
and suboesophageal ganglia* which are connected by the very short ^circuni-
oesopha<real connectives that pass around the oesophagus (see Figure -h •
One landmark to be especially noted is the frontal ganglion, which in all
insects lies in front of the brain and between the ci banal and pharyngeal
muscle bundles. Just posterior to this frontal ganglion the alimentary
canal swells into a bulb which is the pharynx and which forms the sucking
pump From the pharynx large bundles of muscles extend to the dorsal wa
o7the Lad. It is 'these muscles which function to

cause it to operate as a pump in sucking up the blood ot the host. There
is nothing in any way unusual about them or their arrangement.

Anterior to the frontal Ganglion are several muscle bundles originating
upon the cUeus anT inserting4 upon the dorsal -all of the oral chamber
These are the cibarial muscles, as is confirmed by their innervations whic

WllAnteriorirStodthestr muscle bundles the dorsal wall of the mouth chamber
is formed by a sclerotized plate, somewhat in the form of an inverted Y.
From the apex of each arm a muscle extends forward to a little, external
p[a?eihich we have previously identified as the labrum. These muscles may
be regarded as the compressors of the labrum and are essentially e sam
as are to be found in most insects. The plate to which they insert, and
which forms the dorsal wall of the chamber, is here called the palatal
date It is morphologically equivalent to the plates which are to be found
t the dorsal wa?l of the oral chamber, in one form or another, in a wide

variety of insects.

There is nothing here that is in any way unusual or markedly different
from what is to he seen in any typical insect.

Disregard for the moment the pair of structures which lie one on each
side of the palatal plate. They present one of the most obscure problems
involved in the mouthparts of these insects and we shall discuss them later.

Look again at Figure 5 in which the relations of the various structures
are shown and note the position of the ventral trophic sac and the point at
which it joins the cibarium.

In Figure 8 the ventral wall of the head has been dissected away -to re¬
veal the trophic sac and its associated muscles. The piercing stylets are
concealed within this sac and only the branching ends of the apodemes upon
which the muscles of the stylets insert, and which are formed from the
walls ot the sac, appear at the posterior extremity.

Note that from these apodemes slender muscles extend forward to attach
to a sclerotized transverse band in the ventral wall of the sac at about
the midpoint of its length. Also, certain muscles extend from the apodemes
and attach to the obturaculuin, a structure which will be discussed later,
and there is a relatively huge, transverse muscle at the base of the troph¬
ic sac, this originating at each of its ends upon the obturaculum. Aloncr
the trophic sac are inserted certain slender muscles which originate upon
the outer head wall.

In Figure 9 the base of the trophic sac is shown as with the walls cut
away to reveal the apodemes of the stylets which are contained within the
sac. It must be emphasized that the stylets are outgrowths from the wall
ot the sac and so are these apodemes, the sac being closed apically. There
are no muscles inside the sac. The sac is an invagination of the body wall
and anything inside it is morphologically outside the body. We may note
in passing that certain illustrations of these structures which have been
published show the apodemes as being inside the sac and thus in effect in¬
dicate an arrangement that is a morphological impossibility.

Let us now turn our attention to the stylets themselves. There are
three of them— a dorsal, a middle, and a ventral. Th-e dorsal and ventral
stylets are flattened and relatively broad; the central stylet is cylindri-
cal and very delicate. The three are very closely appressed together and
extend forward so far that when at rest their anterior ends lie just within
the mouth chamber. They enter the chamber through a slit in its floor If
they are extended to reach a blood vessel the base of the sac must be pushed
tar forward to lie close to the apex of the head.

The homologies of these stylets have been much disputed.

The nature of the median stylet is, however, quite clear. It carries
the extension of the salivary duct and except for the lumen of the duct is
solid. There is no reason whatsoever to consider it as anything more than
a greatly elongated papilla which arises from about the mouth of the sali-

1J' 1S bere cailed the salivary stylet. It has been consid-
ered to be the hypopharynx, but only because of a complete misunderstand¬
ing which has been widely spread among morphologists. The salivary duct
does not belong to the hypopharynx and the opening of this duct defines a
point that belongs to the labium.

Nor can there be much doubt concerning the ventral stylet. It is un¬
questionably derived from the labium and its muscles are innervated by
nerves that seen, clearly to be labial. Presumably the stylet is formed
lorn the terminal portion of the labium, the part generally called the lig-

vUn li \hifre n°, i£bial palp1' To understand the structure one mult

h?-iCn re ablUm aS bein^ puiied back into the head, part of it
forming the floor of the consequent invagination and part of it bein<r free

and extendmg forward to form the tree stylet. We consider that the “floor
the trophic sac is lormed from the labium as far forward as the point

indicated as the "limes labial is," or boundary of the labium in Figure 8.

There now remains the question ol the identity ol the dorsal stylet. In
the earlier work upon which this presentation is based this stylet was con
sidered to be the hypopharynx. At that time, however, the innervations had
not yet been studied. In the light ol further studies we have been forced
to conclude that this stylet is actually formed from the fused maxillae.
The reasons for this decision will be discussed when we consider the ner¬
vous system of the head.

The hypopharynx here, on the basis of evidence presented by the nerves,
is represented by the dorsal wall of the trophic sac and the floor of the
oral chamber as far posteriorly as the opening into the pharynx. Muscles
inserting upon this area are innervated by nerves which can be nothing but
nerves of the hypopharynx.

We are Left now with the problem of the pair of structures lying inside
the head just in from the apex, one on each side of the palatal plate and
to which we referred earlier.

In our previous work we considered these structures to be vestiges ol
the maxilLae, but in the Light of the information now available this opin¬
ion can not be maintained. The maxillae have already been accounted for
and there remain but two possibilities in regard to the structures in ques¬
tion. It wouLd seem that they must be either completely new structures with
no antecedents in other insects or that they must be derived from some pre¬
existing movable structure and the only such structures that exist in this
part of the head of any insects are either the mandibles or certain develop¬
ments from the dorsal wall of the cibarium to which the name "premandibles, "
or the better name "inessores," have been applied. We discard recourse to
the first suggestion— that they are completely new structures— as a matter
of principle and concentrate upon the thought that they are derived 1 rom
some pre-existing movable structure provided with muscles and nerves.

First let us°describe them as they occur in Haematopinus (Figure 10).
Here they consist of a pair of somewhat wedge-shaped, sclerotized bodies,
lying within the head cavity, one on each side of the palatal plate. Each
of them is connected by a strong and undoubtedly chitinous ligament to the
head wall just at the margin of the labrum and each is also connected by a
slighter ligament to the margin of the palatal plate. Each is provided
with one very large muscle that originates far back on the lateral wall oj
the ocular lobe of the head and inserts upon the doubtful structure bv waj
of a long and strong apodeme. A much smaller muscle originates upon the
sclerotized wall of the pharynx. Both these muscles are innervated by
branches of a nerve, which is one of the first pair to arise from the sub-
oesophageal ganglion (Figure 2P0 .

In the light of this innervation we are forced to conclude that the
structures in question are actually the mandibles. They cannot have been
derived from such structures as the messores, which are developed from the
dorsal wall of the cibarium and whose muscles are innervated by branches
from the labral nerve.

How these mandibles have become withdrawn into the body and how they
have acquired two connections to the body wall are problems which can only
be solved by careful and detailed embryological studies that are still to
be made. But that no interpretation other than that here presented is con¬
sistent with their innervations is clear.

These structures vary somewhat in form and position in different groups
of the lice, in some species lying close to the ventral side of the head
(Figure 11) and close together, but they seem to retain the same connec¬
tions to the head wall and to the palatal plate and to retain also the same
musculature. It is possible that a minutely detailed study ol them would
offer some evidence that would have a bearing upon problems of relation¬
ships within the group but at the present time this seems impracticable.

Structures Associated With the Mouthparts
Pawlowsky's Glands and Salivary Glands
Figures 5. 6, 7, 9, 10, 11

These are a pair of small, glandular structures each of which opens
through a short duct into the trophic sac. Their function is unknown, as
are their homologies. They are innervated hy branches of what we here con¬
sider to be the hypopharyngeal nerves. There exists very little informa¬
tion — and that mostly very vague — concerning glands of the head region
other than the salivary glands and at present we can make no statement con¬
cerning Pawlowsky's glands other than to record their position and their
innervation. We have suspected that they may be homologous with what have
been called "pharyngeal glands" in other insects, but an extended study
would be necessary to determine whether or not this is the case.

In those lice which have been investigated in regard to the matter there
are two pairs of salivary or labial glands. These lie in the thorax, dor¬
sad of the alimentary canal. In Pediculus humanus the members of one pair
are small and oval or somewhat kidney-shaped, those of the other pair are
tubular and folded. From each gland a duct extends forward and unites with
the duct from the other type of gland on the same side of the body. These
two ducts then unite into the common duct near the point of their entry
into the head. This common duct then discharges at the apex of the median,
salivary stylet which has been described.

The Obturaculum
Figures 7, 8, 12

The mouthparts themselves are not the only peculiar feature of the head
in these insects. We have referred earlier to a structure which we have
called the "obturaculum, " literally the plug or cork, which represents a
most peculiar development that, as far as at present known, occurs only in
this Order.

The extreme posterior portion of the head is filled by a mass of tissue
which in histological preparations has the appearance of being minutely fi¬
brous and which may be regarded as connective tissue. This tissue forms a
plug, or a transverse diaphragm, which fills the occipital foramen and ex¬
tends into the thorax. There it more or less completely envelopes the
ganglia which are concentrated in that region. The anterior end of the
plug is concave and cup-like. Through this plug the alimentary' canal, the
tracheae, the salivary duct, and the tracheae enter the head in the manner
of tubes through the cork of a bottle.

The base of the trophic sac extends into the concavity of the anterior
end of the obturaculum and certain of the muscles which have their inser¬
tions on the apodemes of the trophic stylets originate upon the walls of
the cup. Thus, the large, transverse labial muscle, which inserts upon the
ventral surface of the trophic sac, originates in this manner as do the
folded maxillary retractors and the labial retractors. The importance of
this will be apparent upon reference to the illustration of the head of
Llnoinathus vltuli (Figure 12). Here it will be seen that the trophic sac
extends Jar luck into the thorax. Upon what would its muscles originate if
the obturaculum were not present? This mass of connective tissue is evi¬
dently a part of the solution of the problem of functioning which is pre¬
sented hy the peculiar trophic mechanism and its significance in connection
with this will be discussed later.

The origin and homologies ot the obturaculum are at present quite in
doubt. It seems evidently to be a mass ot connective tissue, but at pres¬
ent there is very little information concerning the occurrence of such tis¬
sue in insects. Other work at present in progress has shown that a sheath

cibarium and cibarial muscles pharynx ami pharyngeal muscles supraoesophageal ganglion

Internal structures of head in Haematopinus suis

Figure 5

labial retractor muscle' folded labial retractor muscle

labrum

mandibular ligament ^

haustellum

^ palatal plate

eibarium _

labral

compressor muscle

cibarial muscles ^rsSS

1st antennal
segments _

mandibular vestige

japodeme of
mandibular

muscle

- mandibular muscle

Jtgpo pharyngeal
suspensor

-frontal ganglion

trophic sac

tormal muscle _

pharynx^- -zZZ."- _ _

mouth angle
retractor muscle- --

pharyngeal muscles

mandibular

muscles -

supraoesophageal

ganglion-

hypopharyngeal

stator muscle---"' '

transverse labial muscle-

labial

protractor muscle-''

folded maxillary
retractor muscle

antennal muscles

- ocular point

__ op//c nerve

—» — '

- ... oesophagus

suboesophageal
--ganglion

_ obluraculum

\
I
I

origins of transverse J
~ - ^ labial muscle i

_ thoracic mall !

salivary duct

Structures in the head of Haematopinus suis, dorsal aspect

Figure 6

manditnilar ligament

labrum

huustellum

palatal plat?

ibarium

_ apodeme of

mandibular

muscle

mandibular muscle

,1st antennal
segment

- ocular point

."^clypeofrontal suture

Jormal muscle
posterior
hypopharyngeal
muscle

_ labial adductor

muscle

_ obluraculum

r origins of transverse
labial muscle

thoracic mall

salivary duel

mandibular vestige

pharyngeal
opening ---

Paivlowsky’s glands'

mandibular

muscles ----

hypopharyngeal

stator muscle -

trophic sac -

labial protractor muscles .

(- flexor of glossa
paraglossa )

transverse
labial muscle

folded

maxillary

retractor muscle**'

labral

compressor muscle -■

ci burial muscles ^

Structures in the head of Haematopinus suis, pharynx and brain removed

Figure 7

mandibular ligament ^

labrum

labial compressor muscle l

man dib u lar nest ige

apodeme of
mandibular
muscle.

pharynx _ __

ocular point.

formal muscle _

mandibular

muscled-

hypo pharyngeal
stator muscle _

obturaculum _

labial protractor muscle
(-flexor of glossa and
paraglossa )

folded labial
retractor muscle-^----

haustellunj

, ventral fold

trophic sa

- -^palatal plate

~ --- mandibular muscle

1st antennal

''•--.segment

pharyngeal muscles

-- xientral pharyngeal
muscle

'--ventral labial
adductor muscle

- transverse labial muscle

---- - maxillary

protractor muscle

Structures in the head of Haematopinus suis, ventral aspect

Figure 8

maxillary apodeme-

folded maxillary

retractor muscle<

salivary bulb -

a- —

maxillary stylet- -

labial protractor muscles
( -flexor of y l ossa and
paraglossa )

labial adductor muscle

salivary styleL _

_ Jabial apodeme

salivary stylet

\transverse labial
muscle

W '•^maxillary

protractor muscle

_ J\y popharyngeal
surface

anterior by popharyngeal

dimes labialis

labial stylet -

Base of trophic sac in Haematopinus suis

_ maxillary stylet
Figure 9

haustellum ^

labrum

origin of labial
compressor muscle .

palatal plate _

apodeme of
mandibular
muscle

cibariunv'

mandibular ligament

,■ mandibular nest ige

mandibular

origins

''mandibular muscle

A. dorsal aspect

haustellum^. .

.labrum

labral compressor muscle ^

palatal plate _

apodeme of
mandibular

muscle _

trophic sac opening -

/ nandibular ligament

jnandibular vestige

mandibular

origins

'mandibular muscle

B. ventral aspect

The mandibles in Haematopinus suis

Figure 10

haustellum „

(interior mandibular ligament

limes
labial is

anterior hypopharyngeal stator muscle

labrum -

trophic sac opening _

jnaiulibular vestige

dorsal labral
/ retractor muscle

labral compressor

muscles ^

posterior

mandibular

lifjament

hypo pharyngeal
plate ,

trophic sac.

labial

retailor

muscles-

mandilnilar

muscle

Paudowsky's gland'

labial
adductor
muscle - '

labial protractor muscle '

(= flexor of glossa and paraglossa )

- palatal plate

posterior
mandibular
ligament

Cibarial region in Pediculus humanus, trophic sac removed

Figure 11

Inbrnm _

cibarium _

palatal plate

Jabral compressor muscle

--^dorsal labial retractor muscle

2nd coxa

.trophic sac

folded maxillari/+ labial /
retractors muscle

| cibarial muscles

frontal
ganglioir

formal muscle^

pharynx _

mouth angle
retractor muscle

lateral
pharyngeal
muscle

1st coxa

1st antennal segment

- Jabial protractor muscles
( = flexor of glossa and
^ paraglossa )

■x

maxillary
protractor muscle

' -dransncrsc labial muscle

~ - — obturaculum

supraoeso phageal
ganglion

oesophagus

suboesophageal

ganglion

obturaculum

antennal

muscles

hypo pharyngeal
suspensor

pharyngeal muscles

Structures in the head of Linognathus vituli (Linnaeus)

Figure 12

of connective tissue which envelopes the thoracic ganglia and extends into
the head occurs in a cockroach and offers the material from which the ob¬
turacuium could have been derived.

The Functioning of the Mouthparts

Some idea of how the trophic mechanism may function can be derived from
contemplating the arrangements that have been described.

It seems probable, from purely mechanical considerations, that the slen¬
der muscles which extend forward from the branches of the maxillary and
labial apodemes and attach to the walls of the trophic sac must act to pull
the stylets forward, but it seems improbable that they can furnish the en¬
tire drive necessary for the complete exsertion of the stylets. We may
therefore sumiise that the action of these muscles is aided by blood pres¬
sure. That pressure might very well be developed by the conti*action ot the
powerful dorso-ventral muscles of the abdomen. The pressures thus devel¬
oped would be transmitted through the obturacuium to the head cavity and
thus brought to bear upon the trophic sac.

In any case, however the necessary force is developed, the stylets are
pushed out of the trophic sac and forced into the tissues of the host. It
is evident from the structure of the stylet bundle that the blood does not
pass through any channel formed by the stylets themselves. II it did so it
would merely pass into the trophic sac itself and not into the pharynx. We
may assume, therefore, that the stylets merely serve to pierce a blood ves¬
sel. The membranous area at the apex of the head can be unrolled and ap-
pressed closely to the skin of the host. It seems probable that the blood
is drawn up around the stylet bundle and into the mouth by the action ol
the pump formed by the pharynx.

The retraction of the stylets offers some interesting mechanical prob¬
lems. Presumably the muscles which accomplish this are those indicated in
the illustrations as the folded retractors ot the labium and maxillae.
These muscles when at rest are actually folded upon themselves and raise
the question of how such a folded muscle can function. This may possibly
be accounted for in the following way.

It seems probable that in forms such as Linognathus vttuli, where the
trophic sac extends far back into the thorax, a rather peculiar muscle
wou Ld be required to permit of the extreme degree ot relaxation and exten¬
sion that would be necessary when the stylets are fully extended and that
this muscle would have to be extremely long. We suspect that when the mus¬
cles are extended they are under little or no strain. Note that these mus¬
cles originate on the" obturacuium which is composed of a tissue that seems
to be extremely elastic, as can be seen in dissections of fresh material.
We may suppose then that when the stylets are to be withdrawn the retrac¬
tor muscles give merely an initial pull and that the completion ol the re¬
traction of the parts is accomplished by the elasticity of the obturacuium.
As the retraction comes to completion the retractor muscles simply lold
upon themselves. We see no other way of accounting lor the mechanics of
such a peculiar arrangement.

The Antennae

The antennae are normally five-segmented and no species of sucking louse
is known that has more segments than this. In a few species there is ap¬
parently a fusion of the two terminal segments, leading to an apparent
four-segmented condition. In one genus, Haematopinoid.es, the antennae are
definitely four-segmented. In a few species the three terminal segments
are but weakly separated from each other and the antennae have the appear¬
ance of being three-segmented.

Each of the two terminal segments usually bears upon its posterior bor¬
der a ring-like structure that may be regarded as a sensorium, but in some
forms these are lacking.

No especially peculiar developments of the antennae are known. In some
forms there is a slight sexual dimorphism which involves segment three in
the male. This segment may have its apical, preaxial angle somewhat pro¬
longed and this prolongation may terminate in a recurved spine or a short,
stout seta or some other modification of setae may occur. In some cases a
modified seta may here be present unaccompanied by any other modification.
No instance is known where the antennae of the male are developed into such
extreme clasping structures as occur at times in the Mallophaga.

The Eyes

Definitely developed and clearly recognizable eyes are present in only a
few species of the sucking lice. In these species each eye possesses a
single lens, beneath which is a pigment spot. The presence of such eyes
has been accepted as a taxonomic character and has been much used in the
definition of genera.

Webb in a recent paper has shown definitely the existence of eye ves¬
tiges in Haematopinus suis, each eye consisting of a very small and obscure
lens, "beneath which there are two or possibly four ovoid crystalline bodies
surmounting a cluster of pigmented sensory cells ending in nerve fibres."
He has identified in all probability quite correctly — what he considered
to be eye vestiges in a considerable number of species of Haematopinus ,
Linoinathus, Solenopotes, Ratemia, Prolinoinathus, Hybophthirus, and Ant-
arc tophthirus. They are definitely present in Pedtculus, Phthirus, Ped-
icinus, Microthoracius, and Pecaroecus.

Unfortunately, the great majority of all the species of sucking lice are
known only from preparations made from dried specimens and in the°course of
making these preparations any spots of eye pigment are destroyed and the
lenses themselves at times become difficult or even impossible to recognize
and to differentiate from any small, clear spot. For this reason the ex¬
istence of eyes cannot at present definitely be demonstrated in many forms
where they may actually exist. For this the study of properly preserved
material must be awaited.

It is clear, however, that the presence of recognizable eyes cannot form
a sound basis for any system of classification within the Order Anoplura,
although in some instances it may offer good key characters for identifica¬
tion.

Webb has referred to these eyes as "ocelli," but their position indi¬
cates clearly that they are the morphological homologues of the compound
eyes of other insects and not of the structures for which the term ocelli
is ordinarily employed. Morphologically, they are vestigial compound eyes.

The Thorax
Figures 13, 14

In association with the absence of wings the thorax is very much re¬
duced. While three separate segments are definitely present they are
closely fused and certain marked modifications have taken place. The meta¬
thorax is the most reduced of the three segments and may be represented
only by small lateral elements. Only one pair of spiracles is present
this belonging to the mesothorax.

An understanding 0f the structure of the thorax in these insects rests
upon the recognition of certain landmarks. In the vast majority of insects

* Webb, J. E.

o logical Society

1948. Eyes In the Siphuncul ata.

of London, Volume 118, Part III, p

Proceed

age 575.

i n

the 7o-

prothoracic pleural apophysis and pit

mesonotum

natal pit

pi r aide

pronotuin

Structures of the thorax

Figure 13

occipital apophysis of head

prothoracic pleural apophysis,

' pit and phragma

coxae prothoracic sternal apophysis and pit

B. ventral aspect

Structures of the thorax in Haematopinus suis

Figure 14

there occurs on each side, on the so-called pleurite of the prothorax, an
invaginated tubular internal process called the prothoracic pleural apophy¬
sis. The external opening of this apophysis lies in a fold or sclerotized
stiffening rib that is known as the pleural fold or pleural ridge. The
latter is best characterized as the prothoracic pleural phragma. The point
of origin of this apophysis in the body wail is a precise landmark which
definitely defines the "pleurite" — morphologically the subcoxa — of this
segment. A similar phragma, but without such a deliuite apophysis, occurs
ou^each of the other two segments. The principal coxal articulation in all
insects is with the apex of this phragma.

On the basis of the landmark afforded by the prothoracic pleural apophy¬
sis and of other minor indications it is clear that the apparent notum ol
the thorax in the Anoplura is composed for the most part ol the so-called
pleurites, morphologically the subcoxae. The true notum, at the very most,
includes merely a narrow median area, usually unsclerotized, and a lateral
extension of this area on each side which includes the spiracle. In some
species the notum is reduced almost completely to a median notal pit, the
subcoxae having fused with each other across the mid-line of the dorsum,
even to the extent of enveloping and isolating the notal element about the
spiracles.

Perhaps the nearest approach to a normal thorax that is to be seen in
any sucking louse is that found in the genus Solenopotes (ligure 1JA) .
Here the pronotum is reduced merely to a median told, but the mesonotum is
of considerable extent in its anterior portion, although its posterior por¬
tion forms merely a median pit which lies in the subcoxai phragma. In
Haematopinus suis (Figure 14) the notum is reduced to a slight median line
throughout, with a very large median pit which is entirely enclosed by the
mesothoracic subcoxai phragma, and the spiracles seem to form isolated is¬
lands of notal origin. In Pediculus humanus (Figure 13B) there is a some¬
what broader notal "area left, this being membranous and enclosing the notal
pit, but the spiracles are even more completely isolated, being surrounded
by a sclerotized area of the mesothoracic subcoxa.

Apart from the modifications ot the mouthparts this reduction oi almost
complete obliteration of the notal areas ol the thorax is perhaps the most
characteristic feature in the morphology of the sucking lice.

The ventral side of the thorax is always quite simple. Nothing at all
remains of the primitive ventral arcs of the subcoxae and the whole ventral
area normally is membranous except for a single, median, sternal plate, even
this not bein'1, present in some species. It is not clear whether this plate
belongs to any one of the segments or is a fusion product derived from all
of them. In a few species, such as in the genus Raematopinus , it is clear
that the plate belongs at least in part to the prothorax, a pair ol sternal
apophyses being present which fuse with the prothoracic pleural apophyses
in a manner like that found in many other insects. These sternal apophy
ses are present in only a relatively small number of the sucking lice. .

In many Anoplura the sternal plate becomes free from the body around its
margins, especially at its posterior end. In some species it is lacking
and^in a few it is merely an irregular, sclerotized area.

The Legs

The principal modifications of the legs are found in the tibiae and tar¬
si and are connected with the grasping of the hairs of the host. There is
a sin 'le tarsal claw on each leg, except in four species. In these four
suedes the anterior tarsus bears a short, somewhat hooked structure which
arises at the side of the usual claw. This structure has the appearance ol
a claw but there is difficulty in being certain about its true nature and
the opinion is here held that it is not actually movable.

The tarsus is at the most but weakly two- segmented and even this appear¬
ance of segmentation is usually obliterated. In almost all forms the tar¬
sus is closely joined to the tibia and is but doubtfully movable upon the
tibia. The tibia is usually more or less produced at its morphologically
ventral apical angle, a thumb-like process being formed that opposes the
claw and thus participates in grasping the hairs of the host.

Usually the anterior legs are smaller than the others and have a slender
claw. In the majority of forms the middle legs are somewhat larger than
the first, with stouter claw, and in most species the posterior legs are
much broadened and flattened, with a stout claw. There are, however, var¬
ious combinations of these characters and in a few species the legs are all
of essentially the same size and form.

There is never anything that can be called a pulvillus, but frequently
there is a small, membranous lobe or blade-like process on the inner margin
of the tarsus. In Haematopinus there is a peculiar, sclerotized plate in
the "palm" at the base of the tarsus.

Certain very peculiar developments found only in the genus Euhaematopi-
nus will be discussed in connection with that genus.

The Abdomen

The primitive number of segments in the abdomen of insects is here con¬
sidered to be eleven. The anus is borne upon the eleventh segment, the
gonopore of the female upon the eighth (usually appearing as between the
eighth and the ninth) and the gonopore of the male upon the tenth. If the
full complement of abdominal spiracles is present there are eight pairs,
borne upon segments one to eight. There is a rather general tendency for
the number of abdominal segments to be reduced by the reduction or obliter¬
ation of segments either at the anterior or posterior ends of the body or
at times in both. In order to determine the homologies of segmental struc¬
tures in any insect it is necessary to rely closely upon established land¬
marks. Of these landmarks the gonopore in either sex may be regarded as
the most useful. In the Anoplura the spiracles are of but little "aid since
no known species has more than six pairs.

Among the Anoplura the genus Hoplopleura seems most nearly to retain
evidence of the full complement of abdominal segments. Here the paratergal
plates, which will be discussed later, are present on eight segments and
the eighth of these segments bears the gonopore in the female and also
bears a pair of gonopods. The ninth segment is well developed, but the
tenth and eleventh are reduced to mere membranous areas around the anus.
The first segment is evidently much reduced, but can still be distinguished.
Since in this genus all the evidence is harmonious and consistent we need
have no hesitation in accepting it. We may therefore take the position of
the vulva in the female, which lies between the gonopods of segment eight,
as the initial point for all reckonings of segmentation of the abdomen.

In some forms, such as the genus li nognathus, the abdomen is membranous
almost throughout except for the tergum of segment nine and certain sclero—
tized areas on the ventral side in association with the genitalia. In oth¬
ers, such as Haematopinus, the derm of the dorsum is slightly sclerotized
almost throughout with small plates of denser sclerotization and with
sclerotized paratergal areas along the lateral margins. But in the major-
ity of known species, the abdominal segments, both dorsally and ventrally
bear distinct, segmentally arranged, sclerotized plates in addition to the
paratergal sclerotizations. These tergal and sternal plates are usually
associated each with a row of setae and the arrangement of the plates and
rows of setae is to some degree distinctive of the various genera. Thus,
in l olyp lax the lemale bears two plates and two rows of setae on most of
the segments both dorsally and ventrally, while in Hoplopleura there are

aonnaLly three plates and three rows of setae on most of the segments.
There are some species in which the plates are entirely lacking, but the
rows of setae remain. There is a certain measure of sexual dimorphism in
regard to the numbers of the plates and rows of setae in certain of the
genera.

In the majority of known species there is at the lateral margin of each
of at least some of the abdominal segments a distinct, sclerotized plate
which bears or is at least closely associated with the spiracle, although
these plates may be present on segments on which the spiracles are lacking.
In earlier literature these plates were characterized as "pleural plates, "
but since the spiracles may be regarded as belonging morphologically to the
dorsum the term "paratergal plates" or the term "iaterotergites" is prefer¬
able. They are here called paratergites.

The number and form of these plates is of very great importance for the
recognition of both genera and species. In some genera, such as Hoplo-
pleura, Schizophthirus , and Pterophthirus, they form an overlapping series
along each side of the abdomen, with their posterior margins free from the
body and at times extraordinarily prolonged.

The External Genitalia of the Female

As already pointed out, the gonopore of the female belongs to the ven¬
ter of the eighth abdomiual segment. Morphologically, the gonopore and the
vulva are not necessarily the same thing, since the gonopore may be re¬
tracted into the body with the consequent formation of a genital chamber
or vagina into which surrounding elements may be withdrawn, even to the ex¬
tent of involving the venter of one or more segments, however, in the Ano-
plura there seem to be no complications involved which disturb the appar¬
ently normal segmental arrangements.

The sternal plate of segment eight, lying anteriorly to the transverse
fold which forms the vulva, is always or almost always in some degree
sclerotized to afford support for muscle origins and this sclerotization at
times involves also the venter of segment seven. These sclerotizations,
which are usually in some degree united if more than one segment is in¬
volved, collectively form what is here called the genital plate.

The’gonopods, which represent the primitive paired segmental appendages
of semnent eight, are practically always indicated at least by the presence
of a cluster of setae on each side at the end ol the vulvar fold. In the
majority of species they are definitely represented by a pair of flattened,
apically free lobes which at times are of considerable size. The mesal
margin of each gonopod is continued into the fold that torms the lip or an¬
terior margin of the vulva and this lip is at times beset with a fringe ol
delicate processes or is at other times simple.

Just within the vulva is the vaginal chamber. In some species the dor¬
sal wall of this chamber is slightly sclerotized. Into this dorsal wall
opens the duct of the spermatheca, if the spermatheca is present. In some
species the spermatheca seems not to be developed, as in Pediculus and
Haematopinus. It is very strongly developed in Pthirus and in at least
most of the species of Lino^nathus and occasionally in other torms. in
some instances where it appears to be lacking this may be occasioned merely
bv its weak sclerotization or perhaps even by its destruction in the making
of the preparation. It is a structure which calls for further investiga

U°The spermatheca itself, at its best development, is a swollen, sclero¬
tized structure at the inner end of the sclerotized spermathecal due lg
ure 124) and there seems never to be more than one. Usually, the openin
of the duct through the wall of the vagina is surrounded by a small area ol
dense sclerotization, if the duct is present.

The venter of segment nine bears near each lateral margin a tuft of
setae, one or more of which may be enlarged and flattened and is also at
times produced at each posterior lateral angle into a distinct lobe. These
structures may — and very probably do — represent vestiges of the gonopods of
this segment.

The sucking lice are therefore to be characterized as possessing at
least vestiges of an ovipositor.

The External Genitalia of the Male
Figure 15

In all insects the go nopore of the male belongs to the venter of seg¬
ment nine of the abdomen. We cannot here enter into the much disputed
problem of the homologies of the parts associated with these genitalia and
their morphological origins. A study of the general problem, made by Dr.
Joel Gustafson, * has been published and the following conclusions are de¬
rived from it.

Fundamentally, the structure of the external genitalia of the males of
insects is a simple matter. There is the median gonopore itself. There is
a pair of the primitive segmental appendages belonging to segment nine
which form claspers, each consisting of a coxite and a one-segmented mov¬
able piece which is considered to represent the coxal style; and there is
another pair of movable appendages, likewise belonging to the ninth segment,
and derived morphologically from the primitive eversible sacs of this seg¬
ment. These are the parameres. This simple basic set of structures may,
however, be modified in fearful and wonderful ways.

First of all the gonopore may be borne at the apex of what we may regard
as a papilla which may become so greatly enlarged that it forms a sac, here
called the genital sac. This sac, when at rest, is usually retracted into
the body. The gonopore may be surrounded by a sclerotization of the walls
ot the sac and a penis is thus formed. Furthermore, other sclerotizations
of the walls of the sac may form as supporting structures for the penis and
these structures may become extremely complicated. The entire structure
included in all this is the aedeagus.

The pair of structures formed from the primitive segmental appendages of
segment nine may form elaborate clasping structures for holding on to the
female during copulation, or they may be reduced or even lost.

The othei pair of movable structures, the parameres, may become very
closely associated with the base of the aedeagus or may even seem to become
a part of the aedeagus itself. Or they may be lost. Only rarely do both
the claspers and the parameres appear clearly developed in the same insect
and consequently doubt has at times been expressed — or in fact the definite
opinion has been expressed — that two pairs of movable structures do not
actually exist. But the fact that both pairs were a part of the primitive
genitalic complex is indisputable and there are insects in which both are
re tained.

Because of these factors of loss of some parts, coincidentally at times
with unusual development of other parts, the homologies of the parts are at
times much obscured. But if we cling closely to principles and identify
the Landmarks consistently, genitalia of even the most complicated appear¬
ance can be shown to consist merely of modifications of these few basic
parts.

Now to apply these ideas to the Anoplura.

The structures appear in a very simple and even somewhat reduced form in
the male of Pedlculus (Figure 15) as shown in their expanded position when
engaged in copulation.

.. * Gustafson, Joel F. 1950. The Origin and Evolution of the

the Insecta. H l cro «n tonol ogy 15:2.

Genitalia of

basal apodeme

Genitalia of the male of Pediculus humanus

Figure 15

We have here a very large genital sac. Near the apex of the sac is the
gonopore, which is borne upon a sclerotized, tubular process, the penis,
formed as a continuation of the walls of the sac. Close to this penis is
another sclerotization of the walls of the genital sac, for which the term
"statumen penis" was employed by Nuttall — and which presumably serves to
aid in supporting the penis. Arising from the dorsal wall of the sac,
toward its base, is a fold of the wall which is partially sclerotized and
forms a quite large, Y-shaped structure, the apex of which is free. For
this, no better term than "pseudopenis," which in the past has been em¬
ployed, is available. Partially fused with the base of the pseudopenis
there is on each side a short, strongly sclerotized piece which is apically
free. These pieces are here interpreted as the parameres, a term that has
commonly been employed for similar structures throughout the group and
which is in all probability morphologically correct. Articulated to the
base of the sac is a long, flat, sclerotized apodeme which extends into the
body. This piece has commonly been called the basal plate, but the term
genital apodeme would be better.

When the genitalia are at rest they are retracted into the body, with
no more than the apex of the pseudopenis extruded. The parts are then nec¬
essarily folded upon each other in such a manner that it is difficult to
determine their exact relationships and in our systematic work, in which
the details of the genitalia of the male are of great importance for the
separation of species, we can as a rule do little more than merely to rec¬
ognize the sclerotized parts as seen in the retracted position.

The changes in the structure of the genitalia are very largely concerned
with changes in the details of form of the various sclerotized parts, the
parameres, the pseudopenis and the sclerotizations of the genital sac. In
some species apparently certain additional structures are formed from the
walls of the sac which are not yet completely understood and remain to be
elucidated in the future, but this happens in only a few species. In gen¬
eral, if one will recall the simple basic plan of structure, it will not be
difficult to understand the illustrations which will be presented in con¬
nection with the species which are to be treated in detail.

The Tracheal System
Figure 16

As has already been pointed out the thorax always bears but a single
pair of spiracles, these belonging to the mesothorax. They have at times
been ascribed to the prothorax, but this is definitely in error, since no
insect, with the very dubious exception of some of the Apterygota, pos¬
sesses prothoracic spiracles. It has at times been claimed that a pair ex¬
ists on the metathorax in some of the sucking lice, but this too is defi¬
nitely in error.

No species of Anoplura possesses more than six pairs of abdominal spir¬
acles, these being on the third to eighth segments. In many species the
number is less. In one genus, Neol inognathus , there is but one pair, this
being on the eighth segment.

The tracheal system itself is very simple. There is but one tracheal
trunk on each side of the body and there are no tracheal sacs. The accom¬
panying illustration of the tracheal system of Haematopinus suis will suf¬
fice (Figure 16).

The spiracles, however, present some points of special interest. While
they have in the past been illustrated for a few species, the only compre¬
hensive studies of them are those of Webb.* Webb has studied them to some

* Webb, j. E.
49-119; 1947, p

1948, Proceeding

194fl, Pro c ee d ing s o f the Zoological
roceedings of the Zoological Society
s of the Zoological Society of London

Society of London 116:
of London 116:575-578;
118: 582-587.

spiracles of

VI _

t-V/z -

L VIII

IV _

V _

mesothorax _

abdominal

segment

III _

Tracheal system of Haematopinus suis

Figure 16

extent in histological preparations and also has given an extensive account
of them as they appear in such preparations as are used for taxonomic pur¬
poses.

It is not possible in a general work such as this to go into this sub¬
ject in all its details, but since Webb has proposed a partial classifica¬
tion of the Anoplura on the basis of the spiracles they must be considered
at some length.

It may be noted at the beginning that the thoracic spiracles usually
differ somewhat from those of the abdomen, usually being definitely larger
— as is common in almost all insects — and also slightly different in de¬
tails, although of essentially similar character. They are ordinarily
somewhat difficult to study because of the presence of other structures
that somewhat obscure them and we shall here concentrate attention upon the
abdominal spiracles.

In its simplest form an Anopluran spiracle consists of the following
parts. There is an oval or circular opening which leads into a somewhat-
expanded chamber called the atrium. This chamber possesses quite strongly
sclerotized walls and may be subglobular or more or less elongated. On its
inner end it is constricted, usually very suddenly, and then continues into
a tapering second chamber which finally becomes much attenuated. At the
point of greatest attenuation there is usually attached to its wall a mi¬
nute, sclerotized rod which is the apodeme of the occlusor muscle. At this
point the trachea begins with its usual ctenidial threads.

The occlusor muscle originates upon the body wall and by contracting
seems to bring about a pinching of the attenuated inner end of the atrium“
thus closing the spiracle. In some forms the apodeme of the occlusor mus¬
cle seems to be lacking and the muscle inserts directly upon the wall of
the atrium.

Webb has described a gland which accompanies the atrium and discharges a
waxy material that coats the inner walls of the chamber. This gland, how¬
ever, will not appear in ordinary' taxonomic preparations. Webb has indi¬
cated that in some cases what he considers to be the duct of this gland
has taken over the duty of forming the narrowed air passage into the tra¬
chea, while a portion of the attenuated atrium has been left as a side
chamber that ends blindly. This side chamber is at its greatest develop¬
ment quite small. On the basis of Webb's own illustrations and upon gen¬
eral morphological grounds, it is here maintained that Webb has misunder¬
stood the situation. The present writer does not believe that any such
change as that indicated has occurred. It seems much more probable that
the atrium at times is produced on one side somewhat past the internal
opening into the spiracle and thus forms a slight diverticulum.

The walls of the atrium may be beset with little points or ledges which
project into the chamber on the inner side, as Webb has clearly “shown in
histological sections. These ledges vary greatly in their size and extent
and in the degree and manner in which they may anastomose with each other.
Viewing a spiracle from the side they are indicated as dark lines which
sometimes form rings about the chamber or form a network. At times they
correspond with deep furrows on the wall of the chamber that may result in
constrictions, which divide the alrium in ring-like sections as shown in
Figure 17.

The simplest spiracle which appears in the extensive material at hand is
that o I Sctpio aulncodi (Figure 17). Here we have an almost spherical atri¬
um, with apparently no internal ledges. It constricts very abruptly into
the slender prolongation that leads to the opening of the trachea. A very
similar arrangement, is seen in Haematoptnoides squamosus (Fi<mre 17) Tn
kultnotnathus aculeatus (Figure 17) the appearance is the same except that
the second chamber is somewhat swollen and has thickened walls. The next
step seems to be represented in Hae.modlpsus ventricosus and Neohaematoptnus

Haematopinoides squamosus Hesperoctenes

Phthirpediculus propitheci

Neohaematopinus sciuropteri

Scipioaulacodi

Hoplopleura acanthopus

Neolinognathus praelautus Polyplax spinulosa
Spiracles of Anoplura and Polyctemdae

Ctenophthirus cercomydis
Figure 17

sciuropteri (Figure 17) where the transition from the atrium to the second
chamber is more gradual and shows annular constrictions which appear as
slight ledges inside the chamber. In Phthirpediculus propithec i (Figure
17) and Hoplopleura acanthopus (Figure 17) the walls of the atrial chamber
present a number of slight furrows marking the position of internal ledges
which anastomose to some degree. In Enderleinellus longtceps (Figure 18)
the chamber presents deep, annular constrictions which are widely spaced.
In some species, such as Neol inognathus praelautus (Figure 17) and Ender¬
leinellus malaysianus (Figure 18) the lines marking the internal ledges be¬
come more or less anastomosed, forming a network which marks out cell-like
areas on the surface.

In Microthorac ius mazzai (Figure 18) the lines mark out very distinct
cells, but here we have a quite distinctive feature in the fact that at the
points of intersection of the internal ledges the ledges form pointed proc¬
esses projecting into the atrial chamber.

Very beautiful examples of a network and enclosed cell-like areas is
furnished by Linognathus setosus (Figure 18) and Linognathus stenopsis
(Figure 18) . In these species, what seems to represent the diverticulum
formed by the spiracular gland appears as a slight swelling at one side of
the base of the second chamber. In Solenpotes cap illat us °( Figure 18) the
atrium is elongated and almost cylindrical, with the lines which mark the
courses of the internal ledges forming a quite distinctive angular pattern,
but in other species of this genus the atrium is more nearly spherical al¬
though the pattern of the lines is similar. In this genus also there is a
tendency for the spiracle to be borne in a slightly elevated, more or less
sclerotic tubercle, but this is at times much reduced or but little evident.

In Hybophthirus notophallus (Figure 18) the atrial chamber is short and
almost cylindrical, with its walls marked by very short and non-anastomo¬
sing lines indicating the presence of short and interrupted internal ledges.
The second chamber is but little narrower in diameter than the atrium.

In Pediculus humanus (Figure 18) the atrium is more or less conical with
the opening at the apex and is marked by longitudinal lines. Inside it is
beset with slender points. A quite similar arrangement appears in Pthirus.

In Haemtopinus suis (Figure 18), as in other species of this genus, the
atrium is elongated, apparently including the second chamber of the area
usually occupied by the second chamber, and is marked throughout by little,
circular areas which correspond with multitudinous points°which line its
inner wall.

The most distinctive spiracles in the Order are those of the lice of
max ine carnivores. These are here illustrated (Figure 18) as they occur in
Antarctophthirus callor hini, a species in which the membranous derm permits
them to be clearly seen. Here the spiracle opens through a short, mem¬
branous papilla. Slightly in from the apex of the papilla is a peculiar,
sclerotized collar which marks the beginning of a short, simple, sclero-
tized tube that constitutes the atrium. The spiracle then expands’ into the
second chamber, which may be membranous or partially sclerotized and forms
a sort ot bulb. To the wall oi this bulb attaches the lar^e apodeme of
the occlusor muscle and then the trachea begins. This type of spiracle is
very distinctive of the little group of species considered to form the fam¬
ily Echmophthimdae.

The significance of these structures in the intra-ordinal classification
Anoplura will be considered in the chapter on classification
a possible bearing upon the question of their value for such
there are here presented illustrations (Figure 17) of the spiracles
genera ol the Hemipterous family Polyctenidae, which are ectopara¬
sites occurring on bats. It may be noted that in their general
they quite closely resemble the spiracles of some of the
display the same characteristics of an atrium and

of the
having
poses,
in two

As
pur-

features
Anoplura. They
second chamber, with

Solenopotes capillatus

Hybophthirus notophallus

Enderlemellus longiceps

Linognathus

stenopsis

Microthoracius mazzai

Pedicinus obtusus

Enderlemellus malaysianus

Pediculus

Haematopinus suts

aspect

ventral

aspect

Spiracles of Anoplura

Figure 18

the walls of the atrium beset internally with ledges. The principal dif¬
ference lies in the presence of a very well-developed diverticulum which
may possibly represent the atrial gland described by Webb for the Anoplura.

We may here call special attention to the spiracles of Enderleinellus
malaysianus (Figure 18) and Enderleinellus loniiceps (Figure 18). It may
very well be that at some time in the future these two species will be re¬
ferred to different genera, but with the utmost degree of splitting they
will stand as still belonging to closely related genera. Yet the pattern
of these spiracles does not suggest any close relationship.

Internal Anatony

There still remain numerous details of the internal anatomy of the Ano¬
plura which have not been well investigated and which cannot here be consid¬
ered. This chapter is therefore somewhat incomplete. It will be con¬
ceited only with the major features, especially as they may have a bearing
upon the understanding of external morphology and thus aid in the problem
of taxonomy.

The Head

It was necessary to consider certain aspects of the internal anatomy of
the head while we were attempting to explain the homologies and functioning
of the mouthparts. What remains to be reviewed consists chiefly of the
nervous system.

We have briefly mentioned the musculature of the antennae. In insects
which are provided with a tentorium the muscles of the antennae originate
upon it. But since there is no tentorium in the Anoplura the muscles orig¬
inate upon the head wall. There is some variation in the position of their
origins, in Haematopinus all being upon the dorsal wall, while in some
other species a portion of the origins are upon the ventral side.

This is a matter of some general interest, since the claim has been made
by one morphologist that the origins of muscles do not shift. There can be
no possible question of the fact that they do so in these insects.

Pawlowsky's glands, as previously noted, are a pair of rather large,
apparently glandular structures, opening separately through the dorsal wlall
of the trophic sac (ligure 11) . They are innervated by a branch from the
hypopharyngeal nerve and thus may definitely be regarded as belonging to
the hypopharynx and consequently to the clypeal segment. A general study
of the occurrence of glands in this region in other insects will be requir¬
ed before their homologies can be determined.

We ma) here call attention once more to the peculiar mass of connective
tissue which occurs in the posterior portion of the head and which we have
called the obturacuium. It extends into the thorax, where it envelopes the
thoracic ganglia of the nervous system. It serves as the point of origin
of various muscles and in this respect, if in no other, presents a quite
unique development. Again, no conclusions concerning this mass of tissue
can be drawn until a general study of connective tissue in this region of
the body of other insects has been made. The present knowledge of the oc¬
currence of connective tissue in the bodies of insects is exceedingly
sLight. & J

The salivary glands have already been noted and described in connection
with the morphology of the head and mouthparts.

The muscles found in the head have been considered to some extent in con¬
nection with the description of the mouthparts. We shall not here describe
them in detail, muscle by muscle, relying rather upon the illustrations
which have already been presented, but we shall refer to them again in con¬
nection with the nervous system.

The Thorax
Figures 13, 14, 19

It has already been pointed out that the true notum of all of the thor¬
acic segments is much reduced. Apart from the ganglia of the nervous sys
tem, which will be considered later, the thorax is occupied mostly by the
muscles connected with the legs. We shall here content ourselves with a
general description of the disposition of these muscles which will aid to
some degree in understanding the external structures.

In Figure 19 A the muscles are shown as they appeal' when the dorsal derm
has been removed. In Figure 19B are shown the muscles which are revealed
by the removal of the first layer of dorsal muscles.

Note in Figure 14A the prothoracic pleural ridge or phragma. Upon this
phragma there originate on each side of the body a little cluster ol muscles
which insert upon the anterior margin of the sternal plate, these being
properly designated as the dorsoveutral prothoracic muscles. Just anterior
to the phragma a little cluster of muscles originates and these muscles pass
forward to insert upon the posterior border of the head near its lateral
margin . From each of the two apophyses at the posterior bordex- of the head
narrow muscle bauds extend posteriorly to the median notal apophysis or pit.

The muscles of all the legs converge to points of origin upon the trans¬
verse, sclerotized band which includes the notal pit or apophysis, this
band is apparently formed by a continuation and fusion of the pleural ridges
or phragmata of both the mesothorax and metathorax.

There are three muscles inserting upon each coxa. Since the muscles of
the coxae in other insects — there usually being four of them — have been
named according to their functions as promotor, remotor, levator, and de¬
pressor muscles and since these various functions seem here to be carried
out by but three muscles, it becomes a very dubious procedure to assign
specific names to them and no attempt will be made to do so. They are per¬
haps best called simply anterior, posterior, and dorsal muscles. The dor¬
sal muscle of the anterior coxae sends a small branch to the corresponding
pleurostemal apophysis.

From the notal pit bands of muscles radiate posteriorly to attach at the
posterior border of the thorax (which may involve an element belonging pro¬
perly to the abdomen) and presumably function to move the thorax in rela¬
tion to the abdomen. ,

Ventrally there are very few muscles (Figure 19B) . Attention may be

called to a single small muscle strand extending from each coxa to the mar-
r\u of the sternal, plate; to a anaLl strand which passes across between the
prothoracic pleurostemal apophyses; to another quite large strand which
extends across between the posterior coxae.

The musculature within the legs themselves shows no especially interest-
in f features and will not be discussed here.

The Abdomen

The segmentation of the abdomen has already been discussed.

Apart from the muscles connected with the genital organs the principal
muscles of the abdomen are found in the relatively huge dorsoventral bun¬
dles arranged in a row on each side of the alimentary canal and extending
from the termite to the sternite of each segment. It may very well be that
upon these muscles depends to a large degree the compression of the abdomen
which sets up internal pressures in the body fluids by which various move¬
ments— such as perhaps the extrusion of the trophic stylets— are at least

in part accomplished. . , , , , .

The muscles of the reproductive structures will be considered in connec¬
tion with the account of these structures.

prothoracic pleuro- sternal apophysis

notal ^
Pit

stomach- — "

- occipital apophysis

.spiracle

A. thoracic musculature , dorsal muscles in place

spiracle

notal

pit

stomach - ^ ^ ^ ''^salivary gland

B. thoracic musculature,dorsal muscles removed, and tracheal system

prothoracic pleuro- sternal apophysis

■v

^ occipital apophysis

Musculature of thorax of Haematopinus suis

Figure 19

The Alimentary Canal
Figures 20, 22

The very slender oesophagus passes through the obturaculinn and then ex
ponds abruptly into the relatively huge mid-gut. Keiliu and Nuttall* indi¬
cate the presence of caeca at the anterior end of the mid- gut, but these do
uot appear in Haematopinus suis. This large intestine then passes directly
through the body without any convolutions until it becomes the small intes¬
tine. The small intestine is somewhat convoluted and variously swollen,
these swellings perhaps depending upon the contents. It passes finally in¬
to a swollen, bulbous structure, the rectal sac, then narrows again to the
anus. Upon the rectal sac are four or more pad-like or plate-like struc¬
tures which are the rectal "glands" or rectal pads as found i n other insects.

The Organs of Excretion

There appear to be but four Malpighian tubules in Haematopinus suis and
four are recorded by Keilin and Nuttall* for Pedtculus humanus. These dis¬
charge into the alimentary' canal near the anterior end of the small intes¬
tine. Keilin and Nuttall have indicated that excretory products may be
stored in certain "nephrocy tes" or cells of the fat body, but this is ques¬
tioned by Wigglesworth .

The Heart
Figure 20

The dorsal vessel is of a peculiar form in the sucking lice. It con¬
sists mostly of the very long and slender aorta which continues posteriorly
almost to the end of the abdomen and then forms a bulbous swelling, from
which muscles extend apparently to the body wall.

The Reproductive Organs of the Female
Figure 21

Each ovary consists in Haematopinus suis of five ovarioles and the same
number is recorded by Keilin and Nuttall for Pediculus humanus. Apparently
at any moment only one of these tubules will contain an egg ready for de¬
position. The ovarioles are distributed along an oviduct which leads to
the uterus from each ovary. Opening into each oviduct there are in Haema¬
topinus suis apparently three quite large accessory' glands.

The common oviduct or uterus is a very large structure with apparently
muscular walls. The duct from each ovary opens into the uterus far back
toward the posterior end of the latter.

At its posterior end the uterus opens into the vagina and from the vagina
various small muscles pass to the body wall.

The spermatheca has already been described. It apparently does not
occur in all species of the Anoplura and the details ol its histological
structure remain still to be studied.

The Spermatheca

A single spermatheca is present in the female of so many species of
.Anoplura "that it seems justifiable to regard this structure as a part of
the basic structure of the group, although in some species it seems to have
been lost. Thus, it appears to be lacking in Pediculus, Haematopinus, and
some other genera.

- * Keilin, D. , and G. H. F. Nuttall. 19 30. Parasitology 22:Plate XI.

oesophagus „

caecum _

ventriculus _

accessorg glands _ _

tnalpighian tubules

heart- - - - "

vagina - - '

Internal abdominal structures of female of Pediculus humanus

salivary glands

jiorta

ovarioles

jyjlorus

rectal plates

heart muscles

"'-rectum

Figure 20

uterus

ovarioles

oviduct - -

vaginal muscles^

__ accessory gland

~ vagina

Reproductive organs of female of Pediculus humanus

Figure 21

oesophagus

. mycetome

_j)as deferens

. _ pylorus

^'s^vas deferens

vesicula seminalis

aedeagus

caecum .

ventriculus _

testes -

malpighian tubules

rectal plates.

rectum--'"

Internal abdominal structures of the male of Pediculus humanus

Figure 22

In appearance it consists of a deLicate, weakly scierotized tube which
extends from the dorsal wail of the vagina for a short distance and termin¬
ates in a membranous or more or less scierotized bulb. The opening into
the vagina is commonly surrounded by a crescentic or circular ring ol strong
sclerotization and this ring is at times almost the sole evidence ol' the
presence of the spermatheca.

The bulb is commonly extremely delicate and subject to distortion in
preparations and even to loss. Because of these circumstances it is gener¬
ally of somewhat dubious taxonomic value. In one group, however, that of
certain species of the genus Knderleinellus, it is extremely helpful, its
variations presenting the only apparent means of separating the females of
certain of the species. It merits an extended study, throughout the Ano-
plura, which is scarcely practicable in a work of the present type.

For an example of a well-developed spermatheca the reader is referred to
the illustration of the genitalia of the female of Pthtrus pubis (Figure
124).

The Reproductive Organs of the Male
Figures 22, 23

The number of testes can be interpreted either as four with the members
of each pair connected together by a short and slender tube, from which the
vas deferens issues, or as two, each of which is much constricted to form
a dumbbell-shaped structure, with the vas deferens issuing from the con¬
stricted portion. Each vas deferens proceeds for some distance as a slen¬
der tube and then expands to form a thick tube which has been considered by
various authors to represent the vesicula seminalis. The two vesiculae lie
close together, appressed against the ventral side of the alimentary canal,
and are twice convoluted, the convolutions extending forward almost to the
anterior border of the abdomen. Posteriorly, the vesicles unite into a
common tube which forms the ejaculatory duct.

The ejaculatory duct terminates at the apex of the scierotized penis, so
that when the genitalia are extruded as in copulation it must traverse the
length of the aedeagus.

The musculature of the genitalic parts has been studied in connection
with this work only in Haematopinus suis (Figure 24) and here only suffi¬
ciently to give a general idea of the mode of operation. It is probable
that other species will show some departure from the arrangements here il¬
lustrated.

A relatively massive muscle inserts upon the base of the basal apodeme
ventrally and originates upon the terminal sternite of the abdomen. Appar¬
ently this muscle, probably with the aid of blood pressure, serves to ex¬
trude the genitalia. From the dorsal side of the basal apodeme arises a
mass of muscle bundles which curve toward the mid-line where those from the
two halves unite and finally insert upon the base of the genital sac.
These are undoubtedly concerned with the retraction ot the sac. From each
side of the sac a series of muscle bundles extends to the ventral body wall,
where the bundles originate upon the genital plate. These apparently also
serve as retractors. Another small muscle inserts on each side near the
apex of the sclerotization of the genital sac and inserts on the termi¬
nal sternite of the abdomen close to the apex of the body. The function of
this is not entirely clear, but it is evidently concerned with the retrac¬
tion of the complex.

The Nervous System
Figure 24

For an understanding of the elements of the nervous system as related to

testes

vas deferens

_ vesicula seminalis

— — aedeagus

ejaculatory duct

Structure of male reproductive organs of Pediculus humanus

Figure 23

nerve to labral
compressor muscle

nerve to formal muscle _

nerve to mouth angle
retractor muscle _

nerves to minor
mandibular muscles*. '

supraoesophageal
ganglion -

nerves to major
mandibular muscles

undetermined - "
structure X

suboesophageaL " "
ganglion

salivary nerve-''

nerve to folded
maxillary
retractor muscle''

nerve to transverse''
labial muscle

prothoracic nerves' ’

nerves to cibarial muscles

Jrontal ganglion

recurrent nerve

_ _ antennal nerve
_ optic nerve

'' / //'X

^ nerves to
Pawlo iv sky 's glmuts

nerve to hypophargngeal
stator muscle

- - ventral nerve cord

' ''''^nerves to folded
maxillary and
\ labial protractor muscles

•V

^ \ '

nerve to folded labial
retractor muscle

^ mesothoracic nerve

abdominal nerves

s metathoracic nerves

Nervous system of Haematopinus suis

Figure 24

the segmentation of the head the reader is referred to a series of papers
hy Laura M. Henry* in which the significance of the innervations is dis¬
cussed in detail. The conclusions there presented are here adopted without
reservation and have been employed as the basis for an understanding of the
head and mouthparts of these lice.

Some description of the arrangement of the nervous elements in the head
has already been given in connection with the consideration of the mouth-
parts, but we may review this again in connection with the nervous system
as a whole.

From the supraoesophageal ganglion there originate three pairs of nerves.
Of these, one pair innervates the eyes and one the antennae. Those of the
third pair are the nerves of the labrum and the stomodaeum and innervate
all muscles which insert upon these parts. All muscles which they inner¬
vate are therefore to be identified as inserting upon structures that be¬
long morphologically to the labral segment, which is segment one of the
body. These nerves pass forward and branch. A mesal branch from each
leads to the frontal ganglion and close to the point of origin of this
branch is a lateral hranch which passes to the muscle which we identify
as the mouthangle retractor. The main nerves continue forward and give off
laterally a branch which innervates what we consider to be the tormal mus¬
cle, a muscle that inserts upon the posterior- lateral angle of the palatal
plate. Continuing, the main nerves innervate the muscle that we consider
to be the compressor of the labrum.

From the frontal ganglion a single median nerve extends posteriorly,
this being the recurrent nerve which cares for the innervations of the an¬
terior portion of the alimentary’ canal, that is to say of all those por¬
tions derived from the stomodaeum. We have not followed it in complete de¬
tail, but Florence** has indicated that it forms a small ganglion which may
be identified as the occipital ganglion, and then branches.

The circumoesophageal connectives are very short and the opening en¬
closed by them for the passage of the oesophagus and other structures is
extremely small.

Arising from the suboesophageal ganglion, somewhat toward the meson, is
a pair of nerves which through branches supplies the two pairs of muscles
that we identify as belonging to the mandibles.

Next comes a pair of nerves which arise from near the side of the sub¬
oesophageal ganglion and which, through branches, innervate all the struc¬
tures that we identify as belonging to the hypopharynx, these including
Pawlowsky' s gland and certain muscles. From each of these main nerves,
near its base, there arises a short stalk that terminates in a little,
knob-like structure, indicated in Figure 25 as X. We have no evidence as
to what this structure is.

There follows a pair of nerves which we consider to be formed by a fu¬
sion of the maxillary and labial nerve trunks and which innervate the mus¬
cles assigned to the maxilla and labium. The last pair innervate the sali¬
vary glands and are connected by a small cross-nerve to the labial-maxil¬
lary' trunk.

Following the suboesophageal ganglion the ventral nerve cord is very
slender for a short distance and then expands into the huge mass of fused
ganglia in the thorax. This ganglion mass is very evidently composed of
the three thoracic ganglia and includes also all the abdominal ganglia.
From its posterior- lateral angles there arise on each side four nerves
which supply the structures of the trbdomen. We have not traced these nerves
beyond the point of determining that the last nerve of the series, which is

♦ Henry, Laura m! 1947, M icroent on o logy 12: 3: 65-82; figs. 23-29; 1947,

Mi croentono lo gy 12:4:83-110; figs. 30-45; 1948, M i cr o e n t ouo lo g y 13:1:1-26;
figs. 1-9; 1948, Microentomology 13:2:27-48; figs. 10-16.

** Florence, Laura. 1921. Cornell University Agr icul tur al Sxp er inent
Station, Ithaca, tew fork. Memoir 51, pages 642-743.

the largest, cares for all the reproductive system. There is evidently a
considerable decree of fusion of the nerves to the various segments and
this remains still to be worked out.

We have not traced the innervation ot the heart. It is therefore evi¬
dent that there is more work still to lie doue before all the details of the
nervous system are finally established.

The Mycetome
Figure 22

The structure called the mycetome has been noted by several authors in
various species of Anoplura and knowledge concerning it has been summed up
by Steinhaus in his book "Insect Microbiology" (1946). The entire subject
of the mycetome is a rather complicated one and will not lie reviewed here.
It will suffice for our purposes to uote it as a visible disk of cells ly¬
ing on the mid-gut in the larvae and in the adult male. In the adult female
the enclosed symbionts leave the region of the stomach and migrate into the
ovaries. The mycetome has been observed in Pediculus humanus, Pthirus
pubis, a species of Pedtctnus, Haematopinus suis, eurysternus, and asini,
Linognathus vituli and setosus, and iu Polyplax spinulosa. The position
of the mycetome appears generally to be essentially as is indicated for the
male of Pediculus humanus in Figure 22.

CHAPTER III

Growth and Development

The Egg

Figures 25, 26

As far as is known the sucking lice, with one single exception, always
attach their eggs to the hairs of the host. This single exception is a
louse of man which will attach eggs to the clothing. Even here the method
is essentially the same as in other forms, the eggs being attached to pro¬
jecting fibers whenever possible. The attachment is by means of a drop of
cement which surrounds a single hair, or in some instances several hairs,
and encloses one end of the egg itself. Florence (1921) has indicated that
the cement originates in the collaterial or accessory glands illustrated in
Figure 21.

The free end of the egg is always provided with an operculum that breaks
away to permit the exit of the larva and, as far as observations go, the
egg is always oriented with this end away from the base of the hair upon
which the egg is attached. The eggs are always arranged singly along the
hair, although at times they may be placed in a very closely spaced series.

The total amount of information concerning the eggs of the Anoplura is
relatively small. The eggs of a number of species have been illustrated
from time to time but these illustrations are usually but little detailed
and the structures of the egg shell have only rarely been studied in de¬
tail. Such information as there is has been derived mostly from the egg of
Pediculus humanus.

In many forms, such as Pediculus humanus, the operculum is beset with
small, knob-like tubercles, each of which carries a minute external opening.
This pore communicates with a somewhat expanded chamber, which in turn com¬
municates with the interior of the egg by a pore. However, this arrangement
is by no means present, at least in such form, in the eggs of all species.

Unfortunately, the total amount of material available in connection with
the present work has been quite small and the eggs of only a few species
can be illustrated. These are sufficient, however, to show that there is
considerable variation in details and to suggest that an extended study
would be worth while and might throw some light on the problems of general
classification. It may be remarked that eggs can be obtained with no dif¬
ficulty from the skins of mammals in museum collections, even if remains of
the lice themselves are not present. However, this presents difficulties,
since as many as three species of lice may occur upon a single host and
consequently doubt will arise in many instances as to the ascription of the
eggs to the proper species. Also, there is the difficulty that a large
percentage of the eggs will have hatched and consequently the operculum
will be lacking. In such examples the most distinctive characters of the
eggs will have been lost.

On the basis of the material that is available the following notes are
ottered. In Figure 25C is shown the egg of Pediculus humanus corporis as
attached by an irregular drop of cement to a cluster of fibers of the cloth¬
ing of the host. The operculum is shown as detached and flattened. It
wiLL be noted that there are 15-20 of the little air cells or cellulae,
these being arranged in an eccentrically placed group. In eggs of what is
here for the moment called Pediculus chapini (Figure 26D) the number of
cells is lewer , but material has not been available to permit the dissec¬
ting away of an operculum in such a manner as to reveal the exact number.

Eggs of Anoplura

Figure 25

A. Neohaematopinus sciuropteri

B. Haematopinoides
squamosus

C. Linognathus breviceps

Eggs of Anoplura

In Pthirus pubis (Figure 25F) the cellulae seeiu to occupy practically the
entire area ot the opercular surface and are large and elongated. In
Hoplopleura oenomydis (Figure 25A) the cellulae are moderately large and
are arranged in a single row extending around the operculum close to the
margin. In Hoplopleura crypt lea (Figure 25B) the cellulae are very small,
being scarcely larger than the enclosed pore and there seem to be about 16
of them. In Pahrenholzia microcephala (Figure 26D) there seem to be only 3
or 4 cellulae and these are very small and are displaced towai'd one side.
In L inognat hus breuiceps (Figure 26C) there are perhaps 10 cellulae, these
occupying the centi’al area. In Enderleinellus osborni (Figure 2f»t ) there
are no cellulae and no evident pores. In Haematopinus suis (Figure 26E)
there are no evident cellulae.

The egg shell appears sculptured in some species. Thus, in the egg of
Hoplopleura oenomydis (Figure 25A) it is mai’ked by two sets of diagonal
ci'osslines which divide the surface into little, lozenge-shaped areas that
can be seen only in favorable specimens and by proper adjustment of the
light in the microscope. In Haematopinus suis the entire shell, including
the operculum, is marked with minute, clear, cellular areas. In Hoplopleura
crypt ica the shell bears somewhat irregular, cellular areas. In other
species here illustrated there seem to be no markings of any kind. In the
egg of Neohaematopinus sc iuropteri (Figure 26A) — or at least on the basis
of circumstantial evidence an egg presumed to belong to that species — the
available specimens have lost the operculum, but the base of the egg is
very distinctly marked by transverse folds or ridges.

The form of the cement pedicel varies. Its attachment to the hair may
be very short or may be much extended. It may be rough and irregular or
smooth. The attachment to the egg may be strictly apical or may extend up
one side of the egg. In the egg of Haematopinoides squamosus (Figure 26B),
which seems usually to be attached to a cluster of hairs of the host, the
cement itself is transversely wrinkled on the side away from the attachment.

The number of hairs enclosed within the cement seems frequently to be
variable. In general, it appears on the basis of the small amount of avail¬
able material that ordinarily a single hair is involved. There seems to be
no special choice as to the size of the hairs. Thus, eggs are at hand of
Fahrenholzia microcephala, taken from the same host individual, some of
which are attached to small, slender hairs and others to large, spiny hairs.
In the latter case the cement is spread very thinly' around the hair.

In the form of the eggs there seems in general to be nothing especially
distinctive, although certainly in some species this is not true. The egg
of Haematopinoides squamosus in the material at hand is somewhat constric¬
ted medially. The egg of Pediculus chaptni is quite pointed basally. The
egg of Heohaematopinus sc iuropteri is noticeably long and slender. In most
of the species here illustrated the actual form of the egg is probably more
slender than is indicated, since dried eggs usually collapse with consequent
apparent broadening.

Developmental Stages
Figures 27, 28 , 29 , 30

The existing knowledge of the immature stages of the Anoplura is quite
scanty, there being probably not more than a half-dozen species of which
all the stages are known. TTie only species at hand of which all stages are
represented are Pediculus humanus, Pediculus mjQber§i (Figure 27) and
Pedicinus obtusus (Figure 28).

There seem to be a total of four instars. It will be noted that in
these species the principal changes which occur, apart from the development
of the genitalic structures and alteration of proportions, are associated
with the development of the paratergites . In Pediculus mjH>ber£i these are

2nd instar

Nymphal stages of Pediculus mjobergi Ferris

3rd instar

Figure 27

2nd instar 2

Nymphal stages of Polyplax spinulosa (Burmeister)

Figure 29

recognizable in the second instar, while in Pedicinus obtusus they do not
appear until the third instar. Also, in these species there are certain
changes in the antennae, the last three segments being scarcely or not at
all indicated in the first three instars. This is not generally true, the
first stage usually having five antennal segments as does the adulf.

Of other species, there are at hand only a few immature individuals, no
complete developmental series being represented. Consequently, it is dif¬
ficult to assign these specimens to their proper instars. On the basis of
present indications, however, it appears that th*e only species in which the
immature stages present any especially peculiar developments are those of
the genus Hoplopleura. In this genus there seems to be a general tendency
toward the development of sclerotized tubercles on the antennae and on the
ventraL side of the head and also of unusual setae on the abdominal margins.
It is not clear in which stage the paratergites appear in these forms.

The appearance of the immature stages in the genus Hoplopleura is such
that an incautious worker might very well be led to suspect the presence of
more than one species. Illustrations of some of these stages of this genus
are herewith presented.

intermedia

First stage of species of Hoplopleura

pelomydis

CHAPTER IV

The Taxonomic Status of the Sucking Lice

Historical Review

The sucking lice are for the most part very small forms and require
special methods of preparation for study under the microscope. Such meth¬
ods were scarcely available until comparatively recent times and in fact
have reached a satisfactory decree of development only within the last
thirty- five years; even yet very few entomologists have become fully aware
of them or have become proficient in their use. Moreover, the members of
this group of ectoparasites were long regarded as merely "disgusting para¬
sites" and therefore scarcely worthy of the attention of anyone possessed
of esthetic feelings. And still beyond that, it requires a special effort
to secure material of any large representation of the group, since the col¬
lector must first obtain specimens of the mammals upon which they occur.
For all of these reasons they have until relatively recent years received
but little attention from entomologists. Their taxonomic history repre¬
sents a series of stumbling attempts to assign them to some generally ac¬
ceptable position in the scheme of classification of the insects and to de¬
vise a system of classification within the group itself. Even yet there is
no absolute agreement in regard to either of these aspects of their classi¬
fication.

Fahrenholz,* in a paper published in 1936, gave a detailed review of the
taxonomic history' of the group, which has been utilized in preparing the
following summary. This has been supplemented from other sources and
brought up to date, especially with the aid of a work on the history of the
development of insect classification published in 193V. **

We need not concern ourselves with anything thatmayhave been done prior
to the appearance of the tenth edition of Linnaeus' "Systema Naturae" of 1758,
upon which our present system of classification and nomenclature is ground¬
ed. In this work Linnaeus adopted the ancient Latin name Pediculus for a
genus into which he threw almost everything that could conceivably be called
a "louse," placing this genus in the insect Order Aptera along with the
mites, fleas, and the primitively wingless group now called the Thysanura.
The genus contained a weird assortment of forms, including biting lice,
sucking lice, Corrodentia, a beetle triungulinid and a Hippoboscid fly.

In 1775, Fabricius, in his "Systema Entomologiae", included the genus with
essentially the same composition in his Order Antliata, placing in it a mis¬
cellaneous assortment of forms that included mites and some flies, and this
system was followed in his "Systema Antliatorum" which was published in 1805-

In 1806, Latreille recognized the Order Parasita, which was composed of
the two groups of lice — the biting lice being placed in the genus Ricinus
and the sucking lice in the genus Pediculus.

Also in 1806, Lamarck, in his "Histoire Naturelle des Animaux sans Ver-
tebres, " placed these genera in the Order Arachnides Antennistes, in the
strange company of the centipedes, the myriopods, and certain insects now
generally referred to the Order Thysanura.

In 1815, Leach, in the "Edinburgh Encyclopaedia," placed the lice in the

•Fahrenholz, H. Zeitschrift filr Parasitenkunde, Volume 9, Part 1, pages
50 - 57 . 1936.

*• Wilson, H. F. , and M. H. Doner. The Historical Development of Insect
Classification. 1937.

uew Order Anoplura, this including two genera of sucking lice, Fediculu s
and Huematopinus; and one jfenus of biting Lice, Nirmus. This arrangement
was adopted also by him in "Zoological Miscellany" in 1817, where he placed
the sucking lice in the f;unily Pediculidae and the biting lice in the lam
ily Nirmidae.

In 1818, Nitzsch, in a paper entitled "Darstel lung der Familien uud (jat-
tungen der Th ierinsekteu, " published in Volume ill of Geimar's Magazin fiir
die Entomologie, recognized these two groups but placed the sucking lice in
the Hemiptera Epizoica and the biting lice in the Orthoptera Epizoica.

In 1823, Dumeril, in his "Considerations generates sur le classe des in-
sectes, " placed the sucking lice in the group Khinaptera along with the
fleas and at least some mites.

In 1825, Latreille, in his "Families uaturelles du regne animal," re¬
tained the Oilier Parasita and divided it into two groups, Siphunculata for
the sucking lice and Mandibulata for the biting lice.

In 1826, Kirby and Spence, in Volume IV of their "Introduction to Ento¬
mology," placed all the lice together in the Order Aptera, still along with
the Thysanura, the inyriopods and various arachnids.

At some time between 1835 and 1840, burmeister, in his "Handbuch der Eu-
tomologie, " which was published in five volumes during these years, placed
the sucking lice under the Order Rhynchota and the biting lice in the Order
Mallophaga, this apparently being the first use of the latter name.

In 1842, Denny, in his "Monographia Anoplurorum britanniae," restored
the Order Anoplura, dividing it into two suborders. For the sucking lice
he employed the subordinal names Rhynchota or Haustellata and for the bit¬
ing lice the names Mallophaga or Mandibulata, in each case apparently as
alternatives.

In 1874, Giebel, in his great work "Insecta Epizoa, " placed the sucking
lice as the family Pediculina in the Order Hemiptera, calling the group
Hemiptera Epizoa.

In 1880, Piaget, in his monumental "Les Pediculines, " very frankly a-
voided the use of any ordinal names, but regarded the sucking lice as con¬
stituting a single family, the Pediculidae, and the biting lice as two fam¬
ilies, Liotheidae and Philopteridae, the three families being considered as
of equal rank.

In 1896(7), Haeckel referred the two groups of lice to the Order Phthi-
raptera and this name has recently been taken up by Weber and by Eichler,
these authors using it to include the suborders Anoplura, Mallophaga and
Rhynchophthirina, the last name being that which was proposed by Ferris for
the reception of the genus Haematomr/zus.

At some time during the latter part of the nineteenth century the idea
became established in the minds of entomologists that the sucking lice are
related to the Hemiptera. Thus in the earlier editions of Comstock's
"Manual for the Study of Insects," and in fact in an edition as late as
1913, they are placed as the Suborder Parasita under the Hemiptera. In
Kellogg's "American Insects" (1908) they are thus placed. On the other
hand, the Mallophaga were recognized by Comstock as a separate Order as

early as 1895-

In 1903, Cholodkovsky, in "Zoologischer Anzeiger, " Volume 27, recognized
the sucking lice as an independent Order and proposed the name Pseudorhyn-
chota for them.

In this same journal and volume Shipley proposed the ordinal name
Eliipoptera for them in an attempt to establish a uniform system of endings
for the names of the insect Orders.

Also in this same journal and volume Borner, in a projected classifica¬
tion of the insects, recognized the sucking lice as the Order Siphunculata,
crediting the name erroneously to Meinert.

In 1908, Handlirsch, who probably had a broader understanding of insect

classification than any other man who has ever lived, recognized the suck¬
ing lice under the ordinal name Siphunculata and he adopted this assignment
in°a section of Schroeder's "Handbuch der Entomologie, " which was published

in 1923.

In 1908, Dalle Torre, in Wytsmann's "Genera Insectorum, ' published a
catalogue of the sucking lice, calling them the Order Anoplura.

In 1910, Mjoberg, in a work entitled "Studien iiher Mallophagen und Ano-
pluren," which appeared in Volume 6 of "Arkiv for Zoologi," reverted to the
opinion that these two groups constitute a single Order, for which he ap¬
parently used the name Siphunculata, while employing the subordinal names
Anoplura for the sucking lice and Mallophaga for the biting lice.

In 1916, Lancelot Harrison, in a paper published in Volume 18 of the
"Proceedings of the Cambridge Philosophical Society," also supported this
view.

Apparently as a result of the conclusions of these last two authors, the
writers of various textbooks — notably Imms in his "General Textbook of
Entomology" — have adopted this opinion and have united the two groups, Imms
employing the ordinal name Anoplura for both.

Ferris, in his series of papers entitled "Contributions Toward a Mono¬
graph of the Sucking Lice"— published in the Stanford University Publica¬
tions, Biological Sciences Series, over a period from 1919 to 1934 — consid¬
ered the sucking lice to belong by themselves in the Order Anoplura.

Fahrenholz, in a paper published in 1936 in "Zeitschrift fur Parasiten-
kunde, " Yolume 9, followed this procedure, but included in the sucking lice
the peculiar genus Haematomyzus, which has but one species, the louse of
the elephants. Ferris had shown earlier that this insect is not a sucking
louse, being an insect with biting mouthparts and probably more closely re¬
lated to the Mallophaga, to which Order he assigned it. Fahrenholz retained
it in the sucking lice purely on the grounds that functionally it is a
sucking form. He divided the Anoplura into two Suborders — Rhvnchophthirina,
a name previously proposed by Ferris as a Suborder of the Mallophaga for
inclusion of Haematomyzus ; and the new name Inrostrata, for the true sucking
lice.

In 1939, Weber, in "Biologisches Zentralblatt, " Volume 59, took up
Haeckel's name Phthiraptera, as already noted, placing under it as suborders
the three groups Anoplura, Mallophaga, and Rhynchophthirina, the last named
including only the louse of elephants. In 1946, Eichler, in "Archiv fur
Naturgeschichte," Neue Folge, Volume 10, Heft 3, also adopted this arrange¬
ment .

In 1946, Webb, in "Proceedings of the Zoological Society of London,
Volume 116, adopted the opinion that the lice constitute a single Order, to
which he applied the name Anoplura.

In 1949, Hopkins, in Volume 119 of the same journal, followed Weber's
procedure of placing all the lice in the Order Phthiraptera.

Throughout all this time the authors of various textbooks of entomology
and parasitology, none of whom had any personal acquaintance with the in¬
sects involved, have adopted one or another of the various arrangements and
one or another of the various names. It would be useless to consider these
textbooks, since they contribute nothing original.

The problem of the name to be used for the sucking lice involves two as¬
pects, one being purely nomenclatorial, the other zoological.

The problem of nomenclature being the simpler may be disposed of first.
To the question of what name should be employed for the sucking lice there
is no absolute answer. The International Rules do not cover situations of
this kind, since they do not extend to the names of groups above the super-
family and even here are not definite. The only rule that can be followed
is some rule of reason. The subject is subjudice at the moment, to be set¬
tled, perhaps, at Copenhagen in 1953-

Apparently the first ordinal name employed for all the 1 ice was Paras i ti ,
proposed by Latreille in 1796. If priority holds, this name would have to
be maintained either for al L the lice together as combined in a single Or
der, or for one of the groups if the Order be divided. But priority does
not necessarily hold in such circumstances and there is the objection to
this name that it is entirely too broad in its connotations. Furthermore,
the name Parasitidae has been employed also for a family of mites.

The second available name seems to be Anoplura, proposed b) Leach in
1815. This name has come into quite general use and conveys no suggestion
of applicability to any group other than the lice. The opinion here held
is that there is no reason why it should be replaced by any of the names
which were later proposed.

Now, since this name was originally proposed for the biting and the
sucking lice as combined in a single group, the question arises as to which
group should inherit it if the Order is divided. The opinion is here main¬
tained that since the biting lice were very early removed as the Order
Mallophaga and were thus supplied with a name that has long been employed,
the reasonable procedure is to retain the name Anoplura for the sucking
lice if they be recognized as an independent Order.

We come now to the zoological question involved, that of whether the two
groups should be recognized as separate Orders or merged in a single Order.

This problem has no objective solution, whatever answer is adopted being
merely one of opinion . The two groups exist, but the rank to be assigned to
them depends solely upon the mental processes of the protagonists of either
opinion and upon subj ective ideas concerning philosophical questions involved
in the general process of classification. The groups are nearly enough
alike to justify union, or they are sufficiently different to justify sepa¬
ration, either step being dependent upon the weight assigned to similarities
of habit on the one hand or to differences in morphology on the other.

The author, being a morphologist, claims the greater weight for mor¬
phology. And from the point of view here adopted this argues for ordinal
separation.

The principal distinction between the sucking lice and the biting lice
(the latter including the Rhynchophthirina) lies in the mouthparts. The
feeding mechanism of the sucking lice, which will be described in detail in
a later section, differs very greatly from the mechanism found in the biting
lice and there are no known transitional forms between the two groups. Nor
is this all. The arrangement of the thoracic elements is quite different
in the two groups.

As a matter of fact, the idea that the biting lice and the sucking lice
are related finds but little actual morphological support, although there
is a suggestion of some sort of remote relationship. The feeling of rela¬
tionship0 seems to arise chiefly from the fact that the biting lice offer
the only known source from which the sucking lice could have been derived.
The writer is not impressed by Webb's insistence upon the evidence afforded
by the tracheal system, in the light of the very profound differences in
the feeding mechanism.

The opinion is here maintained, therefore, that the sucking lice should
be recognized as the Order Anoplura.

The Characteristics of the Order Anoplura

An insect Order the members of which are, as far as known, ectoparasites
exclusively upon mammals, living in the hair, feeding upon the blood of the
host throughout their entire life cycle, and attaching their eggs to the
hairs of the host. Metamorphosis of the type commonly called hemimetabolic,
consisting chiefly of changes in size or proportions of parts, in the de¬
gree of sclerotization and in the development of the sexual apparatus.

Antennae normally five-segmented, although at times apparently only
three-segmented or four-segmented.

Mouthparts highly modified, being formed of three stylets which are re¬
tracted into a trophic sac, lying beneath the pharynx, these stylets being
protrusible .

Thorax with the three segments closely fused but recognizable, the true
notum being reduced in all segments to a narrow, median, membranous area or
at times to a median pit and slight lateral extensions which surround the
spiracles. The apparent thoracic nota formed almost entirely of subcoxal
(=pleural) elements and these at times fused entirely across the notum,
thus almost completely obliterating the true notal plates. Never more than
one pair of thoracic spiracles, these belonging to the mesothorax.

Ovipositor of the female present, but reduced to at most a pair ot flat¬
tened lobes which represent the gonopods of segment eight and a pair of
small, sclerotized areas or mere tufts of setae which probably represent
the gonopods of segment nine. Male never retaining the gonopods of segment
nine^and consequently without claspers, but usually retaining the parameres,
the genital structures being retracted into the body when at rest.

Legs modified as clasping organs by having the distal, ventral angle of
the tibia prolonged into a thumb which opposes the claw. Tarsus normally
one-segmented and at most only obscurely two-segmented. Tarsal claws al¬
ways single except in a few instances where a very doubtful second, claw¬
like structure is present on the anterior legs.

NOTES. The extraordinary mouthparts, which are among the most highly
modified that are to be found in the Insecta, are the principal basis for
the recognition of the Order as distinct from the Mallophaga. No transi¬
tion between such mouthparts and those of the Mallophaga is known to exist.
However, other distinctive characters are to be found in the thorax. In
the Mallophaga the pro thorax is always definitely separated from and mov¬
able independently of the other segments and the dorsa of at least the pro¬
thorax and the metathorax seem always to be composed of the true notal
plates. In the Anoplura the prothorax is always fused with the mesothorax
in such a manner that it cannot be independently movable, and the thoracic
dorsum throughout is composed primarily of the pleural (=subcoxal) elements.
While it is highly probable that the two groups go back to some common an¬
cestry, that ancestry- is entirely hypothetical and even if definitely known
would still not preclude their being considered as separate Orders.

The feeling that the two groups should be united into a single Order
seems to rest chiefly upon the similarities of habit and metamorphosis.
Thus, the members of both groups attach their eggs to the hairs of their
host, but exactly the same procedure is adopted by certain mites and by
flies of the family Oestridae. Webb has considered the two groups to be
closely related because of resemblances in their spiracles and tracheal
systems. No comparative study of the spiracles through various Orders was
made. Thus spiracles in some respects very- similar to those both of Ano¬
plura and Mallophaga appear in members of the families Polyctenidae and
Cimicidae of the Order llemiptera. We have here a situation very- similar to
that of the presence of "combs" of setae, such as those on the fleas, which
occur in various ectoparasitic groups but which are also found in other
forms that are not parasitic when once those forms are examined by the same
methods which are employed in the case of the ectoparasites.

It is quite true that no other group than the Mallophaga is ‘known to
which the sucking lice may be closely related, but this in itself is no ar¬
gument for combining the two groups into a single Order.

Concerning the general relationships of the sucking lice, it may simply
be noted that if there is in fact some connection with the Mallophaga then
the Anoplura are probably remotely related to the Corrodentia, since the
Mallophaga seem thus to be connected.

CHAPTER V

The Classification Within the Order Anoplura

For many years after the formal naming of the genus Pediculus by Linnaeus
all the known species of lice, both biting and sucking, were referred to
this genus. It was not until 1806 that the biting lice were removed and
placed in the genus Rtclnus. It was then not until 1816 that the sucking
lice remaining in Pediculus were in part transferred to two new genera,
Haematopinus and Pthirus. The next genus of sucking lice, Pedlcinus, was
not named until 1844 and the next, Ecdinophtdirius, not until 1871. As
late as 1880 Piaget recognized only these genera, most of the species being
referred by him to Haematopinus. In 1891 Osborn named the genus Haematopin-
oides and in 1896 the genus Kuhaematopinus, which is definitely a synonym
of Haematopinoides. During all this time — perhaps fortunately — no attempts
were made to develop a higher classification, Giebel, in 1874, placing all
the species in the family Pediculina and Piaget, in 1880, using the same
arrangement with the alteration of the family name to Pediculidae.

As late as 1904 but seven genera had been named as belonging to the
suckiug lice, and of these one was a synonym and one, Haematomyzus, is now
known not to be a sucking louse.

In 1904, Enderlein began the publication of a series of papers which
represent the first really intelligent work done on the group. He recog¬
nized the superficial character of the genus Haematopinus and hegan the
process of dismembering it. He uamed a total of eight new genera, of which
six were removed directly from Haematopinus and two were based upon new
species. At the trine when Enderlein began his work there were, according
to a count later made by Fahrenholz, but 6 ‘S known species, one ol which was
later shown not to belong to the Anoplura, and nearly 60 were referred to
the genus Haematopinus. Up to this time there was no need for a system of
classification above the generic level. Enderlein essayed the beginnings
of such a system and this was later followed by Dalla Torre and by Ferris.

Eliminating the genus Haematomyzus , the sole member of the family Haema-
tomyzidae established by Enderlein, which we now know is not a sucking
louse, Enderlein's system was as follows:

FAMILY Pediculidae Leach FAMILY Haematopinidae Enderlein
Subfamilies Subfamilies

Haematopininae Enderlein
Linognathinae Enderlein
Euhaematopininae Enderlein

Pediculinae Enderlein
Pedicininae Enderlein

FAMILY Echinophthiriidae Enderlein

In 1909, Enderlein, in a paper on the lice of marine mammals published
in the report of the Deutsche Sudpolar Expedition, Volume X, divided the
Echinophthiriidae into two subfamilies, Echinophthiriinae and Antarctoph-

thiriinae. „ it . , . „ . .

In 1908, Dalla Torre published a catalogue of the Anoplura m Wytsman s
"Genera Insectorum." This work, while useful as a first attempt at a cata-
logue, was an entirely mechanical bit of bibliography involving no knowledge
of^the insects on the part of the author and represented a none- too- thorough
search of the literature. Only 65 species were listed. The classification

adopted by Enderlein was followed. „ .
In 1916, Ferris published a "Catalogue and Host List ol the .Anoplura in

the "Proceedings of the California Academy of Sciences, (Fourth^ Series),
Volume VI. The system proposed by Enderlein was here followed. Some names
were overlooked and there was some false synonymy, but the number of known

species at this time must have been quite close to the 120 listed.

In 1929, Ewing, in a book entitled "A Manual of External Parasites,"
proposed an extended classification. This still included Haematomyzus and
the family Haematomyzidae . Excluding the Haematomyzidae the system was as
follows:

FAMILY Haematopinidae Enderlein
Subfamilies

Enderleinellinae Ewing
Hybophthiriinae Ewing
Uoplopleurinae Ewing
Linognathinae Enderlein
Neolinognathinae Ewing
Haematopininae Enderlein
FAMILY Haematopinoididae Ewing
Subfamilies

Haematopinoidinae Ewing
Hamophthiriinae Ewing

FAMILY Pediculidae Piaget
Subfamilies

Pediculinae Enderlein
Pedicininae Enderlein
Phthirpediculinae Ewing
FAMILY Phthiriidae Ewing

Contains but one genus Phthirus
FAMILY Echinophthiriidae Enderlein
Subfamilies

Echinophthiriinae Enderlein
Antarctophthiriinae Enderlein
Lepidophthiriinae Ewing

In 1931, Ferris, in an extended paper in "Parasitology," Yolume XXIII,
showed that Haematomyzus does not belong to the sucking lice. He trans¬
ferred this genus doubtfully to the Mallophaga and proposed for it the sub¬
ordinal name Rhynchophthirina.

In 1932, Bedford, in a check list of the parasites of birds and mammals
of South Africa, published in the "18th Report of the Director of Veterinary
Services of the Union of South Africa, " considered the biting lice and the
sucking lice together to constitute the Order Anoplura, with three Subor¬
ders, Mallophaga, Rhynchophthirina, and Siphunculata. Under the Siphuncu-
lata he adopted the system proposed by Ewing.

In 1936, Fahrenholz, in a paper in "Zeitschrift fur Parasitenkunde, "
Volume IX, Heft 1, proposed a classification in which he considered the
suckincr lice to constitute the Suborder Inrostrata of the Order Anoplura.

Under this Suborder he proposed the following arrangement:

FAMILY Echinophthiriidae Enderlein
Subfamilies

Echinophthiriinae Enderlein
Antarctophthiriinae Enderlein

FAMILY Haematopinidae Enderlein
Subfamilies

Haematopininae Enderlein
Linognathinae Enderlein
Polyplacinae Fahrenholz

Haematopinoidinae Ewing
Lemurphthirinae Fahrenholz
Hamophthirinae Ewing
FAMILY Neolinognathidae Fahrenholz
Containing the single genus

N eolinognathus
FAMILY Pediculidae Leach
Subfamilies

Pedicininae Enderlein
Pediculinae Enderlein

In 1946, Eichler, in "Archiv fur Naturgeschichte, " Neue Folge, Volume X,
Heft 3, adopted the name Phthiraptera of Haeckel, for an Order which in¬
cluded the three Suborders Mallophaga, Anoplura, and Rhynchophthirina. Un¬
der the Suborder Anoplura he proposed the following arrangement. If we
substitute the term "superfamily" for his "family series" we shall be more
nearly in accord with usual entomological practice.

FAMILY Echinophthiriidae Enderlein Polyplacinae Fahrenholz

Subfamilies Lemurphthirinae Fahrenholz

Echinophthiriinae Enderlein Hamophthirinae Ewing

Antarctophthiriinae Enderlein

FAMILY SERIES Pediculiformia Eichler FAMILY Pediculidae Leach
FAMILY Haematopinidae Enderlein Subfamilies

Subfamilies Pedicininae Enderlein

Haematopininae Enderlein Pediculinae Enderlein

Linognathinae Enderlein Phthirinae Ewing

In 1946, Webb, in a paper in the "Proceedings of the Zoological Society
ot‘ Loudon, " Yolume 116, Part 1, proposed a partial system ol‘ classi Ideation
of the sucking lice "based solely on differences in spiracle structure "
as he himself said, but influenced to a considerable degree by considera¬
tions based on the distribution of the lice according to hosts. The system
as there presented was only partial, since some genera were not placed in it.
Webb's system, as far as it has been carried at the time of the present
writing, is as follows: The biting lice and sucking lice are uuited in the
Order AnopLura and are regarded as Suborders, the names Mallophaga and
Siphuuculata being employed for them. The Siphunculata, so lar as treated,
are divided into six families — Docophthi ri idae, Pediculidae, Eulinognath-
idae, Linognathidae, Haematopinidae, and Echinophthiri idae . No subfamilies
are named.

In the opinion here held this classification is in part correct. But in
part it also rests upon some very peculiar concepts of taxonomy. Thus the
louse of elephants, Haematomyzus elephant is, is not only regarded as a
sucking louse but is even placed in the family Haematopinidae on the basis
of spiracle structure alone!

In 1949, Hopkins, in the "Proceedings of the Zoological Society of
London," Volume 119, recognized the biting and sucking lice together as
constituting an Order Phthiraptera with the three Suborders Anoplura, Mal-
lophaga, and Rhynchophthirina. Under the Anoplura he recognized but three
families — Pediculidae, Haematopinidae, and Neolinognathidae.

In attempting to develop an intraordinal classification ol the sucking
lice, here considered to be the Order Anoplura, consideration has been
given only to morphology. It is here held that the problems associated with
the distribution of the lice on their various hosts should be approached
only after conclusions have first been reached by way of morphological
studies. It may be very agreeable in the end to find that the conclusions
drawn from the lice themselves are in accord with those drawn 1 rom their
hosts and if the two sets of conclusions serve to confirm each other we may
be justified in feeling that some approach to truth has been achieved. But
to use A to develop B and then to turn about and use B to develop A is a
process that presents certain illogical aspects. This is especially true
if a priori ideas concerning A and B have been employed. On the other
hand, if A has been competently and logically developed and then B is found
to agree closely with A there may very well be situations where, a point in
A beiug doubtful, B may at least be used as a tentative aid in testing A or
in supplementing it.

Thus in developing a classi fication of the sucking lice, if on morpholog¬
ical grounds a certain set of relationships is clearly indicated and then a
definite correlation with the distribution of the lice according to hosts
appears in harmony with these indications, we may be justified in employing
the evidence of distribution to supplement the evidence or morphology where
the latter is inadequate or confused. But we are certainly not justified
in employing the evidence trom distribution if it completely denies or is
otherwise not in harmony with clear evidence from morphology. Especially
is this true when the evidence from distribution is itself based upon ques¬
tionable concepts. , . , . , .

For example, the genus Polyplax is a large group which is confined to

rodents of the family Muridae, except for one or two species which occur on
shrews. To say that since the shrews and the Muridae are not closely re¬
lated to each other lice of the genus Polyplax should not occur on shrews
and that therefore these lice actually from shrews do not belong to the
renus Polyplax, despite the morphological evidence, is an utterly illogical
procedure'. To say that such genera as Schizophthirus and Ancistroplax can¬
not be related to the genus Hoplopleura because of the wide differences in
the assumed relationships of the hosts would be to deny the plain evidence

of morphology.

But in the case of the Enderleinellinae we have a different situation.
The genus Enderleinellus, which is of considerable size, is known only from
Sciuridae. There are two peculiar genera, M icrophthirus and Verneckia, which
in certain respects are like Enderleinellus but which depart from that genus
in the lack of a certain structure that we are inclined to accept as of
morphological significance. But in certain other respects these two genera
are like Enderleinellus and all three generaareas far as known confined to
the squirrels. The totality of the evidence thus indicates that the absence
of the structure in question is probably not sufficient to indicate that
the two genera which lack it should be excluded from the Enderleinellinae.
We are not here denying the evidence from morphology, we are merely supple¬
menting it where it is weak, for experience in other fields of taxonomy in¬
dicates that the loss of a structure in the course of evolution may occur
without disturbing or invalidating the evidence of relationship that is de¬
rived from other sources. What it does disturb is really nothing more than
the ease with which keys for identification may be developed.

In attempting to develop a classification of the sucking lice certain
considerations should be kept in mind.

1. This group is probably something of a remnant of what it once may
have been. The living mammals represent but a very small part of all the
species of mammals that have existed in the past. Let us reflect upon the
number of proboscideans that are known only as fossils. What were their
parasites? The single louse-like parasite of the elephants is all that is
left to accompany the dwindling line of its hosts. The same situation may
very well hold in regard to other lice.

2. Being a very small, very' specialized, and perhaps remnant group, we
have left but few or none of the connecting links that one may hope to find
in the Orders of insects that still contain thousands of species.

3. There has evidently been a very large amount of evolution by loss
among the Anoplura and this leaves us with a rather small complement of
structures with which to work.

4. We still know probably not more than half of the species of sucking
lice that exist in the world.

5. Under these conditions we have but limited bases for judgment as to
the relationships of such forms as Hybophthirus notophallus, the louse of
the aard vark; Haematopinoides squamosus , the louse of certain moles; the
two known species of Neolinognatlius from elephant shrews; as well as some
other peculiar forms. Each of these genera contains the solitary' or almost
solitary representative of some ancient line and we have no basis for judg¬
ment as to whether its peculiarities are merely those of an individual spe¬
cies or were common to the members of what may once have been a much larger
group .

Actually, whatever basis for classification we may adopt, the grouping
of the genera of the sucking lice is not at all clear. Certainly no satis¬
factory arrangement can be obtained by focusing attention upon a single set
of characters. We must take into consideration the totality of the charac¬
ters which are available.

The opinion is here held that in the light of our present knowledge of
the Anoplura any system of classification within the Order which elaborate¬
ly divides it into superfamilies, families, subfamilies, tribes, subtribes,
genera, and subgenera, can be nothing more than pretentious nonsense. Such
minute subdivision can very well wait until a substantially larger propor¬
tion of the species is known.

In the light of these considerations the system here adopted will be
rather conservative, only families, subfamilies and genera being employed.
It is presented with a full realization of the possibility that it may very
well be inadequate and at some points may actually be wrong. However, there

is the thought that the number of species concerning which error is most
likely to exist constitutes only a small part of the whole. Thus, there
may be question concerning the seven species referred to the Fedicininae
and to certain isolated species. But the great mass of the species and
genera fall into clearly recognizable groups.

The genera of Anoplura are reasonably clear and uncomplicated, except as
they may be complicated subjectively by workers who see a new genus in
every slight departure from the normal form. These genera to a consider¬
able degree fall into groups, the central pattern of which is rather evi¬
dent, but which becomes clouded as we pass out from it. In fact, around
their peripheries these groups become so misty that no very definite out¬
lines can be detected and it is here that the principal difficulties arise.
To name families that cannot be defined is to create difficulties lor all
future workers, to overemphasize the differences that exist and exaggei'ate
the boundaries among these differences.

At an earlier time, when 1 ittle or nothing was known about the morpholo^
of these insects, certain ideas concerning their classification seemed valid
which now appear at least dubious. Thus it was assumed that the presence
of a distinct lens marking each compound eye separated the Fediculidae from
all the other lice and defined a family. We now know that such lenses are
present in various other forms which seem not to be closely related to
Pediculus and this once apparently beautifully defined family becomes ex¬
tremely difficult to define. Evolution by loss has occurred so extensively
in the Anoplura that the persistence of a structure, may indicate only the
retention of what was once a widespread character in all the ancestors of
the group. It is therefore difficult to properly evaluate the remnants
that are left.

One other quite secondary, but nevertheless important, problem arises.
This has to do with the names which should be employed for the families and
subfamilies that are recognized. This is a problem which is inherently
difficult and which, probably because of this difficulty, is not even
touched upon by the "International Rules of Zoological Nomenclature."

Should priority rule? If so, what constitutes' priority? Does it date
from the first recognition of the existence of a group in any status what¬
soever or from the first unequivocal employment ot a group name with a par¬
ticular ending? Are family and subfamily or any lower category names such
as tribes and^ subtribes to be regarded as nomenclatorially equivalent, m
the nammer in which generic and subgeneric names are equivalent? What de¬
termines the type of any category? Should it be the oldest genus referred
to the category — as some maintain — or should it be the genus upon which the
category name was first based — as others maintain?

And who is to be cited as the author of a group name— if this recogni¬
tion must willy-nilly be accorded to someone? Shall it be the ^person who
first recognized the group and employed a group name of any^rank? Or shall
it be the person who used the name which is now employed? Thus, is the
author of a subfamily name to be regarded as the author of the name if it
is elevated to family rank, or shall it be the person who first used the
name to characterize a family? If a subfamily is raised to 1 am ily rank and
then reduced again to a subfamily of some other family, who is the author
of the name? Or should we refuse to bother about the name of the author of

^There are no fixed answers to these problems and in the absence ol an¬
swers one is left very' much to follow whatever course seems reasonable.

The writer of these lines has puzzled long over all these questions and
has come to no answers to any of them that are entirely satisfactory to
himself. As far as classification is concerned, there is on the one hand
the undesirability of obscuring facts by too comprehensive groupings or on
the other hand of going beyond all reasonable limits by the indiscriminate

naming of groups that cannot be supported by any cogent reasons.

As to nomenclature, the stand here adopted is in part based upon prior¬
ity in actual recognition of a group, whatever category may have been as¬
signed to it, and in some degree upon arbitrary action in refusing to rec¬
ognize a group — such as the Haematopinoididae — which would assign a family
name to a large group on the basis of a very aberrant member.

As to authors' names, it is here held that the matter is too unimportant
to bother with, or above all to argue very much about one way or another.
The names of authors are indicated in accompanying discussions but not con¬
joined with the names of the groups in section headings.

THE SYSTEM HERE EMPLOYED
Family Echinophthiriidae

This family, which was established as such by Enderlein in 1904, is re¬
tained. It was united with the Haematopinidae by Hopkins but this step is
regarded as erroneous on morphological grounds. Two subfamilies — Echinoph-
thiriinae and Antarctophthiriinae — were recognized by Enderlein and accept¬
ed by most later workers, but are here rejected since it does not appear
that anything is gained by recognizing them.

Family Haematopinidae

First established by Enderlein to contain the vast majority of the suck¬
ing lice and later accepted by all workers with various restrictions. It
is here accepted for two genera, Haematopinus and Pecaroecus.

Family Hoplopleuridae

This family contains within it the genus Haem.atopinoid.es which, under
the synonymic name Euhaematopinus, was placed by itself in the subfamily
Euhaematopininae of the family Haematopinidae by Enderlein and later, under
its correct name Haemal opinoides, placed by Ewing in the family Haematopin¬
oididae. This family is rejected as zoologically unjustified and referred
to the Hoplopleuridae. It may very well be argued, and perhaps correctly,
that the family name Haematopinoididae should be employed for the family
Hoplopleuridae, to which it is here referred, but it is a very aberrant
form which is quite unsuited to be used as the type of the family.

The group of which H op l op l eur a is here taken as the type was recognized by
Ewing in 1929 as the subfamily Hoplopleurinae of the family Haematopinidae.
Since at—the same time Ewing recognized the subfamily Ender leine.llinae, it
is necessary to make a choice between these two names for the family here
considered to include the two groups. Hoplopleura is chosen as being more
nearly central than is Enderleinellus .

Subfamily Enderleinellinae Ewin»r

v to

Established by Ewing in 1929 as a subfamily of the Haematopinidae. Here
accepted as a subfamily of the Hoplopleuridae, to contain three genera.

Subfamily Hybophthirinae Ewing

This group was named by Ewing in 1929 as a subfamily of the Haematopinidae.
Webb has considered the members of this group to be closely related to
Haematopinus and has placed them in the family Haematopinidae, but since he
also placed in this family the genus Haematomyzus — which is not even an
Anopluran — the importance of the spiracular characters which he employed
may be somewhat discounted.

The three genera here placed in the Hybophthiri inae are, in their tree
paratergal pLates of the abdomen, much more closely similar to the members
of the Hoplopleur idae than to any other group. They agree among themselves
in the possession of a claw-like structure arising beside the claw on the
anterior legs and in this appear to be related most closely to each other.

Subfamily Pedicininae Enderlein

It is concerning this group that there will probably be the greatest
difference of opinion and the writer does not expect agreement with the
point of view here adopted.

The group was first recognized by Enderlein as a subfamily of the tamily
Pediculidae, which was restricted to Pediculus, Pthirus, and Pedlcinus.
The genus Pedlcinus has, ever since, been placed as a subfamily ol the
Pediculidae because of two factors. One ol these factors is undoubtedly
that of the feeling that anything occurring on the Primates should be re¬
lated to Pediculus. The other factor is that of the common possession of
eyes by Pediculus and Pedlcinus and this character was employed as the dis¬
tinguishing mark of the Pediculidae. But in the light ol' what is now known
it does not appear that this character by itself is especially significant,
since it is known that genera such as Pecaroecus and Hicrothoracius — which
certainly do not belong with Pediculus — have eyes and they can even be dem¬
onstrated in Haematopinus although they are not there so evident.

In fact, present evidence indicates merely that the presence of eyes
represents merely the retention of a primitive character, once common to
all the sucking lice and their ancestors.

If we remove the eyes from consideration, the remainder of the body in
Pedlcinus contraindicates any association with Pediculus. In fact, with the
eyes removed all the characteristics of the body seem much closer to what
is found in the Hoplopleuridae . The free paratergal plates and the absence
or great reduction of the gonopods of segment nine in the female are more
nearly in harmony with what appears in this family.

We might raise Pedlcinus to family rank, but this is merely to increase
the number of families by emphasizing the difference in one structure.

With these thoughts in mind the author has decided to "take the bull by
the horns" and remove the genus Pedicinus from close association with
Pediculus , assigning it to the family with which it shares the majoritj ol
its features, regardless of whether or not it ouiht to be related to the
Pediculidae .

Subfamily Polyplacinae

Established by Fahrenholz in 1936 as a subfamily of the Haematopinidae .
It is here considered to contain the genus Eamophthirus , which was employed
by Ewing in 1929 as type of his subfamily Ramophthirinae of the family
Haematopinoididae and thus on grounds of priority it might be argued that
the name Ramophthiriidae should be employed for it. However, the genus
Hamophthirus is very badly described and quite unsuitable to stand as the

type of any group. , , „ , . ,

It contains also the genus Lemurphthirus which was employed by fahrenholz

in 1936 as type of the subfamily Lemurphthirinae of the family Haematopin¬
idae. This subfamily is here rejected.

Family Linognathidae

First recognized by Enderlein as the subfamily Linognathina^ of the tam-
ily Haematopinidae and retained in this position by later workers. It is
here elevated to family rank.

Family Neolinognathidae

This group was first recognized by Ewing as the subfamily Neolinognathinae
of the family Haematopinidae and later raised to family rank by Fahrenholz.

Family Pediculidae

This family was apparently established under a family name, with the fam¬
ily ending as now employed, hy Piaget in 1880, although it had earlier been
recognized by Giebel under the name Pediculina. It is here utilized for
two genera only, the genus Pedicinus — which has usually been assigned to the
Pediculidae — here being regarded as belonging rather to the Hoplopleuridae.

CHAPTER VI

Review of the Families, Subfamilies, Genera and Species of the Anoplura
Family ECHINOPHTOIRllDAE Enderlein

1904. Enderlein, Zoologischer Anzeiger 28:136. (Names the family.)

1909. Enderlein, Deutsche Siidpolar Expedition 10:506. (Divides the family

into two subfamilies, Echinophthiriinae aud Antarctophthiriinae.)

1910. MjSberg, Arkiv f8r Zoologi 6:177. (Names the family Lepidophthiri-

idae . )

1916. Ferris, Proceedings of the California Academy of Sciences (Series 4)
6:180. (Rejects the family Lepidophthiriidae as a synonym of Ech-
iuophthiriidae. )

1928. Freund, Die Tierwelt der Nord-und Ostsee, Teil XI dx. (Reviews the

family and accepts the two subfamilies Echinophthiriinae and Lepi-
dophthirinae .)

1929. Ewing, A ManuaL of External Parasites, page 148. (Recognizes three

subfamilies, Echinophthiriinae, Antarctophthiriinae, and Lepidoph-
thiriinae. )

1936. Fahrenholz, Zeitschrift fiir Parasite nkunde 9 : 56 • (Recognizes the two
subfamilies Echinophthiriinae and Antarctophthirinae. )

1946. Eichler, Archiv fur Naturgeschichte , Neue Folge, 10:345-398. (Recog¬
nizes the family with two subfamilies, Echinophthiriinae and Ant¬
arctophthiriinae and places the family in the "Family Series Echi-
nophthiri formia. ")

DESCRIPTION OF THE FAMILY. .Anoplura which occur exclusively upon marine
mammals of the Suborder Pinnipedia of the Order Carnivora. Eyes not exter¬
nally evident. Antennae four- or five-segmented. Body more or less thick¬
ly beset with setae which are in some species modified into scales, in
others only somewhat flattened and in at least one species normally cylin¬
drical. Abdomen never with sclerotized tergal, paratergal, or sternal
plates, membranous or leathery. Abdominal spiracles of a distinctive type,
with a lom>', slender, and more or less membranous atrial chamber, the walls
of which donot bear transverse markings. Female with the gonopods of seg¬
ment ei^ht never forming free lobes. Males with the genital sac forming a
flattened, median plate, alongside which lie the flattened parameres.
Thorax never with a sternal plate which is apically or marginally free, al¬
though at times with an irregular sternal sclerotization. Middle and pos¬
terior legs always with a very stout tibiotarsus, in which the tibiotarsal
division Is scarcely or not at all evident. Anterior legs usually small
and slender, with the tibiotarsal division evident, but in at least one

species similar to the others. ... . ,

NOTES. Nothing is to be gained by dividing this family into two sub¬
families, and still less by dividing it into three as has been proposed.
Division on the basis of the presence or absence of scales is contraindi¬
cated by the fact that Proechinophthirius fluctus (Ferris), which lacks
scales is in other respects very similar to Antarctophthtrus callorhini
(Osborn) in which the vestiture of scales is less than that to be found in
other members of the latter genus. A division on the basis of number of
antennal segments also is contraindicated, since such a division would cut
directly across any division on the basis of the presence or absence of
scales/ Nor is anything to be gained by division on the basis of the dis
tribution of the species for, upon whatever basis such division may be made,
it does not follow the relationships of the hosts.

Key to the Genera of ECHINOPHIHIRIIDAE

1. Antennae ^segmented . ANTARCTOPHTHIRUS

Antennae 4-segmented . 2

2. Abdomen with scales . LEPIDOPHTHIRUS

Abdomen without scales . - . 3

S. Leers all of same size and shape, with stout ti bio tarsus and claw .

. ECHINOPHTHIRIUS

Anterior legs much smaller than the others and with small, slender claw
. PKOECHINOPHTHIRIUS

Genus ANTARCTOPHTHIRUS Enderlein

1906. Antarctophthirus Enderlein, Zoologischer Anzeiger 29:661.

1910. Arctophtirius Mjoberg, Arkiv for Zoologi 13:177.

1934. Antarctophthirus, Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 7:484.

1941. Achimella Eichler, Archiv fur Naturgeschichte, Neue Folge, 10:375-
GENERIC TYPE. The typ.e of Antarctophthirus, by original designation, is
Antarctophthirus oimorhini Enderlein. The type of Arctophtirius Mjdberg
is, by original designation, Eaematopinus trichechi Bohemann. The type of
Achimella is, by original designation, Eaematopinus callorhini Osborn.

CHARACTERS. Echinophthiriidae with five-segmented antennae. Anterior
legs small and with slender claw; middle and posterior legs very large and
stout and with stout claw. Body more or less beset with flattened, scale¬
like setae.

NOTES. There is no justification for the recognition of the genus Arc¬
tophtirius Mjoberg. There is perhaps some justification for the recogni¬
tion of Achimella, but that evidence is not considered sufficient.

Key to Species of ANTARCTOPHTHIRUS

1. Thoracic sternum beset only with slender setae; scales on the abdomen

very few, arranged more or less in patches on the dorsum of the abdo¬
men and on the posterior portion of the venter; occurring on the

Alaska fur seal . CALLORHINI

Thoracic sternum bearing scales; scales very abundant on the abdomen.. 2

2. Thoracic sternum with a few long setae on its posterior border . 3

Thoracic sternum without long setae along its posterior border . 4

3. Scales of the abdominal dorsum all very uniform in size and shape,

ovoid, the apex slightly pointed; occurring on Phocarctos, Eumeto-

pias, and Zalophus . MICROCHIR

Scales of the abdomen (according to Enderlein) elongated-oval and of
various sizes; occurring on Lobodon . L0B0D0NTIS

4. Ventral side of the head beset with small, flattened, oval setae in ad¬

dition to scales; occurring on Ogmorhinus . 0GM0RHINI

Ventral side of the head with long hairs or with minute hairs only.... 5

5. Ventral side of the head with many long hairs posteriorly . TRICHECHI

Ventral side of the head with only 3~4 minute hairs posteriorly MAWSONI

Antarctophthirus callorhini (Osborn)

1899- Eaematopinus callorhini Osborn, In The Fur Seals and the Fur-Seal
IsLands of the North Pacific Ocean 3:553; figure 1.

1915- Antarctophthirus monachus Kellogg and Ferris, Anoplura and Mallo-
phaga of North American Mammals, Stanford University Publications
(no volume number), page 49; text figures 17A and 18; Plate III,
figure 4.

1934. Antarctophthirus cal lor hint (Osboru), Ferris, Contributions Toward
a Monograph of the Sucking Lice, Fart 7:495; figures 289, 290.

1941. Achlnella cal lorhini (Osborn), Eichler, Archiv fur Naturgeschichte,
Neue Folge, 10:275-

HOSTS AND DISTRIBUTION. The types of Osborn's Haemal op in us cal lorhin i
were from Callorhirius alashanus from the Pribilof Islands. The specimens
upon which the name monachus were based came from Osborn as a loan but bore
no data other than as from "seal," but they were in ail probability from
the type lot. Since that time a tew more specimens have come to hand lrom
the type host on St. Paul and Pribilof Islands in the Bering Sea.

Antarctophthirus lobodoutis Enderleiu

1909. Antarctophthirus lobodontis Enderlein, Deutsche Siidpoiar Expedition
10:510; Figures KK, NN.

HOSTS AND DISTRIBUTION. Known only from the original record from Lobo-
don care inophagus , Booth Wandel Island in the Antarctic.

Antarctophthirus mawsoni Harrison

1937. Antarctophthirus mawsoni Harrison, Australasian Antarctic Expedi¬
tion, 19 11-1914, Scientific Reports, Series C-Zoology and Botany
2:1:11; figure 1.

HOSTS AND DISTRIBUTION. From Ommatophoca rossi, King George V Land in
the Antarctic.

Antarctophthirus microchir (Trouessart and Neumann)

1888. Schinophthir ius microchir Trouessart and Neumann, Le Naturaliste 10:
80; figures.

1906. Antarctophthirus microchir (Trouessart and Neumann), Enderlein,
Zoologischer Anzeiger 29:663; figures 3, 4.

1934. Antarctophthirus microchir (Trouessart and Neumann) , Ferris, Contri¬
butions Toward a Monograph of the Sucking Lice, Part 7:489; fig¬
ures 285, 286.

1939. Antarctophthirus microchir californianus Fahrenholz, Mitteilungen
aus dem entomologischen Verein Bremen, page 42.

HOSTS AND DISTRIBUTION. Types from Phocarctos hookeri, Aukland Island.
Recorded also from Zalophus californianus and Eumetopias jubata from the
coast of California.

NOTES. The supposed variety californianus was named by Fahrenholz pure¬
ly upon the basis of differences in the illustrations presented by Enderlein
and Ferris. These differences are of an order inevitably associated with
differences in the preparation of specimens and involve no actual morpho¬
logical characters. Actually the agreement between specimens from Califor¬
nia and the beautiful illustrations given by Fkiderlein is extremely close.
The supposed variety is here rejected.

Antarctophthirus ogmorhini Enderlein
Figure 31

1902. Echinophthirus setosus (Burmeister) , Rothschild, Report of the
Southern Cross Expedition, page 224. (Misidenti fication >

1906. Antarctophthirus ogmorhini Enderlein, Zoologischer Anzeiger 29:662;
text figures.

1909. Antarctophthirus ogmorhini Enderlein, Enderlein, Deutsche sudpolar
Expedition 10:509; figures 174, 175, 181, 182.

1934. Antarctophthirus ogmorhini Enderlein, Ferris, Contributions Toward a

Antarctophthirus ogmorhini (Enderlein)

Figure 31

Monograph of the Sucking Lice, Fart 7:486; figures 282, 283-
1937. Antarctophthirus ogmorhlnt Enderlein, Harrison, Australasian Antarc
tic Kxpedition, 1911-1914, Scientific Reports, Series C, /oology
and botany 2:1:11.

HOSTS AND DISTRIBUTION . Described as from Otnorhinus leptonyx, Victoria
Land and booth Waude 1 Island in the Antarctic. This is apparently now
placed in the genus Hydrurga.

NOTES. The accompanying illustration was made from the types in the
British Museum.

Antarctophthirus trichechi (Bohemann)

186 5 . Haematoplnus trichechi Bohemann, Vetenskaps Akademie Forhaudl inger,
Kobnhaven 22:557; Plate 35, figure 2.

1909. Antarctophthirus trichechi (Bohemann), Enderlein, Deutsche Siidpolar

Expedition 10:512; figures 172, 173. 185-188. _

1910. Arctophtirus trichechi (Bohemann), Mjoberg, Arkiv tur Zoologi 1U : I3 :

178; figures 90-92. .

1934. Antarctophthirus trichechi (Bohemann), Ferris, Contributions loward
a Monograph of the Sucking Lice, Fart 7:492; figures 287, 288.
HOSTS AND DISTRIBUTION. Recorded by various authors from the walrus,
Odobaenus rosmarus and Odobaenus obesus, from various places in the Arctic.

Genus ECHINOPHTHIRIUS Giebel

1871. Bchinophthirius Giebel, Zeitschrift fur die gesamten Naturwissen-
schaften 37:177-

1934. Bchinophthirius, Ferris, Contributions Toward a Monograph ol the

Sucking Lice, Part 7:475- . , u. .

1946. Bchinophthirius, Hopkins, Annals and Magazine of Natural History,
Series 11, 12:566. (Bears date of 1945, but published May, 1946)
GENERIC TYPE. Pediculus phocae Lucas, which is considered to be a syno¬
nym of Pedlculus horridus von Olfers.

CHARACTERS. With four-segmented antennae. Legs all ol essentially the
same size and form, stout and with stout claw. Abdomen thickly beset with

stout, flattened, but not scale-like, setae.

HOSTS. Occurring as far as known only on seals ot the genera Halichoerus

and Phoca, of the family Phocidae.

NOTES. Ferris 11934) reviewed a considerable amount ol material ann
concluded that this genus contains but one species. Hopkins (1946) sug¬
gested although apparently not on the basis of an examination ol specimens,
that "anaLogy with other genera suggests the possibility that it may prove
necessary to divide the genus again into several species or subspecies.
After reviewing such material as is available, the opinion is here still
held that the genus contains but one species, as far as morphological evi¬
dence goes, although at present nothing certain can be said concerning the
supposed subspecies from seals in Lake Baikal.

Echinophthirius horridus (von Oilers)

Figures 32, 33

1816.

1834.

1838.

1857-

'edlculus horridus von Olfers, De vegetativis et animatis corporibus
animatis reperiundis commentarius, Part 1, page 84. (Fide Fahren

•edlcuLs phocae Lucas, Guerin's Magazin de Zoologie 4:Classe IX;

Plate 121. „ x o •

> ediculus setosus Burmeister, Genera Insectorum. Species 12.
’edlculus annulatus Schilling, Gurlt, Archiv fur Naturgeschichte 23:

281.

Echinophthirius horridus (von Olfers)

Figure 32

1874. Echinophthirius setosus (Denny) (sic), Giebel, Insecta Epizoa, page
42.

1880. Echinophthirius setosus (Lucas), Piaget, Les Pediculines, page 656;
Plate 54, figure 1.

1886. Echinophthirius iroenlandicus Becher, In Die Internationale Polar-
forschung 1882-1883, Beobachtungs Ergebnisse 3:60; Plate 5, fig* 1*
1897* Echinophthirius sericans Meinert, Vetenskabelige Meddelser Kjobenhavn

58:177*

1919. Echinophthirius horridus (von Olfers), Fahrenholz, Jahresbericht des
Niedersachsischen zoologischen Vereins zu Hannover 5-10:22.

1928. Echinophthirius horridus (von Olfers), Freund, In Die Tierwelt der
Nord- und Ostsee, Teil XI dj:6; figures 1, 2.

1934* Echinophthirius horridus (von Olfers), Ferris, Contributions Toward
a Monograph of the Sucking Lice, Part 7:475; figures 277, 278.
1935* Echinophthirius horridus baikalensis Ass, Travaux de la station lim-
nologique de Lac Baikal 6 : 21:25-29 ; 5 figures.

1946. Echinophthirius horridus (von Olfers), Hopkins, Annals and Magazine
of Natural History, Series 11, 12:5 66. (Bears date of 1945, but
published May, 1946)

female genitalia male genital plate male genitalia

Echinophthirius horridus (von Olfers> Figure 33

HOSTS AND DISTRIBUTION. Described by von Olfers as from Phoca vitulina.
The type of phocae Lucas was said to have been taken from the same host
species in a zoological garden in Paris. The type of ser icons Meinert was
from Phoca iroenlandica and the type of iroenlandicus Becher was ascribed
to the same host. The species was recorded by Ferris from Phoca vitulina
from the Shetland Islands, from Scotland, and from specimens taken at the
Hamburg Zoological Garden; from Phoca hispida in the Beaufort Sea off
Alaska; from Phoca richardii at Pacific Grove, California. The species has
been recorded on various occasions from Phoca v itulina, Phoca v ar ieiata,
and Halichoerus ir’jphus from the North Atlantic. Freund recorded specimens
from Phoca baikalensis from Lake Baikal.

NOTES. Hopkins has reviewed the literature and has pointed out that the
name annulata is a nomen nudum and that Burmeister's name setosus was pro¬
posed merely as a substitute for phocae Lucas.

The status of the supposed species iroenlandicus is somewhat obscure.
Ferris has examined specimens from "Greenland seal," but it is possible
that this label meant nothing more than "a seal from Greenland." Mjoberg
recorded specimens from Phoca iroenlandica which he considered to be
horridus. Freund, who reviewed the whole subject, considered it very ques¬
tionable that a distinct species occurs on this host. It is here held that
the evidence is all against the distinctness of the species Echinophthir ius
iroenlandicus Becher.

Freund examined specimens from Phoca baicalensis which he considered to
belong to horridus, this being anterior to the description of the supposed
subspecies baikalensis from this host. We may therefore consider this sup¬
posed subspecies to be a synonym of horridus.

As already pointed out in connection with the discussion of the genus,
the opinion is here held that there is no evidence to justify the naming of
more than one species of this genus.

Genus LEPIDOPUTHIRUS Enderlein

1904. Lepidophthirus Enderlein, Zoologischer Anzeiger 28:44.

1934. Lepidophthirus, Ferris, Contributions Toward a Monograph of the Suck¬
ing Lice, Part 7:498.

GENERIC TYPE. Lepidophthirus macrorhini Enderlein, the only included
species .

CHARACTERS. Echinophthiriidae with four-segmented antennae. Anterior
legs much smaller than the others and with slender claw; middle and poster¬
ior legs very large and stout, with stout claw. Body very thickly beset
with setae of various sizes and shapes, these mostly more or less flattened
and the dorsum of the abdomen also very thickly beset with scales.

Lepidophthirus macrorhini Enderlein

Figures 34, 35

1904. Lepidophthirus macrorhini Enderlein, Zoologischer Anzeiger 28:46;
figures 1-5*

1929. Lepidophthirus macrorhini Enderlein, Bedford, Director of Veterinary
Services, Union of South Africa, Report 1^:607.

1934. Lepidophthirus macrorhini Enderlein, Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 7:499; figures 291, 292.

1937. Lepidophthirus macrorhini Enderlein, Harrison, Australasian Antarc¬
tic Expedition, 1911-1914, Scientific Reports, Series C-Zoology
and Botany 2:Part 1: 13-

HOSTS AND DISTRIBUTION. Known only from the southern sea elephant or
elephant seal, from Kerguelen Island and also from near Cape Town, South
Africa, and from Macquarie Island.

Lepidophthirus macrorhini Enderlein

Figure 34

Lepidophthirus macrorhini Erderlein

genitalia of male

Figure 35

Genus PROECH INOPHTH 1 RIUS Ewing

1923. Proec hlnophthir i us Ewing, Journal of the Washington Academy of
Sciences 13:149.

1934- Proechinophthirius, Ferris, Contributions Toward a Monograph ol' the
Sucking Lice, Part 7: 48U.

GENERIC TYPE, Echinophthirlus fluctus Ferris, the only included species.
CHARACTERS. Echinophthiriidae with four-segmented antennae. Thorax
with uo sclerotized sternal areas. Abdomen beset with setae ol various
shapes and sizes, but none flattened and none being scale-like. Anterior
legs smalL and with weak, slender claw; middle and posterior legs large and
stout, with stout claw.

Proechinophthirius fluctus (Ferris)

Figure %

1916. Echinophthirius fluctus Ferris, Entomological News 27:366; figure.
1923. Proechinophthirus fluctus (Ferris), Ewing, Journal of the Washington
Academy of Sciences 13:149*

1934. Proechinophthirus fluctus (Ferris), Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 7:481; figures 279. 280, 281.
HOSTS AND DISTRIBUTION. The types of the species were taken from the
stuffed skin of an Otariid pup in the collection of Stanford University,
this bearing no data. The pup was identified as being that of a Stellar s
sea lion, Eumetopias jubata, but this may have been an error. The species
has since been taken from the Alaskan fur seal, Callorhinus alashanus, on
St. Paul Island and on the Pribilof Islands in Bering Sea.

Family HAEWATOPINIDAE Enderlein

1904. Enderlein, Zoologischer Anzeiger 28:136.

DESCRIPTION OF THE FAMILY. Anoplura in which the paratergites of the
abdominal segments are strongly sclerotized and form lateral lobes or prom¬
inences but are never marginally free from the body wall. Spiracles with
the atrium beset internally with numerous points. Legs all of substantial¬
ly the same size and with claws of the same size and shape. Clearly de¬
fined eyes present or absent, if absent the head with pronounced ocular
points just posterior to the antennae. Thorax always with a well-defined
notal pit and with a pair of sternal, apophyseal pits. Abdomen usually
with the derm of the dorsum at least very' finely wrinkled, at times slight¬
ly sclerotic, and with sclerotized areas, which, while somewhat more sclero¬
tized than the surrounding derm, are defined more by the absence of this
minute wrinkling. Gonopods of segment eight of the female always well de¬
veloped. Genitalia of the male never with free parameres. Female apparent¬
ly without spermatheca. .

NOTES. At one time this family was considered to include the great ma
iority of all the sucking lice, but it has been progressively reduced until
here it is considered to contain but two genera, Baemtopinus and Pecaroecus,
the latter having but one species. Webb (1936) considered it to include
four genera — Haematopinus , Baemtomyzus, Bybophthirus, and Scipio and pos¬
sibly also the genus Mlcrothoracius. Baemtomyzus, as has already been and
cannot too strongly be — emphasized, does not belong in this Order and the
present writer is unable to see any special affinity between Baemtopinus
and the two rrenera Bybophthirus and Scipio. There has been some temptation
to include Mlcrothoracius in the family, but this has finally been resisted.

Actually the family is very difficult to define, unless it be restricted
to the single genus Baemtopinus . In the past, when the presence of deiin

f/cnitalia of female

Figure 36

ite eyes was recognized only in Pedlculus, Pthirus, and Pedictnus, the
problem was not difficult, hut now that well-defined eyes with a distinct
lens are known to occur in Pecaroeus and morphologically true — although ex¬
ternally scarcely recognizable — eyes have been established in Haenatopinus,
there remains little by which the Pediculidae and the Haematopinidae can be
separated. Perhaps even yet it has not been possible to tree our thinking
from the presupposition that the classification of the lice should follow
the phylogeny of the hosts. This matter will be considered further in con¬
nection with the discussion of the Pediculidae.

Key to the Genera of Haematopinidae

Head with pronounced ocular points posterior to the antennae. . . HAEM ATOP 1NUS
Head without ocular points . PECAROECUS

Genus HAEM ATOP IN US Leach

1815- Haenatopinus Leach, Edinburgh Encyclopaedia, Supplement 1:24.

1842. Haenatopinus, Denny, Monographia Anoplurorum Lritanniae, page 24.
1880. Haenatopinus, Piaget, Les Pediculines," page 6 33 •

1904. Haenatopinus, Enderlein, Zooiogischer Anzeiger 28:138.

1933* Haenatopinus, Ferris, Contributions Toward a Monograph of the Sucking
Lice, Part 6:419.

1942. Haenatopinus , Webb, Proceedings of the Zoological Society of London
118:578.

GENERIC TYPE. Pediculus suis Linnaeus.

CHARACTERS. Haematopinidae in which the head is produced into distinct
ocular points posterior to the antennae, but with distinct eyes lacking.
Legs with a prominent, apically free, and more or less sclerotized process
on the ventral side between tibia and tarsus.

NOTES. The members of this genus occur on hosts of the families Suidae,
Bovidae, and Cervidae of the Order Artiodactyla and the family Equidae of
the Order Perissodactyla.

Key to Species of HAEMATOPINIJS

The species breviculus is omitted from this key because of the inadequate
description.

1. Female with the gonopods of segment eight slender, apically acute,
curved and almost sickle-shaped; known from Equus burchelli in Africa

. ACUT1CEPS

Otherwise . ;2

Female with the gonopods of segment eight elongate and narrow, their
mesal margins straight and divergent; occurring on Cervus unicolor in

India . UONGUS

Otherwise . 3

Genitalia of the male with the penis strongly asymmetrical, strongly
sclerotized at one side, this sclerotization much swollen and forming

a large knob or hook at its base . 4

Genitalia of the male with the penis otherwise formed, V- or Y-shaped

if present . . .

Female with a deeply pigmented, somewhat W-shaped spot in the wall of
the vacrina just anterior to the vulva; occurring on Taurotraius and

Streps iceros . TAUROTRAGI

Female without such a spot . ;••••• .

With 5-8 slender setae in a cluster on the posterior side of each of
the lateral lobes of the abdomen; occurring on Bos bubalus normally
and apparently also on camels . TOBERCULATUS

With not more than 2-3 setae in this position on any segment . 6

6. Head very short and broad, the length only slightly greater than the

breadth: occurring on domestic cattle throughout the world .

. EURYSTERNUS

Head elongate, from 2-2.5 times as long as wide; occurring on Equidae..
. T . . . ASINI

7. An elongate, slender species in which the paratergal plates of the ab¬

domen form prominent, acutely conical points; occurring on Syncerus

caffer . BUFALI

Stout-bodied species, the paratergal plates forming rounded prominences

_ . . . . . 8

8. Head very short and broad, the width of the hindhead more than half as

great as the length of the head . ....9

Head relatively slender, the width of the hindhead normally definitely
less than half the length of the head . 10

9. Margin of the vulva with a pair of marginally serrate lobes between the

bases of the gonopods; occurring on Phacochoerus . PHACDCHOERI

Margin of the vulva with an undivided and simple median lobe between
the bases of the gonopods; occurring on Poramochoerus . LATHS

10. A stout-bodied form with well-developed, broad, paratergal plates and

with the dorsum of the abdomen normally with sclerotized areas; oc¬
curring on domestic pigs . SUIS

A more slender-bodied form with very weakly developed paratergal plates
and with the dorsum of the abdomen membranous; occurring on the wild
boar of Europe . APRI

Haematopinus acuticeps Ferris

1933- Haematopinus acuticeps Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 6:467; figure 275.

1948. Haematopinus acuticeps Ferris, Webb, Proceedings of the Zoological
Society of London 118:581; Plate I, figure G.

HOSTS AND DISTRIBUTION. Known only from the original record, from Hip-
potigris (= Equus ) burchelli, Mpwapwa, Tanganyika Territory, East Africa.
This host is later cited by Webb as Equus burchelli muansae.

NOTES. Whatever may be the status of the two supposed subspecies bur-
chellis and elegans of Haematopinus asini which actually or presumptively
occur on this same host, there can be no question of the status of acuticeps.
The form of the gonopods of the female is alone sufficient to distinguish it.

Haematopinus apri Goureau

1866. Haematopinus apri Goureau, Bulletin de la Societe des Sciences His-
toriques et Naturelles de l'Yonne, Auxerre 20:205.

1870. Haematopinus urius (Nitzsch), Giebel, Insecta Epizoa, page 45; Plate
2, figure 2.

1880. Haematopinus urius (Nitzsch), Piaget, Les Pediculines, page 654;
Plate 48, figure 4.

1933* Haematopinus aperis Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 6:431; figures252B, 253B, 255A-D, 256F, G, M, N.
1939. Haematopinus suis (Linnaeus), Fahrenholz, Zeitschrift fiir Infektion-
skrankheiten, parasitare Krankheiten und Hygiene der Haustiere 55:
2:138; figures 1-5.

HOSTS AND DISTRIBUTION. From Sus scrofa, the wild swine of western
Europe.

NOTES. The confusion of this species with Haematopinus suis (Linnaeus)
will he discussed in more detail in connection with the latter species. It
is sufficient here merely to remark that the lice of Sus scrofa, the wild

boar of western Europe, seem clearly to be distinct from the lice of the
domesticated swine of the present day, which are thought to have been de
rived from pigs of eastern Asia that have been imported into Europe.

The purely uomeuclatorial problem concerning the proper name lor this
species presents some difficult points. In pre-Liunaeau times the name
Pediculus urius was used very evidently for lice from the pigs of Europe
which were derived from the wild swine of Europe, but Linnaeus did not em¬
ploy this name. It was revived by Nitzsch in 1818 and used by him, clearly
as a substitute for the name suis employed by Linnaeus, since suis is placed
as a synonym of urius.

In 1870 Giebel used the name urius for lice of domestic and wild swine.
His illustration was said by him to have been prepared by Nitzsch in 1805.
In the writer's opinion it clearly represents the lice of wild swine. A
case might, therefore, be made out for the use ot the name urius since the
type of the name urius as employed by Nitzsch would fix the application of
the name if suis be regarded as a separate species.

In 1880 Piaget used the name urius. The writer has examined Piaget's
material and used his specimens as the types for the name aperis, which was
published in 1933 *

However, according to Eahreuholz, the name Haematopinus apri Goureau was
proposed in 1866 for the lice of wild swine of Europe. The present writer
has not seen this work and accepts the name on Fahrenholz ' word. In any
case this name certainly reduced aperis to synonymy.

Hopkins has expressed the opinion that this form should be regarded as a
subspecies of suis, but the writer cannot concur in this.

Haematopinus asini (Linnaeus)

Figures 37, 38

1758. Pediculus asini Linnaeus, Sjystema Naturae, Edition X, page 612.

1829. Pediculus asini (Linnaeus). Stephens, Catalogue of British Insects
2: 329 .

1838. Pediculus macrocephalus Burmeister, Genera Insectorum, Rhynchota,
Species 18.

1916. Haematopinus elegans Fahrenholz, Archiv fur Naturgeschichte, Abteil-
ung A, 81:11:22; figure 8.

1916. Haematopinus minor Fahrenholz, Zoologischer Anzeiger 48:90.

1933. Haematopinus asini (Linnaeus), Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 6:464; figures 273, 274, 275C.
1948. Haematopinus asini asini (Linnaeus), Webb, Proceedings of the Zoolog¬
ical Society of London 118:578.

1948. Haematopinus asini macrocephalus (Burmeister), Webb, Proceedings of
the Zoological Society of London 118:578.

1948. Haematopinus asini minor Fahrenholz, Webb, Proceedings of the Zoolog¬
ical Society of London 118:580; figures g-i ; Plate I, figure U.
1948. Haematopinus asini burchellis Webb, Proceedings of the Zoological
Society of London 118:580; figures d-f; Plate I, figures A, B, E.
1948. Haematopinus asini elegans Fahrenholz, Webb, Proceedings of the Zoo¬
logical Society of London 118:580.

HOSTS AND DISTRIBUTION. The type host is Equus asinus. Recorded also
from domestic horses in various parts of the world and, in the opinion here
adopted, apparently occurring normally on zebras. The type host of macro¬
cephalus apparently was the domestic horse. That of minor was Equus bur-
chelli, and it is recorded also from Equus burchelli grant i. The type host
of burchellis is Equus burchelli grant i. The type of elegans was not speci¬
fied, being recorded merely as from Gobabis, Southwest Africa.

NOTES. ^In these species from the genus Equus we have a most extraordi¬
nary’ situation. Webb (1948) has reviewed the group and adopted the point

Haematopinus asini (Linnaeus)

Figure 37

Haematopinus asini (Linnaeus), details

Figure 38

of view that we have to do with a series of "subspecies," some of which
occur upon the same host. They are described by him as differing only in
the form of the head. It may be noted that Haematopinus acuticeps Ferris
occurs also ou Equus burchellt , but this species is very evidently distinct
in every respect, including the genitalia ol the female.

The point of view adopted by the present writer toward the establishment
of subspecies in this group will be considered elsewhere in this paper. It
is the opinion here adopted that in the case of these supposed subspecies
the evidence for their separation from asini is not convincing except in
the case of the form identified by Webb as minor. The present writer exam¬
ined the types of this species in connection with his earlier work and no¬
ticed no difference which would justify the recognition of this form in any
way, but the illustration given by Webb (his Plate I, figure D) suggests
that it may merit some recognition, although it is not evident that the
form illustrated by him is the same as that to which the name was applied
by Fahrenholz.

Haematopinus breviculus Fahrenholz

1939. Haematopinus breviculus Fahrenholz, Mitteilungen aus dem entomolog-
ischen Verein Bremen, page $2; figures 1-3-
HOSTS AND DISTRIBUTION . Based upon a single female, without indication
of host or origin. Said by Fahrenholz to be very close to eurysternus.

Haematopinus bufali (de Geer)

1778. Pediculus bufali de Geer, Memoires pour servir a l'histoire naturelle
des insectes apteres 7:68; Plate I, figures 11, 12.

1781. Pediculus bufali-capensis Fabricius, Species Insectorum, Volume 2.
1816. Pediculus papillosus von Olfers, De vegetativis et animatis corporibus
in corporibus animatis reperiundis commentarius, page 85.

1844. Pediculus phthlriopsis Gervais, In Walckenaer, Histoire naturelle
des insectes apteres 3:366.

1915* Haematopinus neumanni Fahrenholz, Arch iv fur Nat urge schichte , Abteil-
ung A, 81: 11:8.

1935* Haematopinus bufali (de Geer), Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 6:460; figures 270-272.

HOSTS AND DISTRIBUTION. Originally described as from "le Buffle
d'Afrique," from the Cape of Good Hope. Later recorded from Buffelus
caffer and "buffalo" in Nyasaland and the Congo Free State. Mr. G. H. E.
Hopkins has kindly supplied specimens from Buffelus (or Syncerus) caffer
radcliffei on the shores of Lake Edward.

Haematopinus eurysternus (Nitzsch)

Figures 39. 40

1818. Pediculus eurysternus Nitzsch, Germar's Magazin fur die Entomologie
3:305.

1829. Haematopinus eurysternus (Nitzsch), Stephens, Catalogue of British
Insects 2:329.

1916 . Haematopinus quadripertusus Fahrenholz, Archiv fur Naturgeschichte,
Abteilung A, 81:11:19; figures 15-17.

1916 . Haematopinus parviprocursus Fahrenholz, Archiv fur Naturgeschichte,
Abteilung A, 81:11:21.

1933- Haematopinus eurysternus (Nitzsch) , Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 6:448; figures 263, 264.

1950. Haematopinus quadripertusus Fahrenholz, Muesebeck, Journal of Eco¬
nomic Entomology 43:119, 133*

HOSTS AND DISTRIBUTION. Originally described from domestic cattle in
Europe and known from these hosts from many parts of the world.

NOTES. Haematopinus quadripertusus was described from males only, from
cattle at Banjo, in the Cameroons. Haematopinus parviprocursus was de¬
scribed from females only from unknown host from the then German Southwest
Africa. The types of both these supposed species were examined by Ferris,
who came to the conclusion that they represent the same species and that
this species is to be identified as Haematopinus eurysternus.

This question has been reviewed again in connection with the present
work and the opinion previously expressed is still maintained. It was
pointed out by Ferris (1933) that specimens from cattle in tropical coun¬
tries seem to be much larger and of deeper pigmentation than those from
northern areas, although no morphological differences could be detected.
The only basis then for the separation of the cattle lice into two or more
species is that of size. It must be noted, however, that the cattle of
many tropical areas, such as Asia and Africa, are derived apparently from a
different source from the cattle of Europe and North America, the former
apparently having come largely from the "hump- backed" type and there may be
an association with these differences of host origins.

Specimens of the large form have been collected in Florida in the United
States. These were sent to the writer by Dr. C. F. W. Muesebeck of the
United States Department of Agriculture and the information noted above was
imparted to him. He elected to regard these specimens as representing a
species different from that usually found in the United States and has pub-

Haematopinus eurysternus (Nitzsch)

Figure 39

female genitalia

ventral - male genitalia - dorsal

Haematopinus eurysternus (Nitzsch), details

Figure 40

lished the record, using the common name "Cattle tail louse," for this form.

The question of the specific name to be used for this form, if it be
recognized, arises. It would appear that the name quadripertusus Fahrenholz
is available. However, this form is not here accepted as distinct.

Haematopinus latus Neumann

1909. Haematopinus latus Neumann, Archives de Parasitologie 13:595; Fig¬
ures 6-9.

1911. Haematopinus peristictus Kellogg and Paine, Bulletin of Entomological

Research 2:145. (In part)

1912. Haematopinus incisus Harms, Zoologischer Anzeiger 40:293-

1933* Haematopinus latus Neumann, Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 7:477: figures 260C, 262.

1939. Haematopinus latus Neumann, Fahrenholz, Zeitschrift fiir Infektions-
krankheiten, parasitare Krankheiten und Hygiene der Haustiere 55:
151 ; Plate 4, figures 9-11.

1939. Haematopinus latus latissimus Fahrenholz, Zeitschrift fur Infektions-
krankheiten, parasitare Krankheiten und Hygiene der Haustiere 55:
152; Plate 4, figure 12.

HOSTS AND DISTRIBUTION. The types of latus were recorded as from
Potomochoerus africanus at Kaporo, Nyasaland. The types of the supposed
variety latissimus are the specimens recorded by Ferris from Potomochoerus
choeropotamus , Luangwa Valley, northeastern Rhodesia. Recorded from the
same host from Zululand and South Africa and also from Potomochoerus affinis
from the former German East Africa.

NOTES. This species has at times been contused with Haematopinus
phacochoer l Enderlein, but is clearly distinct.

The writer sees no excuse for the naming of the supposed variety latis-
simus. Among the specimens at hand are some of evidently very early adult
females and others of fully adult individuals. There is enough difference
among these specimens to suggest the presence of two forms and this seems
to be the only basis for Fahrenholz' supposed variety.

Haematop inus longus Neumann

1912. Haematopinus longus Neumann, Bulletin de la Society Zoologique de
France 37*141; figure 1.

1933 . Haematopinus longus Neumann, Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 6:469; figure 276.

HOSTS .AND DISTRIBUTION. Known only from the original record, from
Cervus unicolor, at Kota, Nepal, India.

Haematopinus phacochoeri Enderlein

1908. Haematopinus phacochoeri Enderlein, Wissenschaftliche Ergebnisse der
Schwedische Expedition nach dem Ki limandj aro, dem Meru, und dem
umgebenden Massaische Steppen 11:7; figure.

1911. Haematopinus peristictus Kellogg and Paine, Bulletin of Entomological
Research 2:145; Plate 4, figures 3-6 (in part).

1933. Haematopinus phacochoeri Enderlein, Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 6:444; figures 260, 261.

1939. Haematopinus phacochoeri Enderlein, Fahrenholz, Zeitschrift fur In-
fektionsk rankhei ten , parasitare Krankheiten und Hygiene der Haus-
tiere 55:149; figures 9a, 9b.

1939. Haematopinus phacochoeri peristictus Kellogg and Paine, Fahrenholz,
Zeitschrift fur Infektionskrankheiten, parasit&re Krankheiten und
Hygiene der Haustiere 55:150.

HOSTS AND DISTRIBUTION. The types of phacochoeri were from Phacochoerus
aeliani massaicus in the region of Mt. Kilimanjaro and those of peristictus
from Phacochoerus aethiopicus , at Akamanga, North Nyasa, Africa. The
species has been recorded from Phacochoerus sp. in the former German East
Africa, and from phacochoerus sundevali in Zululand. In addition, there
are records given as merely from "wart hog, " in East Africa, northeastern
Rhodesia and southern Rhodesia. Certain records from "buffalo," in Kenya

are open to doubt. . «... • .

NOTES. There has in the past been some confusion ot this species with

Haematopinus latus Neumann, but it appears that these two species are char¬
acteristic of different host genera. Fahrenholz has attempted to resurrect
the name peristictus as a variety, but the types of this name are at hand
and offer no justification tor this procedure.

Haematopinus suis (Linnaeus)

Figures 41, 42

1758. Pediculus suis Linnaeus, Systema Naturae, Edition X, page 611.

1810. Haematopinus suis (Linnaeus), Leach, Encyclopaedia Brittannica, Sup¬
plement 1, page 24. « nn p-

1818. Pediculus urius Nitzsch, Germar s Magazin der Entomologie
1903. Haematopinus irritans Law, Textbook of Veterinary Medicine ^:13-
1911. Haematopinus suis aduenticius Neumann, Archives de Parasitologie 14.

406 ; figure 8. .

1916. Haematopinus suis chinensis Fahrenholz, Fahrenholz, Zoologischer An-

zeiger 48:90.

Haematopinus suis (Linnaeus)

Figure 41

thcrax

antenna

genitalia

'Haematopinus suis (Linnaeus), details

Figure 42

Haematopinus suis germanus (misprint tor germanicus) Fahrenholz, Zo-
ologischer Anzeiger 48:90.

Haematopinus suis sardiniensis Fahrenholz, Jahrbuch der Hamburgisch-
en Wissenschaftlichen Anstalten 34:Beiheft 2:10.

Haematopinus suis (Linnaeus), Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 6:425; figures 252A, 253A, 254.
Haematopinus adventicius Fahrenholz, Ewing, Proceedings of the Hel¬
minthological Society of Washington, page 76.

Haematopinus chinensis Fahrenholz, Fahrenholz, Zeitschrift fur In-
fektionskrankheiten, parasitare Krankheiten und Hygiene der Haus-
tiere 55:Heft 2:138; Plate 3> figures 1-3-

HOSTS AND DISTRIBUTION. The name Pediculus suis was applied specifical¬
ly By Linnaeus to the lice of domestic swine, presumably those of Sweden.
Since that time it has been used for the lice of domestic swine, as recorded
from almost all parts of the world. The only records from undomesticated
swine seem to be those given by Ferris (1933) from Sus cristatus, from
Tanjong Badak, Tenasserim and from Dinapore, India.

NOTES. Concerning this species we have a problem that is compounded of
both nomenclatorial and zoological questions, for none of which is there at
present a satisfactory answer. The story is long and complicated and
nothing more than a mere abstract of it can here be presented.

The reasons for the complications rest largely upon the fact that the
domesticated swine population of Europe has changed within relatively recent
times. The domesticated swine of western Europe, throughout most of his¬
toric times, were presumably derived from the wild swine of that area.
Early in the eighteenth century domesticated swine from eastern Asia were
introduced into Europe and swine of this type have now become the common
domesticated animals of almost all the world.

Because of these facts, references to the lice of swine in European lit¬
erature, certainly prior to such a date as 1758, undoubtedly concerned the
lice of the wild boar of Europe, Sus scrofa. Even at that date, 1758, the
population of domestic swine in Europe was changing.

Now, if the native swine of Europe originally had upon them a louse of a
species different from that of the domesticated swine of eastern Asia — as
seems to be true — to which of these species should the name suis apply?
There are two indirect approaches to the problem. One of these rests upon
pre-Linnaean history and the other upon post-Linnaean history. On the
basis of pre-Linnaean history it is clear that the name suis should apply
to lice from the wild swine of Europe. But on the basis of post-Linnaean
history the name has been used for almost 200' years to apply to lice de¬
rived from the swine of eastern Asia, which even at the time of Linnaeus
were beginning to replace the other type.

No one knows what Linnaeus actually had in hand when he named the species
Pedicuius suis. The probabilities certainly are that he used the name for
lice of swine of the older European race, although it must be noted that
the Asiatic swine had been introduced into Sweden a few years before the
appearance of the tenth edition of the Systema Naturae and it is possible —
since they undoubtedly attracted a considerable amount of attention — lice
may have been taken from them and come to Linnaeus' notice.

From the point of view of the present writer the reasonable procedure is
to accept the name suis for the lice of domestic swine as we now find them
and to consider that as used by Linnaeus the name covered a compound
species. We may legitimately argue that the general usage of the last 150
years or more constitutes a de facto restriction of this name to the form
for which it has commonly been used and that to change this at the present
date is unnecessary and confusing.

Furthermore, it should be noted that as early as 1818 Nitzsch revived
the old, pre-Linnaean name urlus as a substitute for suis, and that later

1916.

1917.

1933-

1934.

1939.

evidence has indicated that lie actually had at hand the lice ol the wild
swine. This name might be considered thereby to have become established.
In any case, in 1866 these lice were definitely given the name Haemtopinus
aprl Goureau.

Fahrenholz and Ewing have opposed the point of view here held and have
maintained that the name suts should be transferred to the lice of the wild
swine of Europe, thus leaving it to be detennined what name should be used
for those from the present-day domesticated swine. Fahrenholz has main¬
tained that the proper name to be used is Haema top inus cbinensis Fahrenholz,
while Ewing has supported Haematopinus advent ictus Neumann, both overlooked
the earlier name Haematop inus trritans Law.

The problem can in the end be settled only by a ruling of the Interna¬
tional Commission on Zoological Nomenclature, and lor the present the writer
refuses to disturb the long-established nomenclature.

So much for the nomenclatorial problem. The zoological problem is also
involved and complicated.

To begiu with, it appears that the lice of the wild swine of Europe dif¬
fer to such a degree from those ot domesticated swine tliat they utiy justi¬
fiably be called a distinct species. After we have eliminated the lice of
the wild swine of Europe from consideration, there remains the question ol
the forms to be found among the lice of domesticated swine. Among these
there is present a considerable variety ot size, lorm, pigmentation, and ol
some morphological characters. At least three of the forms have been first
named as varieties and then elevated to the rank of species by Fahrenholz.

Ferris (1933), on the basis of an examination of specimens from numerous
parts of the world, gave an extended account of the variations to be found
among them. Here and there appeared individual collections which, if taken
by themselves alone, might justify some nomenclatorial recognition. But
after these specimens are removed we are left with a residue ol material
which presents almost every possible combination of the characters seen in
the others, indicating a completely interlinked genetic chain which con¬
forms to the concept of species here adopted and previously discussed. In
the end the problem can be settled — if at all — only by genetic studies.
For the present the single name Haemtopinus suis (Linnaeus) will here be
used to cover this complex of forms.

Haematop inus taurotragi Cummings

1914. Haematoptnus taurotragi Cummings, Bulletin of Entomological Research

5:155; figures. _

1933. Haematopinus taurotragi Cummings, Ferris, Contributions I oward a
Monograph of the Sucking Lice, Part 6:453; figures 265, -66.

HOSTS AND DISTRIBUTION, type from Taurotragus oryx ( =Boselaphus oreas) ,
from a menagerie in England. Later recorded by Bedford from the same host
at various placed in South Africa and also from Strepsiceros kudu in South
Africa.

Haematopinus tuberculatus (Burmeister)

Figures 43, 44

1800. Pedtculus bufall-europaei Latreille, Histoire naturelle et parti culi-
ere des crustaces et des insectes 8:96 (nomen dubium,) .

1839. Pedtculus tuberculatus Burmeister, Genera Insectorum, Khynchota,

1862 Haematopinus tuberculatus (Burmeister', Lucas, Annales de la Socieie
Entomologique de France (series 2) 10:529; Plate 11, number 2.

1869 . Haematopinus punctatus Rudow, Zeitschrift fur die gesamten Naturwis-
senschaften 34:16*.

Haematopinus tuberculatus (Burmeister)

Figure 43

neutral male genitalia dorsal female genitalia

Haematopinus tuberculatus (Burmeisterb details

Figure 44

1919. Haematopinus bufali-europaei (Latreille), Fahrenholz, Zeitschnft
fur angewandte Entomologie 6:154. .

1933. haematopinus tuberculatus (Burmeister), Ferris, Contributions Toward
a Monograph of the Sucking Lice, Part 6:455; figures 26 / 269 -
HOSTS AND DISTRIBUTION . Burmeister' s name was based upon specimens irom
the "common buffalo or buffalo of India," presumably Bos bubalus, i and t.here
are manv records of the species from this host in India, China, the Philip
pine Islands and Guam. In addition, there are records of the species ; from
the yak, Bos trunniens, and from camel in Australia and Africa. Latreill s
snecimens were recorded as from buffalo Irom Ital} . . , ,

NOTES. There is a strong probability that the species named 3 La rei
(1800) is the same as that named by Burmeister in 1839, but the writer has
In the oast declined, and still declines, to accept the synonymy involved.
A verv considerable literature has grown up under the nam e tuberculatus,
while' under the clumsy name bufali-europaei there have been but occasional
nassinii references since it was first proposed.

P Specimens from camel have been examined by the writer. They seem
differ from the typical form only in smaLler size.

Genus PECAROECUS Babcock and Ewing

1939- Pecaroecus Babcock and Ewing, Proceedings of the Entomological Soci¬
ety of Washington 40:197.

GENERIC TYPE. Pecaroecus javalii Babcock and Ewing.

CHARACTERS. Haematopinidae with distinct eyes, these being represented
by a facet and a pigment spot just posterior to each antenna. Legs without
a sclerotized plate in the "palm" of the tibia. Thorax without a sclero-
tized sternal plate. Abdomen with small, sclerotized, tubercle-like para-
tergites present.

NOTES. The single known species of this genus is a strange form which
seems rather definitely to be related to Haematopinus.

Pecaroecus javalii Babcock and Ewing
Figures 45, 46

1938* Pecaroecus javalii Babcock and Ewing, Proceedings of the Entomologi¬
cal Society of Washington 40:199; figures 103 -

HOSTS AND DISTRIBUTION. Known from a single record, from Pecari aniula-
tus, between Juno and the Pecos River, State of Texas, United States.

NOTES. Specimens from the type lot have been made available through the
kindness of Mr. Babcock and the accompanying illustrations are based upon
these.

Family HOPLOPLEURIDAE Ferris, new family

DESCRIPTION OF THE FAMILY. Anoplura in most of which there is no exter¬
nal indication of eyes, lenses being present in only one genus. Abdomen,
in all but one species, with paratergal plates on at least one segment,
these plates always at least in part with an apical portion which is free
from the body and not forming a mere cap over the apex of lateral lobes of
the abdomen. Thoracic sternal plate usually developed, commonly with its
posterior apex free, but at times weakly sclerotized and at times lacking.
Ventral prothoracic apophyseal pits lacking. Abdominal tergal and sternal
plates very commonly present and well developed. Antennae normally five-
segmented, in one species described as three-segmented, at times with the
terminal three segments tending to be more or less fused together, frequent¬
ly sexually dimorphic, the male having either the apical preaxial angle of
segment three more or less produced and bearing one or more short, stout,
retrorse setae dorsally, or having such a seta on the dorsal side of each
of the last three segments. Female with the gonopods of the ninth segment
usually short and not strongly developed, never elongated and leaf-like.

NOTES. As here understood this is the largest family of the Order, most
of its members occurring on rodents, but a few on Insectivora and Primates,
and one on Ungulata.

The principal objection that may be urged against this family is the in¬
clusion in it of the genus Pedicinus, which has long been associated with
the Pediculidae. This matter will be discussed at length under the sub¬
family Pedicininae and it may here simply be noted that in the writer's
opinion the genus is more closely related to the Hoplopleuridae than it is
to Pediculus .

The general pattern of the Hoplopleuridae is reasonably consistent, be¬
ing marked especially by the development of the paratergal plates of the
abdomen, which always have at least an apical point free from the body wall.

The famiiy name is chosen from a genus which is large and in which the
characters of the family are especially well developed.

The family is here divided into five subfamilies, which may be separated
by the following key.

Pecaroecus javalii Babcock and Ewing

Figure 45

genitalia of female

Pecaroecus javalii Babcock and Ewing,

Figure 46

100

Key to the Subfamilies of Hoplopleuridae

Eyes present externally as a pair of distinct lenses; occurring on Old

World monkeys of the superfamily Cercopithecoidea . PEDlCININAE

Without any external indication of eyes . . • “

With a small, claw-like process alongside the true claw on the anterior

legs . ’ . HYBOPHTHIRINAE

Without such a claw-like process on the anterior legs . •••3

Anterior and middle lej/sot the same size and ionn, both small and slen-
der, with slender claw; ventral side of the abdomen in all species
except three, with a pair of small, sclerotized, detached plates on

the venter of abdominal segment two; restrictei to Sciuridae .

. ENUE KLEIN ELLIN AE

Anterior legs the smallest of the three pairs, the second pair at least
somewhat larger than the first and with stouter claw; venter ot ab¬
dominal segment two without such a pair of detached plates, except as
the pa rater gal plates of this segment may be longitudinally divided

into a dorsal and a ventral piece . ••••• •• ** •*.••• *

Second abdominal segment with its sternal plate extended laterally on
each side to articulate with the corresponding paratergal plate......

. HOFLOPLElIKLN AL

Second abdominal segment with its sternal plate never thus lat¬
erally . POLYP LACINAE

Subfamily EN DERLELNELLIN AE Ewing

1929. Ewincr, A Manual of External Parasites, page 132.

DESCRIPTION OF THE SUBFAMILY. Hoplopleuridae with the two anterior pairs
of le.rS slender and with slender claw, the two pairs being of the same size
and shape, the third pair stout and with stout claw. All but three ol the
included species provided with a pair of small, highly sclerotized plates
on the ventral side of the second abdominal segment, each ot these plates
bearing a strongly sclerotized, flat projection. Usua ly very sma
species. All the known species occurring on the rodent family Sciuridae.

NOTES. The members of this subfamily are among the smallest ol e
Anoplura, some of them attaining a length of scarcely, more than .3? mm-
The peculiar plates on the venter of the second abdominal segment are at
present of undetermined homology. In some species each plate articulates
laterally with the corresponding paratergite. It nay be either a detache
oortion of the paratergite arising from some such phenomenon as the ventral
portion of the longitudinally divided paratergites of this
■innpars esDeciallv clearly in the genus Fahrenholzia — or it may be an ex
treme development of the divided sternal plate of the second segment which
appears in s^ch genera as Schizophthirus and Haematopinoides and is at
times indicated in Roplopleura itself. In any case the development seen in

^ ^ iCo" -enera —Hicrophthirus and tterneckia— totaling three species, which
are here assigned to the Enderleinellinae, lack these plates but in o
respects seenTproperly to be associated with Enderletnellus , both on the
basis of general structure and host relationships.

Key to the Genera of Enderleinellinae

The pair of pUtes on the venter of abdomnal se^t two^res^i...^^

^Seteroaal SplaTes°of abdominal segments' three' to 'six iaih connected
ventrallyP»ith the corresponding sternite by a narrow, sclerotized

101

bridge; paratergal plates with their posterior margin not free from
the body wall; occurring on American flying squirrels. . .MICROPHTHIRUS
Paratergal plates of the abdominal segments not thus connected with the
stemites, their apices free from the body wall; occurring on African
squirrels . WERNECKIA

Genus ENDERLEINELLUS Fahrenholz

1912. Enderleinellus Fahrenholz, Zoologischer Anzeiger 39:56.

1919. Enderleinellus, Ferris, Contributions Toward a Monograph of the Suck¬
ing Lice, Part 1:7.

1929. Cyclophthirus Ewing, A Manual of External Parasites, page 195 *

1929. Euenderleinellus Ewing, A Manual of External Parasites, page 197.
1929. Hoplophthirus Ewing, A Manual of External Parasites, page 194.

1929. Rhinophthirus Ewing, A Manual of External Parasites, page 196.

GENERIC TYPE. Pediculus sphaerocephalus Nitzsch = Enderleinellus
nitzschi Fahrenholz.

GENERIC SYNONYMS. Cyclophthirus Ewing, type Haematopinus suturalis
Osborn. Euenderleinellus Ewing, type Enderleinellus larisci Ferris.
Hoplophthirus Ewing, type Enderleinellus euxeri Ferris. Rhinophthirus
Ewing, type Enderleinellus heliosciuri Ferris.

CHARACTERS. Enderleinellinae in which a pair of small, sclerotized
plates is present on the venter of abdominal segment three, these plates
bearing a flattened point which is apically free from the body. All the
known members of the genus occur on hosts of the rodent family Sciuridae.

NOTES. In 1929 Ewing proposed to divide this genus, forming from it a
total of five genera. Actually, if one wishes to overlook the basic simi¬
larities which exist throughout the group' and to seize upon the peculiari¬
ties of individual species, this number of genera could be doubled with no
special effort. It may very well be that when the numerous species that
must still remain to be discovered have been incorporated into the system
some division will be justified and at least a part of the names proposed
by Ewing will deserve to be accepted. For the present nothing much seems
to be gained by such division and all the species are here retained in
Enderleinel lus.

Certain very troublesome problems arise in connection with some of the
groups of species within the genus. These can be illustrated especially by
referring to the group which occurs on New World members of the genus
Sciurus. A very considerable amount of material representing this group
has been assembled, but there remain many host species from which nothing
has yet been collected. Within this material it is possible to recognize a
wide variety of form in the genitalia of the males, although all the dif¬
ferences represent merely modifications of a single pattern. Some speci¬
mens are so different from others that it would appear nonsensical to refer
them to the same species, but there are numerous instances of intermediate
forms differing only in some small detail. We have on the one hand the
constantly pressing temptation to name a new species or "variety" for each
of these slightly different forms, or on the other hand to throw all these
variants into a few species. Either method produces results which are un¬
desirable and which may equally effectually obscure any approximation to
truth. At, the time when the section on this genus was prepared by Ferris
for his "Contributions Toward a Monograph of the Sucking Lice," he was but
an inexperienced student of the systematics of insects. Disgusted by the
results of unrestrained "species mongering" he revolted against the prac¬
tices of certain other workers and went to the other extreme. It now ap¬
pears that this conservatism was in its turn unsupportable and certainly
some of the so-called species recognized by Ferris are actually species
groups.

102

Dr. Fabio Werneck, of the Instituto Oswaldo Cruz, of kio de Janeiro,
Brasil, in the course of a few months spent at Stanford University, inde¬
pendently worked over the extensive collection oi Enderletnel lus there a
vailahle . He detected some of the errors, not only of judgment but ol ob¬
servation, on the part of Ferris and named certain new species Irom the
collection. However, a considerable amount of material was lelt b) him as
still doubtful and remains for further study to elucidate it.

With the addition of the species recognized by Werneck the number ol
species now assigned to the genus is 29, which we may suspect represents
scarcely more than halt the species that exist. El lender, in his review ol
the rodents, recognizes *14 genera in the Sciuridae, these containing ap
proximately 1300 named forms. The vast majority of these forms are listed
as subspecies and as there appears to be an averaged about live subspecies
for each species it may be assumed that about 300 actual species exi^t in
the family. Of these only about 50 have as yet yielded species of Knder-
leinellus, although the evidence indicates that every Sciurid possesses a
member of this genus. It therefore seems probable that the total number ol
species of the genus in existence will probably be between oO and 60. When
the majority of these are knowu we shall be in a much better position than
at present to develop a classification of the group.

Key to Species of ENDERLELNELLUS

1. Paired ventral plates of abdominal segment 2 each produced laterally to

meet the corresponding paratergal plate . ••••• . * .

Paired ventral plates of abdominal segment 2 circular, not thus produced
laterally and completely detached from the paratergites . ••

2 (1). Abdominal segments 4-8 each with a pair of very long setae on the

paratergal plates or at the lateral margins of the body, the lat¬
eral margins thus fringed with pairs of long setae; occurring on

Marmota in North America . •••••; . MARMOT AE

Such long setae present only on the paratergal plates...... . J

3 (2). Female with 2-4 very long setae on the dorsum oi abdominal seg™ent

4, these setae reaching to the apex ot the body; known lrom
Ci tel lus beecheyt, Cttellus variegatus, and Cite ll us teret icaudus

in southwestern and extreme western United States . UbttJKNi

Female without such long dorsal setae; known from many species ; ol
Citellus from Siberia throughout North America . aJiLRALiS

4 (1). Paratergal plates present only on segments 2n4.. . Q’

Paratergal plates present on segments ^2-5 or z-b . .

5 (4). Spiracles present only on segments S~i. . . . . E

Spiracles present on at least segments 3-5 and at times on 3

6 (5). Both sexes with abdominal setae, both dorsally and ventrally , 1' g ,

^ flattened, elongate-cuneiform; known from

Both* sexes with the abdominal setae, both dorsally and ventrally’,
all slender; known from Paraxerus in Africa. .......... - Z/UhAlbS

7 (6). Dorsum of the abdomen very sparsely haired, the lemale with no

more than 4-5 setae on each half anterior to segment, these all

very ’ong; the male with not more than 3-4 such setae and with a

median series of pairs of very small

Dorsum^fn^both' sexes j with seUe more numerous and none much longer

than the length of its segment . . .

q (7) Dorsum of the abdomen with numerous setae, there being as many as

8 UorsumQ ^ ^ almost continuous row across segments 4-6 m both

sexes: known from Lartscus in Borneo . •••••• . . . LAKlbtl

Dorsum of the abdomen with not more than 8-10 setae in any row in

103

the female and less on the male; known from Menetes in Siam .

. MBNETJNSIS

9 (4). Paratergal plates present on abdominal segments 2-6 . 10

Paratergal plates present on abdominal segments 2-5 . 12

10 (9) . Thoracic sternal plate with a slender, median, anterior prolonga¬

tion which extends forward between the anterior coxae; occurring

on the European Sciurus vulgaris . NITZSCHI

Thoracic sternal plate without such anterior prolongation . 11

11 (10). Paratergal plates of abdominal segment 3 bearing a seta which is

as long as the abdomen itself; known from Xerus erythropus in

Africa . EUXERI

No seta on any paratergal plate longer than the plate itself;
known from Dremomys pernyi in China . DREMOMYDIS

12 (9) . Thoracic sternal plate with a narrow, median, anterior prolonga¬

tion extending forward between the anterior coxae . 15

Thoracic sternal plate without such median anterior prolongation..

. 13

13 (12). All spiracles extremely large, their outside diameter equaling

almost half the width of the plates which bear them; known from

members of the genus Calloscturus in the Malayan Region .

. MALAYS! ANUS

Spiracles small, equaling only a small fraction of the width of
the plates which bear them . 14

14 (13) . Paratergal plates of segment 4 with 1 of their 2 setae equaling

2-3 times the length of the plate that bears it ; attributed
(possibly erroneously) to the North American Tamtas striatus...

. TAMIASIS

Paratergal plates of segment 4 with the setae much shorter than
the length of the plates; known from Nannosciurus in Java .

. . NANNOSCIURI

15 (12). Thoracic sternal plate divided longi tudinally into 2 oval, strong¬

ly sclerotized pieces which are separated from each other by a
more or less membranous median area; occurring on New World
squirrels of the genus Sciurus and closely related genera (see

special key); the group of . LONGICEPS

Thoracic sternal plate without such division; known from Sciuro-

t ami as in China . SCIUROTAMIASTS

The Longi ceps Group

The members of this group constitute an extraordinary complex of evi¬
dently closely related forms which occur on New World members of the genera
Sciurus and Microsciurus , the latter being a little genus that is confined
to Central America and northern South America. While a considerable amount
of material is available, it represents only a small portion of the forms
that probably exist. The development of an understanding of this complex
has only begun and consequently uncertainty concerning the species which
should be recognized must remain for a long time to come. All that has
been accomplished thus far is to segregate a few of the forms that are
clearly recognizable, while there are left on hand numerous specimens that
cannot he assigned to one species or another.

In his treatment of the genus Enderlelnellus in Part 1 of his "Contribu¬
tions Toward a Monograph of the Sucking Lice, " Ferris called attention to
the existence of numerous variants of the males of the four species of this
group which he recognized. Later, Werneck worked over this same material
and from it and other specimens available to him later named six more
species, bringing to ten the total number of species recognized in the
group. Neither Ferris nor Werneck was able to point out characters by

104

which the females could be differentiated and the species were based almost
entirely upon characteristics of the genitalia of the males.

A renewed study of the group in connection with the present work has re¬
vealed certain minute characters which serve to make at least some of the
species identifiable in the female also, but even yet there remain some
which can be associated with their males only by being found in company
with them.

The males present what is probably the most unusual complex of struc¬
tures that is to be found in this sex in any of the sucking lice. Material
upon which to base the much-needed dissections to explain these structures
has not been available in connection with this work aud such a study must
be left to the future. All that can be done at present is simply arbitral^
ily to identify the same structure throughout the series without attempting
to discuss its morphological nature.

In the material at hand and among the species which have been named
there are males with genitalic structures so different that there is noth¬
ing to be done other than to regard them as distinct species, but between
these extremes there exist many intermediate forms with which nothing can
be done at present. These must either be left unnamed or referred tenta¬
tively to the nearest recognized species.

Because of these facts separate keys are provided for males and females,
that for the males being the most nearly complete.

Key to Males

Arms of the basal plate bent laterally at the posterior end, but not
expanded at the apex nor with apical lobes or with any subapical spur

or expansion . 2

Arms of the basal plate expanded apically, or divided apically into two

lobes or with a subapical, mesal spur or expansion . ....3

Piece X elongate, widened posteriorly; known only from Sclurus iriseus

in California . KELLOGG!

Piece X very short, transverse; typical of Scturus niter . LONGICEPS

Arms of the basal plate with a distinct little subapical spur which is
set well in from the acute and simple apex; type from Sciurus boothiae

. HONDURENSIS

Arms of the basal plate without such subapical spur . ;-;-4

Arms of the basal plate with the two very small apical lobes containing

the articulation of the corresponding arm of the pseudopenis' and with
a marked subapical, mesal expansion set well in from the apex: type
from Orosciurus in South America . URDSCIURI

Arms of the basal plate with a quite shallow apical emargination . 6

Arms of the basal plate with a deeply V- or U-shaped emargination - ,

Arms of the basal plate with the mesal apical lobe larger than the
other and rounded at the apex; type from Sciurus aestuans in Brasil..

. BRASI LIEN SIS

Arms’ of ’ the* basal" plate with the outer lobe larger than the mesal lobe
and with both lobes apically acute; type from Sciurus nesaeus in Ven¬
ezuela . INSULARIS

Median complex of the aedeagus with a pair of minute, recurved hooks at
the mid-line, just posterior to the gonopore ; type from Sciurus truei

in Mexico . ' . ; . MEXICANUS

Median complex of the aedeagus without such hooks............... -

Arms of the basal plate with the mesal apical lobe shorter than the
other; pseudopenis with its apical stem at least half as long as its

arms; type from Sciurus arizonensis . •.•••••••• . . . AKlZONeHSlo

Arms of the basal plate with the two apical lobes equal m length,

stem of the pseudopenis less than half as long as the arms, at times
forming merely a short projection . 9

9. Arms of the basal plate with the apical lobes short and narrow, not ex¬

panded basally; from Microsciurus in Colombia . MICROSCIURI

Arms of the basal plate with the apical lobes very large and broadened
basally . 10

10. Arms of the basal plate with the mesal lobe definitely broader at the

base than is the outer lobe; type from Sciurus social is in Guatemala.

. .EXTREMUS

Arms of the basal plate with the apical lobes of the same width at the
base; type from Sciurus £riseo$ena in Venezuela . VEHEZUELAE

The distinctive characters in the females are to be found in connection
with the genitalia. The anterior margin of the vulva is beset with small
fimbriae, medially forming a relatively stout and somewhat sclerotized
point. Just anterior to this median point is to be seen a sclerotized
structure which is presumably the spermatheca. Its exact relation to the
surrounding parts remains to be determined from fresh material and here
nothing more can be done than to describe and illustrate its appearance.
In one species of the group ( kelloggi ) it appears to be lacking. In other
species its form is quite distinctive and it ranges in size from a large,
oval body as seen in venezuelae (Figure 41) to a very minute structure as
in microsciuri (Figure 41). This structure maintains its characteristic
shape throughout all the specimens available from the type host and is here
illustrated for nine of the eleven species. A careful special study of the
entire group is needed.

Key to Females
Figure 47

Two species, brasiliensis and urosciuri , are omitted from this key be¬
cause of lack of material.

1. Spermatheca apparently lacking; median tooth of the vulvar fimbriation

strongly sclerotized, this sclerotization extending forward until it

merges with the sclerotization of the genital pJat-' . KELLOGGI

Spermatheca definitely present . 2

2. Spermatheca forming a relatively very large, oval body . VENEZUELAE

Otherwise . 3

3. Spermatheca strongly bent, its ends expanded, the expansion of the an¬

terior end being larger than that of the posterior end . . . . ARIZONENSIS
Not so . 4

4. Spermatheca forming a relatively straight, simple, tapering tube . 5

Not so . 6

5. Spermatheca with a slender, tubular, posterior prolongation .. .LONGI CEPS
Spermatheca larger and stouter, without such posterior prolongation....

. INSULARIS

6. Spermatheca forming an elongate body which is anteriorly swollen, then

reducing by an abrupt constriction to expand again gradually toward
the posterior end, then becoming terminally truncate with a small,

strongly sclerotized terminal appendix . EXTREWUS

Not so . 7

7. Spermatheca forming an elongate body which is more or less constricted

about the middle . MEXICANUS

Not so . 8

8. Spermatheca extremely minute, short, anteriorly swollen, and then con¬

stricting, its posterior end prolonged into delicate tube MICROSCIURI
Spermatheca similar in form but much larger and apparently without the
slender prolongation . HONDURENSIS

106

arizonensis

hondurensis extremus microsciuri insularis

Details of genitalia of females of the Enderlei nellus longiceps group

kelloggi

Figure 47

107

Eaderleinellus arizonensis Werneck

194S. Eaderleinellus arizonensis Werneck, Memorias do Instituto Oswaldo
Cruz 45:288; figures 13— 15 *

HOSTS AND DISTRIBUTION. Type from Sciurus arizonensis from the Huachuca
Mountains in Arizona. Also recorded by Werneck from Sciurus apache from
Colonia Garcia, Chihuahua, Mexico; Sciurus alleni from the Sierra Guadelupe.
and Sciurus nayaritensis from the Sierra Madre, Zacatecas, Mexico.

NOTES. ■'This species was included by Ferris in Eaderleinellus extremus
Ferris.

Enderleinellus brasiliensis Werneck

1937. Enderleinellus brasiliensis Werneck, Memorias do Instituto Oswaldo
Cruz 32:399; figure 11.

HOSTS AND DISTRIBUTION. From Sciurus aestuans at Abaete, state of Para,
Brasil .

Enderleinellus dremomydis Ferris

1919. Enderleinellus dremomydis Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 1:29; figure 16.

HOSTS AND DISTRIBUTION. From Dremomys pernyi, West Szechuan, China.

Enderleinellus euxeri Ferris

1919. Enderleinellus euxeri Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 1:37; figures 22, 23-
1929. Hoplophthirus euxeri (Ferris), Ewing, AManual of External Parasites,
page 194.

HOSTS AND DISTRIBUTION. Described as from Euxerus microdon, which is a
synonym of Xerus erythropus, from Wambugu and Oni, British East Africa.

Enderleinellus extremus Ferris

1919. Enderleinellus extremus Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 1:24; figure 12.

HOSTS AND DISTRIBUTION. Type from Sciurus socialis at Nenton, Guatemala.
Also recorded from Sciurus aureogaster from the states of Vera Cruz and
Oaxaca; from Sciurus deppei from the state of Tabasco; from Sciurus griseo-
flavus, state of Chiapas; from Sciurus negligens , state of Tamaulipas; from
Sciurus poliopus, state of Oaxaca; ail these being from Mexico. All of
these specimens are left in extremus by Werneck.

Enderleinellus heliosciuri Ferris

1919. Enderleinellus heliosciuri Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 1:40; figures 24, 25.

HOSTS AND DISTRIBUTION . Type from Heliosciurus gambianus (as undulatus)
from British East Africa and from the same host (as rufobrachlatus and
multicolor) from Uganda; from Heliosciurus ruwenzori i, Mt. Ruwenzori,
British East Africa; from Protoxerus stanieri , British East Africa.

Enderleinellus hondurensis Werneck

1948. Enderleinellus hondurensis Werneck, Memorias do Instituto Oswaldo
Cruz 45:286; figures 7-9.

108

HOSTS AND DISTRIBUTION . type from Sciurus boot hi ae 1‘rom San Pedro Sula,
Honduras. Also from Sciurus melanin at Boquerou, Colombia, and from
Sciurus uar iegatoides (as gohimarii) from the state of Chiapas, Mexico.

NOTES. This species, with the above records, was included by Ferris in
Under l e Ine l l us kelloggl.

Eriderleineilus insularis Weraeck

1948. Enderleinellus Insularis Werneck, Memorias do Instituto Oswaldo Cruz

4?: 293; fibres 25-27.

HOSTS AND DISTRIBUTION, type from Sciurus nesaeus from Margarita Is¬
land, Venezuela.

NOTES. This species was included by Ferris under Enderleinel lus extremus.

Enderleinellus kelloggi Ferris
1916. Enderleinellus kelloggi Ferris, Psyche 23:10^.

1919. Enderleinellus kelloggi Ferris, Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 1:22; figure 11.

HOSTS AND DISTRIBUTION. From Sciurus griseus, Stanford University,
California.

NOTES. Specimens erroneously recorded as kelloggi are here placed under
hondurensis.

Enderleinellus larisci Ferris

1919. Enderleinellus larisci Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 1:17; figures 7, 8.

1929. Euenderleinellus larisci (Ferris), Ewing, A Manual of External Para¬
sites, page 197. . -

HOSTS AND DISTRIBUTION. From Lariscus insignis (as diversus) , from
Lanchut, southwest Borneo.

NOTES. This species was designated as type of the genus Euenderleinellus
Ewing.

Enderleinellus longiceps Kellogg and Ferris

1915. Enderleinellus longiceps Kellogg and Ferris, Anoplura and Mallophaga
of North American Mammals, Stanford University Publications, Uni¬
versity Series (no volume number), page 44; Plate 2, figure 5>
Plate 4, figure 12; Plate 6, figure 2.

1919. Enderleinellus longiceps Kellogg and Ferris, Ferris, Contributions
Toward a Monograph of the Sucking Lice, Part 1:19; figures 9, 10.
1948. Enderleinellus ^longiceps Kellogg and Ferris, Werneck, Memorias do
Instituto Oswaldo Cruz 4^:284; figures 1—3*

HOSTS .AND DISTRIBUTION. Type from Sciurus niger or Sciurus carolinensis ,
at Lincoln, Nebraska. Recorded from Sciurus niger at Valentine, Nebraska,
and Waterloo, Indiana; from Sciurus carolinensis at Bayou Saint Louis,
Mississippi; from Sciurus kaibabensis from the Kaibab National Forest,
Arizona* from Sciurus aberti at Estes Park, Colorado; from Sciurus alleni,
Co Ionia’ Garcia, Chihuahua, Mexico; from Sciurus oculatus, state of Vera

Cruz, Mexico. . .... ,

NOTES- The records given above are those of Ferns, which have been ac

cepted by Werneck. Specimens from Sciurus apache- in Chihuahua, Mexico, re¬
ferred by Ferris to longiceps have been placed by Werneck in Enderleinellus
arizonensis as have specimens from Sciurus arizonensis in Arizona and from
Sciurus nayaritensis, state of Zacatecas, Mexico.

109

Enderleinellus malaysianus Perris

1919. Enderleinellus malaysianus Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 1:12; figures 3> 4.

HOSTS AND DISTRIBUTION. Type from Callosciurus caniceps (as Sciurus
lucas) from Saint Lukes Island, Mergui Archipelago, Malaysia. Also record¬
ed from the same host (as Sciurus bentinckanus) from Bentinck Island in the
same archipelago, (as Sciurus domelensis) from Domel Island in the same
archipelago, (as Sciurus lancavensis) from Pulo Teratau, and (as Sciurus
davisoni) from Trong, lower Siam; and from Callosciurus prevosti (as Sciurus
borneoensis) at Pulo Kanchut, Borneo.

Enderleinellus marmotae Ferris

1919. Enderleinellus marmotae Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 1:47; figure 30.

HOSTS AND DISTRIBUTION. Type from Marmota monax from Grafton, South
Dakota; also recorded from the same host at Elk River, Minnesota, and Marble
Cave, Missouri, at Sandy Springs, Maryland, and Washington, District of
Columbia.

Enderleinellus menetensis Ferris

1919. Enderleinellus menetensis Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 1:14; figures 5> 6.

HOSTS’ AND DISTRIBUTION. From Menetes berdnorei, Roh Rut Island, south¬
east Siam.

Enderleinellus mexicanus Werneck

1947. Enderleinellus mexicanus Werneck, Memorias do Instituto Oswaldo Cruz
45:289; figures 16-18.

HOSTS AND DISTRIBUTION. Type from Sciurus truei at Chacala, Mexico.
Also recorded from Sciurus nelsoni from the state of Morelos, and from
Sciurus colliae, state of Nayarit, Mexico.

NOTES. The specimens recorded above were recorded by Ferris under Ender¬
leinellus extremus.

Enderleinellus microsciuri Werneck

1947. Enderleinellus microsciuri Werneck, Memorias do Instituto Oswaldo
Cruz 45:287; figures 10-12.

HOSTS AND DISTRIBUTION. From Microsciurus mimulus from unspecified lo¬
cality in Colombia.

NOTES. The specimens upon which this species was based were previously
recorded by Ferris as Enderleinellus kelloggi.

Enderleinellus nannosciuri Ferris

1919. Enderleinellus nannosciuri Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 1:30; figure 17.

HOSTS AND DISTRIBUTION. From Nannosciurus melanotis from Batavia, Java.

Enderleinellus nitzschi Fahrenholz
Figures 48, 49

1818. Pediculus sphaerocephalus Nitzsch, Gennar's und Zincken' s Magazin fur
die Entomologie 3:305- (Not Pediculus sphaerocephalus von Olfers)

110

Enderleinellus nitzschi Fahrenholz Figure 48

1912. Enderleinellus sphaerocephalus (Nitzsch), Fahrenholz, Jahresbericht
des niedersachsischen Zoologischen Vereins zu Hannover 2-4: 52;
text figures 22, 23; Plate 2, figures 5-7. .

1916. Enderleinellus nitzschi Fahrenholz, Archiv fur Naturgeschichte, Ab-

teilung A, 81:29. . .

1919. Enderleinellus nitzschi Fahrenholz, Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 1:8; figures 1, 2.

HOSTS .AND DISTRIBUTION. Originally described from Sciurus vulgaris in
Europe and recorded numerous times from this host. Recorded also from
Sciurus anomalus (as syriacus) from Syria; from Sciurus huds on icus from
Alaska; from Sciurus douglasii from western United States; from Sciurus

frenonti from Colorado. , , , ,

NOTES. Wemeck, who examined all the material recorded above, has agreed

that no specific differences exist among the specimens from the different
hosts .

Ill

head.

antenna

------

female genitalia

Enderleinellus nitzschi Fahrenholz, details

3rd claw

male genitalia
Figure 49

112

Enderleinellus osborui Kellogg and Perris

1915* Enderleinellus osborni Kellogg ,uid Ferris, Anoplura and Mallophaga
of North American Mammals, Stanford University Publications, Uni
versity Series (no volume number), page 43; text iigure 15; Plate
4; figure 11; Plate 6, figure 6.

1919. Enderleinellus osborni Kellogg and Ferris, Ferris, Contributions
Tow;ird a Monograph of the Sucking Lice, Part 1:46; figure 29.

HOSTS AND DISTRIBUTION . Type from Citellus beecheyi (as douilasi) at
Covelo, Mendocino County, California, and recorded from this host species
at various localities in California. Also from Citellus varieiatus (as
buckleyi) in Texas and (as g rammurus ) in Arizona. From Citellus (as
Xerospernophi lus) teret icaudus, Imperial County, California.

Enderleinellus platyspicatus Ferris

1919. Enderleinellus platyspicatus Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 1:26; figures 14. 15.

HOSTS AND DISTRIBUTION . From Funambulus palmarum (as tristriatus) in
Ceylon.

Enderleinellus replicatus Redikorzev

1937. Enderleinellus replicatus Redikorzev, Parasitology 29:4; figure.

HOSTS AND DISTRIBUTION. From Set uropterus volans, Tartar Republic, with¬
out closer indication of locality.

NOTES- Members of this genus very commonly in dying contract in such a
manner that the head is drawn back over the body as the result of opisthot¬
onos, and iu preparing specimens for study they must be unfolded. Unlortu-
nately , in describing this species its author made his illustrations from a
specimen that was thus contracted and consequently his illustration is en¬
tirely useless. It is impossible to determine what the species is actually
like and to include it in the key.

Enderleinellus sciurotamiasis Ferris

1919. Enderleinellus sciurotamiasis Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 1:45; figures 20, 21.

HOSTS AND DISTRIBUTION. From Sciurotamias davldianus, Shensi Province,
China.

Enderleinellus suturalis (Osborn)

1891-

1915.

1916.

1919.

1929.

Eaematopinus suturalis Osborn, United States Department of Agricul¬
ture, Division of Entomology, Bulletin (old series) 7:27; tig. 15-
Enderleinellus suturalis (Osborn), Kellogg and Ferris, Anoplura and
Mallophaga of North American Mammals, Stanford University Publica¬
tions, University Series (no volume number), page 40; Plate 4,
fiiru re 9*

Enderleinellus suturalis occidentalis Kellogg and terns, Anoplura
and Mallophaga of North .American Mammals, Stanford University Pub¬
lications, University Series (no volume number), page 42; Plate 2,
fiim re 3; Plate 4, figure 10; Plate 5, figure 17.

Enderleinellus suturalis (Osborn), Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 1:42; figures 26, 27, 28.
Cyclophthlrus suturalis (Osborn), Ewing, A Manual ot External Para¬
sites, page 196.

HOSTS .AND DISTRIBUTION. Originally described as from Citellus tndecim-

113

lineatus and Citellus franklini (as Spermophilus) at Ames, Iowa. The first
of these has been designated by Kellogg and Ferris as the type host. Ferris
has recorded the species from a long series of species of Citellus as fol¬
lows: eversmanni, Altai, Siberia; mongolicus , Kansu, China; beldingi,
California; elegans, Colorado; franklini, North Dakota; mollis, Nevada;
osgoodi, near Circle, Alaska; townsendi , state of Washington; tridecimline-
atus, Kansas and Oklahoma; madrensis (as Callopsermophi lus) , Chihuahua,
Mexico; nelsoni (as Ammospermophi lus) , California; Cynomys gunnisoni,
Colorado; Cynomys leucurus, Colorado and Wyoming.

NOTES. In the rather extensive material at hand there is a quite wide
range of variation in various details, but a study of the material by Dr.
Edwin Cook has shown no clear basis for recognizing more than one species.
The two species, osborni and marmotae, which have in the past been separ¬
ated from suturalis are the only ones in which the situation seems reason¬
ably clear. Studies are continuing with an accompanying attempt to secure
additional material and it is hoped eventually to publish a detailed report
upon the group.

Enderleinellus tamiasis Fahrenholz

1916. Enderleinellus tamiasis Fahrenholz, Archiv fiir Naturgeschichte, Ab-
teilung A, 81:11:27; text figure 22.

1919. Enderleinellus tamiasis Fahrenholz, Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 5:288; figure 176.

HOSTS AND DISTRIBUTION. Known only from the original record from Tanias
striatus in the Berlin Zoological Garden.

NOTES- It is entirely possible that Tamias striatus, which is a native
of eastern and central United States, is not the normal host of this spe¬
cies, no species of Enderleinellus having yet been recovered from this host
under natural conditions.

Enderleinellus urosciuri Werneck

1937. Enderleinellus urosciuri Werneck, Memorias do Instituto Oswaldo Cruz
32:400; figure 12.

HOSTS AND DISTRIBUTION. Recorded from Sciurus (as Urosciurus) ignivent-
ris from Acajutuba, Rio Negro, state of Amazonas, Brasil.

Enderleinellus venezuelae Ferris

1919. Enderleinellus venezuelae Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 1:25; figure 13.

1948. Enderleinellus venezuelae Ferris, Werneck, Memorias do Instituto Os¬
waldo Cruz 45:292; figures 22-24.

HOSTS AND DISTRIBUTION. Type from Sciurus griseogena from Macuto, Vene¬
zuela. Also recorded from the same host (as meridensis) from Merida, Vene¬
zuela, and from Sciurus gerrardi (as versicolor) from Rio Aurare, Venezuela.

Enderleinellus zonatus Ferris

1919. Enderleinellus zonatus Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 1:32; figures 18, 19. (In part)

1948. Enderleinellus zonatus Ferris, Werneck, Memorias do Instituto Oswaldo
Cruz 45:295; figure 28.

HOSTS AND DISTRIBUTION. type from Paraxerus ochraceus (as jacksoni) ,
Kijabe, British East Africa.

NOTES. Werneck has pointed out that two distinct species were included
in the material recorded by Ferris.

114

Genus MICROPHTHIRUS Ferris

1914. Microphthirus Ferris, Contributions Toward a Monograph of the Sucking
Lice, Part 1:49.

GEKBRIC TYPE, Enderleinellus unclnatus Ferris.

CHARACTERS. Enderleinellinae iu which the pair ol 1 ittie, scleroti zed
plates on the venter ol the second abdominal segment is lack ing . Paratergal
plates of se>jments 3"6 each continuous with the corresponding sternal plate,
being connected with it by a narrow, scleroti zed bridge. Paratergal plates
not free from the body at any point. Antennae with the proximal, postaxial
angle of segments 2-3 produced into a scleroti zed hook.

NOTES. As known at present this genus contains but a single species.
This is a very pecuLiar form, known only from North American flying squir¬
rels of the genus Glaucomys. The assignment of the genus to the Enderleiu-
ellae is based entirely upon the character of the legs, since there is lit¬
tle else to connect it with any other group of the Anoplura. The host as¬
sociation supports this assignment and iu this particular case has been
taken into consideration iu placing the genus.

Microphthirus uncinatus (Ferris)

Figures 50, 51

1916. Enderleinellus unclnatus Ferris, Psyche 23:108; figures 6, 7.

1919. Microphthirus uncinatus (Ferris), Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 1:49; figures 31-32.

HOSTS AND DISTRIBUTION. Known only from the original record from Glauc¬
omys sabrinus at Yosemite National Park, California. The host is a member
of the rodent family Sciuridae.

NOTES. This is one of the very' smallest of all sucking lice, the male
attaining a Length of only about .35 mm. The insects are so small that as
seen upon their° host they are very likely to be mistaken for }oung of one
of the other species which occur on these squirrels.

Genus WERNECKIA Ferris, new genus

GENERIC TYPE- Enderleinellus minutus Werneck. One other species, Ender¬
leinellus paraxeri Werneck, is here included.

CHARACTERS. Enderleinellinae in which the paired sclerotizations 01 the
second abdominal sternum are lacking. Differing from Microphthirus in
which this also is true, in not having the paratergal plates connected with
the sternites by sclerotizations.

Occurring on African squirrels of the genus Paraxerus.

NOTES. Since each of the two included species is known from but a single
male very little can be said about the group. It is by no means certain
that these two species should be placed in the same genus, since thej dil-
fer materially in various respects and it appears possible that they are
independent derivatives from species of Enderleinellus that also occur on
their hosts.

Key to Species of WEFJsECKIA

Genitalia of the male with all parts of the terminal complex slender - ...

. PARAXERI

Genitalia ’of ’the' male’ with the parts of the terminal complex short the
parameres broadened . .

115

Microphthirus uncinatus (Ferris)

female genitalia

male genitalia

Microphthirus uncinatus (Ferris), details

Figure 51

117

Werneckia minuta (Werneck)

Figure 52

Werneckia minuta (Werneck)

Figure 52

1947. Enderleinellus minutus Werneck, Memorias do Instituto Oswaldo Cruz
45:296; figure 30-

HOSTS AND DISTRIBUTION. Known from but a single male, from Paraxerus
jacksont i=ochraceus) , from Kijabe, British East Africa.

NOTES. The single male representing this species was discovered by Wer¬
neck among the type material of Enderleinellus zonatus Ferris and described
by Werneck who, however, did not note the absence of the paired ventral
plates of the second abdominal segment.

Werneckia paraxeri (Werneck)

1947. Enderleinellus paraxeri Werneck, Memorias do Instituto Oswaldo Cruz
45:295; figure 29.

HOSTS AND DISTRIBUTION. Known from but a single female, from Paraxerus
palliatus, from British East Africa.

NOTES. The circumstances surrounding this species are the same as those
connected with minuta.

118

Subfamily HOPLOPLEUKINAE Ferris, new subfamily

DESCRIPTION OE THE SUUFAMILY . Hoplopleuridae in which there is no evi¬
dence of eyes. Paratergal plates reaching an extreme degree ol development,
those of abdominal segments 4-6 enclosing the sides of the abdomen and each
to some extent overlapping that of the segment next succeeding. Sternal
plates of abdominal segment two prolonged laterally on each side to articu¬
late with the corresponding paratergal plate and at times divided mesaliy
by a membranous area into two plates which may' be much expanded. Anterior
legs always small, with slender claw, the middle legs larger with stouter
claw, the posterior legs still larger, generally flattened and with very-
broad claw. Antennae 4—6 segmented, never sexually dimorphic.

NOTES. In its basic pattern this group is quite homogeneous, although
some of its species present some extraordinary' specializations. The group
is as a whole probably the most specialized ol the AnopLura.

The five included genera may be separated by the following key.

Key to the Genera of HOPLOPLEUKINAE

Antennae clearly 5-se^ment'etl . \

Antennae 4-segmented . .4

Paratergal plates of abdominal segment 2 each prolonged into a Made-
like process which projects from the tody wall . PTER0PHT1URUS

Not so . - 3

Sternal plate of abdominal segment 2 divided longitudinally into Z much

expanded plates . SCHIZDPHTHIRUS

Not g0 . . . ....HOPLOPLEURA

Posterior le fs with a membranous, bladder-like expansion on the coxa

Not so . ANCISTROPLAX

Genus ANCISTROPLAX Waterston
1929* Ancistroplax Waterston, Parasitology' 21:161.

1932. Ancistroplax , Ferris, Contributions Toward a Monograph of the Sucking
Lice, Part 5:38.

GENERIC TYPE. Ancistroplax crocidurae Waterston.

CHARACTERS. Moplop leurinae in which the antennae are 1 our-segmented .
Paratergal plates of abdominal segments 3-7 each having the appearance of
bein'1, divided into two equal parts by a longitudinal line of weak scleroti-
zation. Terml and sternal plates of the abdomen strongly developed in
both sexes, the female having three plates and three rows of setae, both
dorsally and ventrally, on segments 3-6; the male having but one plate,
either dorsallv or ventrally, on any segment, but this on segments 4 ,

both dorsally and ventrally having two rows of setae and presenting an ap¬
pearance which suggests that it is composed of two transverse plates which
have partially fused. Tergal plate of segment six of the male having its
posterior angles each produced into a free process which is bent apically
toward the mid-line of the body. Posterior legs strongly expanded and flat¬
tened, with broad claw. Sternal plate of abdominal segment two divided Ion
kritudinally into two expanded plates. First sternal plate of segment three
not produced laterally to articulate with the corresponding paratergites.

NOTFS The affinities of this genus seem to be most closely with Schiz
ophthirus. It seems also to be more or less closely related to the genus
Baemtopinoides.

119

Ancistroplax crocidurae Waterston

Figures 53 > 54

1929. Ancistroplax crocidurae Waterston, Parasitology 21:161; figures.
1932. Ancistroplax crocidurae Waterston, Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 5 : 308 ; figures 188, 189-

HOSTS AND DISTRIBUTION. There is but a single record of this species
which attributes it to a shrew, Crocidura horsfieldi, in Ceylon.

Genus HAEMATOPINOIDES Osborn

1891. Haematopinoides Osborn, United States Department of Agriculture,
Division of Entomology, Bulletin (old series) 7:28.

1896. Euhaematop inus Osborn, United States Department of Agriculture,
Division of Entomology, Bulletin (new series) 5:186.

1929. Haematopinoides, Ewing, A Manual of External Parasites, page 140.
1932. Haematopinoides, Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 5:289-

GENERIC TYPE. Haematopinoides squamosus Osborn. The genus Euhaemato-
pinus Osborn, with Euhaematop inus abnormis Osborn as type, is synonymous,
its type species being synonymous with squamosus.

CHARACTERS. Hoplopleurinae in which the antennae are 4-segmented. Pos¬
terior legs with a membranous, bladder-like expansion arising from the an¬
terior margin of the coxae and a similar structure on the anterior wall of
the tibia. Apart from the usual sclerotizations of the terminal and geni-
talic segments, tergal plates are developed only on segments 1-3 and ster¬
nal. plates on segments 2-3 in the female, while they are present on all
segments except the first in the male. Female with two rows of setae both
dorsally and ventrally on most of the abdominal segments, the male with but
one. Sternite of the second abdominal segment divided longitudinally into
two expanded plates each of which articulates by a lateral extension with
the corresponding paratergal plate. First plate of abdominal segment three
not produced laterally.

NOTES. This genus (as Euhaematop inus) was employed by Enderlein in 1904
as type of the subfamily Euhaematop ininae of the family Haematopinidae and
in 1929 this was elevated to the rank of a family, under the name Haemato-
pinoididae, by Ewing. Actually, the genus is very closely related to
Hoplopleura. The membranous expansions on the posterior legs are unique
structures but do not in any way justify the assignment of the genus to an
isolated position.

Haematopinoides squamosus (Osborn)

Figures 55> 56

1891- Haematopinoides squamosus Osborn, United States Department of Agri¬
culture, Division of Entomology, Bulletin (old series) 7:28; fig¬
ure 16.

1896. Euhaematop inus abnormis Osborn, United States Department of Agricul¬
ture, Division of Entomology (new series) 5:18?; figure.

1922. Euhaematop inus abnormis Osborn, Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 3: 150 > figures 98, 99-
1929. Haematopinoides squamosus Osborn, Ewing, A Manual of External Para¬
sites, page 140.

HOSTS AND DISTRIBUTION. First recorded, undoubtedly quite erroneously,
from a rodent of the family Geomyidae in Iowa. Later described again as a
new genus and species from its true host, a mole, Scalopus aquaticus, at
Ames, Iowa, United States. It has since been recorded from this host from
localities in Kansas, Illinois, and New York. Also specimens are at hand

120

Ancistroplax crocidurae VVaterston

Figure 53

121

Figure 54

Ancistroplax crocidurae Waterston, details

123

thoracic sternal plate male genitalia
Haematopinoides squamosus Osborn, details

female genitalia
Figure 56

3rd leg

head

antenna

124

from another mole, Parascalops brewer l, from eastern United States.

NOTES. Ewiug has recorded finding the types ol Haematopinoides squano: ti¬
lts, which had presumably been lost, and has confirmed the suspicion, previ
ously expressed by Ferris, to the effect that Euhaematopi nus abnormis is
the same species. Apparently Osboni was misled into naming two genera sim¬
ply by differences in the nature of the preparations then at hand.

Genus HOPLOPLEURA Euderlein

1904. Hoplopleura Enderlein, Zoologischer Anzeiger 28:221.

1921. Hoplopleura, Ferris, Contributions Toward a Monograph of the Suck¬
ing Lice, Part 2 : 59 -

1929. Ctenura Ewing, A Manual of External Parasites, page 199 -
1929. Euhoplopleura Ewing, A Manual of External Parasites, page 198.

1929. Ctenopleura Ewing, A Manual of External Parasites, page 199 •

1929. Ferrisella Ewing, A Manual of External Parasites, page 199.

GENERIC TYPE. Pediculus acanthopus Burmeisler.

GENERIC SYNONYMS. Ctenopleura Ewing, type Hoplopleura crypt lea Ferris;
Ctenura Ewing, type Hoplopleura pectinate Cummings; Euhoplopleura Ewing,
type Hoplopleura trisp inosa Kellogg and Ferris; Ferrisella Ewing, type
Hoplopleura ochotonae Ferris.

CHARACTERS, tloplopleurinae in which the sternal plate of abdominal seg¬
ment two is always, and the first sternal plate is usually, extended later¬
ally to articulate with the corresponding paratergites, these two plates
always being narrowly transverse. First sternal plate ot segment three
usually with° two or three enlarged, stout setae in two groups, rarely these
not developed. Paratergites of the abdominal segments never showing any
indication of a partial longitudinal division into dorsal and ventral parts.
Antennae clearly five-segmented. Posterior legs with no bladder-like, mem¬
branous expansions. Paratergites of abdominal segment two not produced in¬
to a long, blade-like extension.

NOTES. This genus, as here understood, forms a very homogeneous group.
In two or three species the first sternal plate of abdominal segment 3 does
not attain the paratergal plate of the segment but any attempt to separate
these species into a different genus on this basis seems to gain nothing
and would result only in the formation of an evidently unnatural grouping.

The four genera named by Ewing and listed above do not, in the opinion
here held, merit recognition, but the genus Pterophthirus, named by Ewing,
is here accepted.

Key to Species of Hoplopleura

1. First sternal plate of abdominal segment 3 extended laterally to ap¬
proximate or articulate with the corresponding paratergal plates....?
First sternal plate of abdominal segment 3 not extended laterally . 2

2 (1). Paratergal plates of abdominal segments 4-6 in the female and ot at

least segment 4 in the male with a pair ot short, stout, apically
flattened and truncate setae; known from Octodontomys in Bolivia.
. DISGREGA

Not so, all setae of the paratergal plates apically acute. ....... .3

3 (2). Paratergal plates of abdominal segment 6 with the dorsal apical an¬

gle prolonged into a thumb-like process, the ventral angle round¬
ed; known from Octodon in Chile . CHILENSIS

Otherwise. #••••••••••••••••••• . • • • . . . • *

4 (3) . Paratergal plates of abdominal segments 4-6 with the posterior mar¬

gin almost straight, the posterior angles each forming but a
slight tooth; known from the squirrel genus Sciurotamias in China
* . EMARGINATA

125

Paratergal plates of abdominal segments 4-5 with the posterior mar¬
gin presenting a deep, median emargination between the two broad
lateral lobes; known from Hydromys in Australia . BIDENTATA

5 (1). Paratergal plates of abdominal segment 8 in both sexes with each

posterior angle prolonged into a tapering apically acute lobe.. 6
Paratergal plates of abdominal segment 8 never with more than one
apical lobe or process, usually with none . 7

6 (5)- Paratergal plates of abdominal segment 2 with each posterior angle

prolonged into a tapering, apically acute point; known from unde¬
termined rodent in Brasil . BRASILIENSIS

Paratergal plates of abdominal segment 2 with the posterior margin
truncate, the angles not at all prolonged into points; female
with the apex of the abdomen bearing a comb of stout setae; known
from Rattus surifer in the Malayan region . PECTINATA

7 (5) • First sternal plate of abdominal segment 3 with two groups of def¬

initely enlarged setae . 8

First sternal plate of abdominal segment 3 without such pairs of
enlarged setae; occurring on Ochotona in Asia . 0CH0T0NAE

8 (7). Enlarged setae of the first sternal plate of abdominal segment 3

arranged in two groups of 3 setae each . 9

Enlarged setae of this plate arranged in two groups of 2 setae each

. . 11

9 (8) . Paratergal plates of abdominal segments $-6 each with the posterior

border bearing 4 prominent, almost equal points or processes. .. 10
Paratergal plates of these segments with merely the apical angles
prolonged into points; occurring on North American flying squir¬
rels of the genus Glaucomys . TRISPINOSA

10 (9). Paratergal plates of abdominal segments 2-6 each borne upon a dis¬

tinct prominence; tergal and sternal setae of the abdomen ex¬
tremely numerous and heavy; known from Tatera liodon in Africa..

. CRYPTICA

Paratergal plates of abdominal segments 2-6 not thus borne upon
prominences; setae of abdominal tergites and sternites slender

and of normal number; known from Tatera boehmi in Africa .

. . VEPRECULA

11 (8) . Paratergal plates of abdominal segment 8 with 1 apical angle pro¬

duced into an elongated, apically free process . 12

Paratergal plates of abdominal segment 8 entirely devoid of any

apical process or with only a very slight lobe . 14

Paratergal plates of abdominal segments 3~6 each with the poste¬
rior margin divided into 4 slender processes of which the dor¬
sal process is longer than the plate itself; probably normal to

Lemniscomys in Africa . PELOMYDIS

Paratergal plates of abdominal segments 3~6, if divided into

lobes, having the dorsal lobe no longer than the others . 13

Paratergal plates of abdominal segment 7 with the posterior mar¬
gin divided into 2 equal major lobes, each of which is divided
into 2 equal minor lobes; known from Arvicanthis in Africa .

. LATICEPS

Paratergal plates of segment 7 with the posterior margin divided
into 2 lobes of which one is apically acute and the other apic¬
ally truncate or slightly emarginate; known from various hosts

in Africa . INTERMEDIA

Paratergal plates of abdominal segment 7 with at least one of the
apical angles produced into a lobe which is in some instances

as long as the body of the plate . 15

Paratergal plates of segment 7 with neither of its apical angles
thus produced . 27

12 (11).

13 (12).

14 (11).

126

15 (14).

16 (15).

17 (16).

18 (16).

19 (18).

20 (18).

21 (20).

22 (21).

23 (15).

24 ( 23).

25 (24).

Paratergal plates of abdominal segment 7 with but one of its ap

* , + s

ical angles produced into a lobe or poiut . lo

Paratergal plates of abdominal segment 7 with each of its apical

angles produced into a lobe or point . 23

Paratergal plates of segments 3~5 each with the posterior mai-gin
divided into 4 slender lobes or processes which equal at least

half the length of the body of the plate . 17

Paratergal plates of abdominal segments 3_5 with the posterior
margin divided into 2 major lobes, each of which may be apical-
Lv more or less emarginate or deeply divided into 2 minor lobes

7. . 7 . is

Dorsal apical process of the paratergal plates of abdominal seg¬
ments 3-7 very long, longer than the body of the plate; known

from Lemniscomys in Africa . ENORMIS

Dorsal apical process on no paratergal plate longer than the body

of the plhte or longer than the other processes . NEUMANNI

Ventral lobe of paratergal plates of abdominal segment 3 divided
into 2 apically rounded, equal lobes; from species ol Oryzomys

in the New World . ;19

Ventral lobe of pai*atergal plates of abdominal segment 3 otherwise
. 20

Ventral lobe of abdominal segment 6 divided into 2 equal lobules;

known from Oryzomys in southeastern United States. .. .ORYZ0MY D1S
Ventral lobe of abdominal segment 6 forming a single, acute point;

known from species of Oryzomys in South America. . .NES0RYZ01YDIS
Ventral lobe of paratergal plates of abdominal segment 3 acute...

. ; . 7 . AFFINIS

Ventral lobe of paratergal plates of abdominal segment 3 broadly

truncate or apically emarginate . 21

Ventral lobe of paratergal plates of abdominal segment 6 acute;

known from the genus Apomys in the Philippine Islands. . APOMYDIS
Ventral lobe of paratergal plates of segment 6 truncate or emar¬
ginate . ^

Ventral lobe of paratergal plates of abdominal segment 6 deeply

emarginate; known from Dasymys yelukus in Africa . SOMERENI

Ventral lobe of paratergal plates ol abdominal segment 6 truncate,

known from various rodents in South America . TRAVASSOSI

Both dorsal and ventral apical lobes ol paratergal plates ol ab¬
dominal segment 7 apically acute . ......24

At least one of the lobes of the paratergal plates of abdominal

segment 7 apically acute . . .

Paratergal plates of abdominal segments 3— ^ with the postenoi
margin divided into 4 nearly equal, apically rounded lobes, oc¬
curring on the iTenera Nectomys and Oryzomys in South America...

. 77 . 7 . QUADRIDENTATA

Paratergal plates of abdominal segments 3_6 with the posterior
margin divided into 2 apically truncate or slightly emarginate
1 b& • 2 1

Paratergal plates of abdominal segments 4-6 with the setae ol the
posterior margin extremely minute; occurring characteristically
on members of the New World genera Peromyscus and Onychomys,
but recorded also from Hus musculus in North America and Eurasia

. . HESPEROMYDIS

Paratergal plates of abdominal segments 4-6 with the setae of
their posterior margin almost equaling in length the depth ol
the median emargination in which they are placed; occurring as
far as known on the genus Cricetulus in China . CKloMlLi

127

26 (23).

27 (14).

28 (27).

29 (28).

30 (28).

31 (30).

32 (30).

33 (32).

34 (33).

35 (34).

36 (33).

37 (36).

Dorsal lobe of the paratergal plates of abdominal segment 7 defi¬
nitely truncate, the ventral lobe acute; from members of the

New World genus Reithrodontomys . REITHRODONTOMYDIS

Both dorsal and ventral apical lobes of the paratergites of ab¬
dominal segment 7 narrowly rounded apically; known from Mus

triton in Africa . .RUKENYAE

Female with 3 rows of setae dorsally and ventrally on most of the

abdominal segments . 28

Female with but 2 rows of setae both dorsally and ventrally on
most of the abdominal segments; probably normal to species of

Tatera in Africa . BISERIATA

Paratergal plates of abdominal segment 6 with the posterior angles

not produced into points . 29

Paratergal plates of abdominal segment 6 with at least one of the

posterior angles produced into a definite point or lobe 30

Sternal plate of the thorax broadly triangular, occurring on the

North American squirrel genus Talhlas . ERRATICA

Sternal plate of the thorax narrowly produced posteriorly; known

from the Asiatic genus Phaiomys . PHAIOMYDIS

Both dorsal and ventral basal angles of the paratergal plates of
abdominal segments 4-5 with a process which projects toward the

mid-line of the body . SI

Basal angles of these paratergal plates without such processes. 32
Outer seta of the pairs of enlarged setae on the first sternite
of abdominal segment 3 longer than the other and distinctly

bent; known from the squirrel genus Callosciurus in China .

. DISTORTA

Outer seta of these pairs not longer than the other and not bent;
known from the squirrel genus Callosciurus in the Malayan area.

. ERISMATA

Paratergal plates of abdominal segment 6 with 4 almost equal,
slender processes on its posterior border; known from Mylomys

cuninghamei in Africa . MYLOMYDIS

Paratergal plates of abdominal segment 6 without such processes..

. . 33

Paratergal plates of abdominal segments 4-6 without setae, even

of the most minute size . 34

Paratergal plates of abdominal segments 4-6 definitely with setae,

even if small, on the posterior margin . 36

Both dorsal and ventral lobes of the paratergal plates of abdom¬
inal segment 3 broadly truncate; occurring on Micromys in Europe

. LONGULA

Both dorsal and ventral lobes of the paratergal plates of abdom¬
inal segment 3 apically acute . 35

Dorsal apical lobe of the paratergal plates of abdominal segments
4— 5 apically broad and emarginate; known from Oxymycterus judex

in South America . FONSECAI

Ventral apical lobe of the paratergal plates of abdominal segments

3-6 acute; from Oxymycterus in Peru . OXYMYCTERI

Apical lobes of paratergal plates of abdominal segment 6 both a-

cute . 37

Apical lobes of paratergal plates of abdominal segment 6 other¬
wise . 42

Apical lobes of the paratergal plates of abdominal segment 3 with
the dorsal lobe apically acute, the ventral lobe broad and
slightly emarginate; occurring on various members of the sub¬
family Microtinae of the Muridae in Europe, North America, and

certainly Asia . ACANTHOPUS

128

38 (37).

39 (33).

40 (39).

41 (40).

42 ( 36).

43 (42).

44 (43).

45 (44).

46 (43).

Apical lobes ot‘ the paratergal plates of abdominal segment 3 otb

erwise . 38

Paratergal plates of abdominal segments 4—5 withadeep, rectangu¬
lar, median emargination in the posterior border, this dividing
the plate into two broad, apical lobes; known from Reithrodon

in Argentina . ARGENTINA

Paratergal plates of abdominal segments 4->5 without such emargina-

tion ol the posterior border . 39

With but 1 or occasionally 2 setae between the ends ol the tergal
and sternal plates of the abdomen and the corresponding para-
teraal plates; known from the squirrel genus Kunambulus in Asia

. . MANICULATA

With not less than 2 and usually more setae between the ends ol
the tergal and sternal plates of the abdomen and the correspond-

ing paratergal plates . . . . 40

Paratergal plates o f abdominal segments 4-6 with the apical angles

strongly produced and acute, the posterior border of the plate

arcuately and quite deeply emarginate . . . . . .

Paratergal plates of abdominal segments 4-6 with the margin be¬
tween the toothed apical angles almost straight, not at all e—
martrinate; occurring on the genus Siimodon in North America....

. . . . . . 7 . HIRSUTA

Parameres of the male apically notched; occurring on the genus

Neotamias (better known as Sutamias ) in North America .

. ARB0R1C0LA

Parameres of the male apically entire; known from the genus Sciurus

in North America . . SCIURICOLA

Paratergal plates of abdominal segment 6 with the dorsal apical
angle forming a slight tooth, the ventral angle without a tooth,

known from Phyllotis in South America . ....REDUCT A

Paratergal plates of abdominal segment 6 with both apical angles

produced into points or lobes . . . 43

Ventral apical lobe of paratergal plates of abdominal segment b

acute, the dorsal lobe more or less truncate . .*44

Dorsal and ventral apical lobes ol paratergal plates ol abdominal
segment 6 essentially similar, truncate or apically serrate or

emarginate . ••***, . . . . .

Paratergal plates of segments 3~5 each with 1 very minute and 1
longer seta on the posterior margin; known from various hosts,

including domestic rats, in various parts of the world - .....

° 0EN0MYDIS

Paratergal plates of abdominal segments 3—5 each with 2 setae
which Tire as long as or longer than the depth of thd emargina-

tion in which they are placed . /“o'c '

Ventral apical lobe of the paratergal plates ol segments j-h a-
cutelv pointed; known from the genus Chrotonys in the Philippine
Islands. . ! . ...CHR0T0MYDIS

Ventral apical lobe of paratergal plates of abdominal segments
3_5 narrowly rounded or truncate apically; known trom nottu s

sabanus in the Malayan area . . . . MALAYSIAN A

Paratergal plates of abdominal segments 4-6 each with 1 very min
ute seta and 1 seta which is about as long as the depth ol the
emargination in which it is placed; known from Rhipidonys in

South America. . . . * V - - ' ‘ *V *2

Parateral plates of abdominal segments 4-6 each with 2 setae ol
nearly equal length, these as long as or longer than the depth
of the emargination in which they are placed ; known fro™
in China . MERIONIDIS

129

Hoplopleura acanthopus (Burmeister)

Figures 57, 58

1839- Pediculus acanthopus Burmeister, Genera Insectorum, Rhynchota, Num¬
ber 5; Plate 1, figure 2.

1842. Haematopinus acanthopus (Burmeister), Denny, Monographia Anoplurorum
Britanniae, page 25; Plate 24, figure 3-
1880. Haematopinus acanthopus (Burmeister), Piaget, Les Pediculines, page
638; Plate 52, figure 4.

1904. Polyplax acanthopus (Burmeist;cr) , Enderlein, Zoologischer Anzeiger
28:142.

1904. Hoplopleura acanthopus (Burmeister) , Enderlein, Zoologischer Anzeiger
28:221; figures 1, 2.

1915* Hoplopleura acanthopus, variety americanus Kellogg and Ferris, Ano-
plura and Mallophaga of North American Mammals, Stanford University
Publications, University Series (no volume number), page 16 ; text
figure 3; Plate 4, figure 2; Plate 5, figure 8.

1916. Hoplopleura acanthopus, variety aequidentis Fahrenholz, Archiv fur
Naturgeschichte, Abteilung A, 81:26; figure 21b.

1921. Hoplopleura acanthopus (Burmeister), Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 2:63; figures 34, 35 •

HOSTS AND DISTRIBUTION. First described from Microtus (as Arvicola )
arvalts, somewhere in Europe. It has since been recorded from Microtus
airestis and Microtus nival is in Europe ; from Microtus constr ictus, Microtus
californicus, Microtus intermedius, and Microtus ochroiaster in various
parts of the United States; from Lemmus alaskensis in Alaska; from Synapto-
mys borealis at Athabasca Landing in Canada; from Pitymys pinetorum in eas¬
tern United States. It has also been recorded from Mus musculus in Rumania
and Mus spicileQus in Sweden, these records very probably being due to con¬
tamination or straggling.

NOTES. An examination of the abundant material at hand offers no excuse
for the naming of the supposed variety americanus. Furthermore, on the
basis of such evidence els was given by its describer there is no apparent
reason for the supposed variety aequidentis.

Hoplopleura affinis (Burmeister)

1839- Pediculus affinis Burmeister, Genera Insectorum, Rhynchota, No. 10.
1842. Haematopinus affinis (Burmeister), Denny, Monographia Anoplurorum
Britanniae, page 36.

1904. Polyplax affinis (Burmeister), Enderlein, Zoologischer Anzeiger 28:
142.

1921. Hoplopleura affinis (Burmeister), Ferris, Contributions Toward aMon-
ograph of the Sucking Lice, Part 2:75; figures 42, 43*

HOSTS AND DISTRIBUTION. First recorded from Apodemus a£rarius and Apo-
demus sylvat icus in Europe. Later recorded by Ferris from both these hosts,
from agrarius in Germany and from sylvatlcus in Manchuria and Siberia. In
addition, Ferris has recorded the species from a number of South American
mammals, including Akodon mollis in Peru; Akodon arviculoides in Paraguay;
Akodon amnicola in Argentina; Phyllotis pictus (as Euneomys sp.) in Peru;
Phyllotls micropus in Argentina. Certain other records by Ferris from
South American mammals are here transferred to other species as will be
noted below.

NOTES. Specimens from Cricetuls incanus from China, previously referred
to affinis are here described as Hoplopleura cricetuli new species, and
specimens from Heithrodon hatcheri from Patagonia, axe referred to Hoplo¬
pleura argentine Werneck.

The records ol affinis from other South American hosts appear to be ut-

130

Hoplopleura acanthopus (Burmeister)

Figure 57

131

132

terly unreasonable, but a re-examination of the material at hand offers no
basis for their separation. It would seem very probable that biologically
more than one species is involved, but nothing morphological appears that
will justify a separation.

Hoplopleura angu^Lata Ferris

1921. Hoplopleura angulata Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 2:73; figures 40, 41.

HOSTS AND DISTRIBUTION. Type from Rhipidomys uenezuelae, somewhere in
Venezuela. Also recorded by Ferris from Rhipidomys sp. (ascertained by
Hopkins to be Rhipidomys leucodactylus) , from the Rio San Miguel, Peru;
from Rhipidomys venustus, Merida, Venezuela; and from Thomasomys cinereus,
Balsas, Peru.

Hoplopleura apomydis Ferris

1921. Hoplopleura apomydis Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 2:84; figures 49, 50.

HOSTS AND DISTRIBUTION. Described as from Apomys bardus, which is Apomys
insiinis, from Malingdang Peak, Mindanao, Philippine Islands.

Hoplopleura arboricola Kellogg and Ferris

1915. Hoplopleura arboricola Kellogg and Ferris, Anoplura and Mallophaga
of North American Mammals, Stanford University Publications, Uni¬
versity Series (no volume number), page 19; text figures 0, 7;
Plate 4, figure 8. (Part) .

1921. Hoplopleura erratica arboricola Kellogg and Ferris, Ferris, Contri¬
butions Toward a Monograph ol the Sucking Lice, Part ^.10,.

HOSTS AND DISTRIBUTION. This species occurs on members of the genus
Neotamias, which has usually been referred to in North American literature
as Eutamias, under which generic name the host records have in the past
been placed. type from Neotamias hindsi, Inverness, Mann County, Califor¬
nia. Recorded from various other species of this genus in California.

1921.

1837.

Hoplopleura argentina Werneck

Hoplopleura af finis (Burmeister) , Ferris, Contributions Toward a Mon¬
ograph. of the Sucking Lice, Part 2:75- (Part; misidentification)
Hoplopleura affinis argentina Werneck, Memorias do Instituto Oswaldo

HOSTS AND DISTRIBUTION, type from Reithrodon sp. from the Republic of
Ar 'entina. Specimens recorded by Ferris from Reithrodon hatcheri from Cor
dilleras, Patagonia, as Hoplopleura affinis, seem definitely to be this.

NOTES. The differences between this form and Hoplopleura affini s are
very slight, as indicated in the key to species, but in accord with the
practice ’here adopted of not recognizing any form below the species, it is

here regarded as a species.

Hoplopleura bidentata (Neumann)

1909. Haematopinus (Polyplax) bidentata Neumann, Archives de Parasitologie

1916. ,o$$llroblZ*faio (Neumann) , Ferris Contributions Toward a Non-
ograph of the Sucking Lice, Part 2:129; figures 86 87.

uh^tc; DISTRIBUTION. Originally described as from Rattus rattus at
Lakfl^s, Sal™ this undoubted an error in host attrrbutron. It

133

has since been recorded from what seems undoubtedly to be its normal host,
Eydromys chrysogaster, from Sydney and an undetermined locality in New South
Wales, Australia.

Hoplopleura biseriata Ferris

1921. Hoplopleura biseriata Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 2:103; fi<pire 64A.

1949. Hoplopleura biseriata Ferris, Hopkins, Proceedings of the Zoological
Society of London 119:477.

HOSTS AND DISTRIBUTION. Originally described as from Malacothrix typicus
at Bothaville, Orange Free State, Africa. Hopkins has pointed out that this
record was undoubtedly due to contamination and has recorded the species
from Tatera brantsi, Tatera lobengulae , Tatera joanae, and Tatera angolae
from unspecified localities in Africa.

Hoplopleura brasiliensis Werneck

1932. Hoplopleura brasiliensis Werneck, Comptes rendus de seances de la
Societe de Biologie de Rio de Janeiro 119:754; figure.

1932. Hoplopleura brasiliensis Werneck, Werneck, Memorias do Instituto 0s-
waldo Cruz 26:235; Plate 45-

HOSTS AND DISTRIBUTION. From an undetermined rodent from the state of
Goyaz, Brasil.

Hoplopleura chilensis Werneck

1937* Hoplopleura disgrega, variety chilensis Werneck, Memorias do Insti¬
tuto Oswaldo Cruz 32:406; figures 15, 16.

HOSTS AND DISTRIBUTION. From Octodon degus from Chile, without closer
indication of locality.

NOTES. On the basis of the very clear illustrations given by Werneck
this appears certainly to be a distinct species and not merely a "variety"
of disgrega.

Hoplopleura chrotomydis Ferris

1921. Hoplopleura chrotomydis Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 2:81; figure 46.

HOSTS AND DISTRIBUTION. Known only from the original record from a skin
of Chrotomys whiteheadi at Irisan, Benguet, Benguet, Philippine Islands.

Hoplopleura cricetuli Ferris, new species

1921. Hoplopleura af finis ( Burmeister) , Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 2:75* (Misidentification)
HOSTS AND DISTRIBUTION. Several specimens, both male and female, from
Cricetulus incanus, Shensi, China; from skin number 172,550 in the United
States National Museum.

CHARACTERS. In all respects practically identical with Hoplopleura af-
finis, as that species is described by Ferris (1921), differing significant¬
ly only in the form of the paratergites of abdominal segment seven. In af-
flnis the paratergites of this segment have the dorsal, posterior angle
produced into an elongated point which is free from the body for most of
its length and the posterior margin of the plate is straight or but slight¬
ly emarginate. In cricetuli, the dorsal lobe — while of the same shape as
in afftnts — is tree from the body onLy at its apex and the posterior margin
of the plate is deeply emarginate, this emargination forming a ventral lobe

134

which is only slightly shorter than the dorsal Lobe but is not lree I roiu
the body.

Holotype and paratypes deposited in the collections ol' Stanl'ord Univer¬
sity. Paratypes deposited in the United States National Museum.

NOTES. Complete illustrations and an extended description ol this spe¬
cies will be presented elsewhere. Enough is presented in the above de¬
scription and in the accompanying key to permit the identi 1 icatiou ol the
species.

HopLopleura cryptica Ferris

1921. Hoplopleura cryptica Ferris, Contributions Toward a Monograph ol' the
Sucking Lice, Part 2:104; figures 65, 6613, D, E, G.

1929. Ctenura cryptica (Ferris), Ewing, A Manual of External Parasites,
page 199.

HOSTS AND DISTRIBUTION. From Tatera Liodon at Kikondu, Uganda.

Hoplopleura disgrega Ferris

1921. Hoplopleura discrete Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 2:132; figures 88, 89.

HOSTS AND DISTRIBUTION. From Octodontomys simonsi, which is placed by-
Hopkins as Octodontomys gliroides, at Orura, Bolivia.

Hoplopleura distorta Ferris

1921. Hoplopleura distorta Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 2:115; figures 72C, H.

HOSTS AND DISTRIBUTION. Recorded as from Rhinosciurus sp . Hopkins, who
has investigated the matter, states that the host is actually Callosciurus

vest itus.

Hoplopleura edentulus Fahrenholz

1916. Hoplopleura acanthopus , variety edentulus Fahrenholz, Archiv iiir
Naturgeschichte, Abteilung A, 81:26; figure 21c.

HOSTS AND DISTRIBUTION. Described as from Evotomys rutilus "aus Sieben-
burgen, Kronstadt." From the description and the accompanying figure it
seems probable that this is nothing more than a slight variant ol Hoplopleura
acanthopus.

Hoplopleura emarginata Ferris

1922. Hoplopleura emarginata Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 3:130; figures 90, 91.

HOSTS AND DISTRIBUTION. Described as from Sciurotamias dav id ianus , a

skin from Shensi, China. ,

NOTES. This is a peculiar form which m some respects more cioseij re

sembles members of the genus Heohaematopinus than it does Hoplopleura. It
is retained in Hoplopleura because of the character ol the sternal plate ol
the second abdominal segment. Future authors may possibly wish either t
remove it to Heohaematopinus or to name a new genus lor it.

Hoplopleura enormis Kellogg and Ferris

1915. Hoplopleura enormis Kellogg and Ferris, Annals of the Durban Museum
1:155; Plate 16, figures 4, 4c.

1921 Hoplopleura enormis enormis Kellogg and Ferns, Ferns, Contnbu

135

tions Toward a Monograph of the Sucking Lice, Part 2:94; figures

57 , 58B, C, 59B.

HOSTS AND DISTRIBUTION. Described as from Arvicanthis dorsalis from
Mfongosi, Zululand, for which the name Lemniscomys £r iselda is now employed.
Later recorded by Perris from Lemniscomys barbarus from Gondokoro, Africa.

NOTES. The supposed subspecies of enormis named by Ferris are here re¬
garded as species.

Hoplopleura erismata Ferris

1921. Hoplopleura erismata Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 2:113; figures 72B, E, F.

HOSTS AND DISTRIBUTION. From Sciurus ferruiineus from Southeast Siam.
This host belongs to the genus Callosc iurus. Recorded also from Callosci-
urus caniceps (as Sciurus davisoni) from Lower Siam and from Callosc iurus
maclellandi (as Tamiops sp.) from Tenasserim.

Hoplopleura erratica (Osborn)

1896. Haematopinus erraticus Osborn, United States Department of Agricul¬
ture, Division of Entomology, Bulletin (new series) 5:186.

1921. Hoplopleura erratica erratica (Osborn), Ferris, Contributions Toward
a Monograph of the Sucking Lice, Part 2:106; figures 67, 68.

HOSTS AND DISTRIBUTION. Type recorded as taken from a gull in Iowa, but
this very evidently in error. The host is clearly Tamias striatus, from
which the species has been recorded by Ferris from Indiana, Tennessee, New
York, and the District of Columbia; all in the United States.

NOTES. The supposed subspecies recognized by Ferris are here considered
to be species.

Hoplopleura fonsecai Werneck

1934. Hoplopleura fonsecai Werneck, Memorias do Instituto Oswaldo Cruz 27:
412; figures 7-12.

HOSTS AND DISTRIBUTION. Type from Oxymycterus judex at Humboldt, state
of Santa Catharina, Brasil.

Hoplopleura hesperomydis (Osborn)

1891- Haematopinus hesperomydis Osborn, United States Department of Agri¬
culture, Division of Entomology, Bulletin (old series) 7:26; fig¬
ure 14.

1915. Hoplopleura hesperomydis (Osborn), Kellogg and Ferris, Anoplura and
Mallophaga of North American Mammals, Stanford University Publica¬
tions, University Series (no volume number), page 17; text figures
4, 5; Plate 1, figure 3; Plate 4, figure 2. (The name Hoplopleura
hesperomydis occ idental is, appearing in the legend of Plate 4,
figure 2, is due to an error.)

1921. Hoplopleura hesperomydis (Osborn), Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 2:70; figures 38, 39.

HOSTS AND DISTRIBUTION. Originally described as from Peromyscus (= Hes -
peromys) leucopus at Ames, Iowa. Recorded by Kellogg and Ferris and by
Ferris from Peromyscus boylei and Peromyscus maniculatus in California.
Recorded by Ferris from Onychomys torridus in California and Onychomys
leucof aster in Colorado. All these localities are in the United States.
Recorded by Ferris from Oryzomys fulvescens at Orizaba, Mexico; from Oryzo-
mys chaparensis at Santos, Bolivia; and from Eli£modontia collisae (which
Hopkins has determined to be Hesperomys callosus) at Goya, Argentina.

136

Ferris has also recorded it from Hus musculus in Virginia, U . S. A., and
Russian Turkestan; from Hus tanaus, which is apparently a synonym of Hus
nusculus, from China; from Hus uagneri, which is also a synonym ol Hus
nusculus, from China.

NOTES. The material recorded above has been examined in connection with
this work and still appears to be Hoplopleura hesperomydis. The normal
hosts of this species seem quite definitely to be species of Peromyscus and
the closely related genus Onychomys. Some of the other records may be due
to contamination. The occurrence of the species on Hus nusculus in the
United States may possibly be due to an actual transfer 1 rom the normal
hosts, but its occurrence on this host in Asia is very strange.

Certain specimens from hosts of the genus Re ithrodontomys, which have
previously been referred to hesperomydis, are here transferred to a new
species, Hoplopleura re tthrodontomydis.

Hoplopleura hirsuta Ferris

1916. Hoplopleura hirsuta Ferris, Psyche 23:112; figures 8, 9A, 10, 11U.
1921. Hoplopleura hirsuta Ferris, Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 2:117; figures 75> 76.

HOSTS AND DISTRIBUTION. Type from Sigmodon hispidus, Raleigh, North
Carolina. There are various records from this host throughout southern
United States, as far west as Yuma, Arizona. Ferris has recorded it from
Sl^nodon ocrognathus from the state of Chihuahua, and from Sigmodon peru-
anus from Peru. Two records by Ferris, each ol a single specimen, 1 rom
skins of Rhipldomys venustus in Venezuela and Xenomys nelsoni in Mexico,
are most probably due to contamination.

Hoplopleura hispida (Grube)

1851. Pedlculus hispidus Grube, In Middendorf f ' s Reise, Parasiten, page
497; Plate 2, figure 2. .

1921. Hoplopleura hispida (Grube), Ferris, Contributions Toward a Monograph

of the Sucking Lice, Part 2:67- . „

HOSTS AND DISTRIBUTION. Recorded as from Lemnus obensis "am Taimyrsee,

in Siberia. . . , .

NOTES. On the basis of the illustration accompanying the original de¬
scription this species seems to be a Hoplopleura and should be identi liable
if taken from the type host.

1915.

1921.

Hoplopleura intermedia Kellogg and Ferris

Hoplopleura Intermedia Kellogg and Ferris, Annals ol the Durban Mus¬
eum 1:153; Plate 16, figures 5, 5a-d. ......

Hoplopleura intermedia Kellogg and Ferris, Ferris, Contributions
Tb ward a Monograph of the Sucking Lice, Part 2:90; tigures S4, _.B*

from Rattus coucha, Mfongosi, Zululand,
Rattus tullberii and Rattus (as Zelotomys)
and from Dendromus insignis from British

C, 56B.

HOSTS AND DISTRIBUTION. Type
South Africa. Also recorded from
hildeiardae, British East Africa

II l i Lit; 5li» tiUC f LJ * A VAOU ** , • i •

East Africa, this last record probably due to contamination

Hoplopleura laticeps Ferris

1921. Hoplopleura laticeps Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 2:92; figures 55A, 56A.

HOSTS AND DISTRIBUTION. Known only from the original record irom Hybomys
(as Arvicanthis) unlvittatus from Benito River, West Africa.

157

Hoplopleura longula (Neumann)

1909. Haematopinus (Polyplax) longula Neumann, Archives de Parasitologie

13:513; figures 15, 17-

1910. Hoplopleura lineata Fahrenholz, Zoologischer Anzeiger 35:715-

1921. Hoplopleura longula (Neumann), Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 2:68; figures 36, 37.

HOSTS AND DISTRIBUTION. Described by Neumann from Micromys minutus at
Colchester, Essex, England, and by Fahrenholz from the same host, presumab¬
ly from Germany.

Hoplopleura maniculata (Neumann)

1909. Haematopinus (Polyplax ) maniculatus Neumann, Archives de Parasito¬
logie 13 : 521 ; figures 21, 22.

1921. Hoplopleura maniculata (Neumann), Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 2:112; figures 71, 72A, D, G.
HOSTS AND DISTRIBUTION. Described from Funambulus (as Sciurus) palmarum,
Rajkote, Rajkote, India, and at the same time erroneously attributed to a
bat. Later recorded by Ferris from the type host from Navapour, India, and
from Ceylon.

Hoplopleura malaysiana Ferris

1921. Hoplopleura malaysiana Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 2:79; figures 44, 45.

HOSTS AND DISTRIBUTION. Recorded as from Rattus vociferans lancauensis,
which is apparently Rattus sabanusj from Lankavi Island, Malay Straits.

Hoplopleura merionidis Ferris

1921. Hoplopleura merionidis Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 2:98; figure 60.

HOSTS AND DISTRIBUTION. Recorded from Meriones psammophilus from Shansi,
China, this apparently being identical with Meriones meridianus.

Hoplopleura mylomydis Ferris

1921. Hoplopleura enormis mylomydis Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 2:97; figure 59C.

HOSTS AND DISTRIBUTION. Known only from the original description from
Mylomys roosevelti, which is apparently a synonym of Hylonys cuninghamei ,
British East Africa.

Hoplopleura nesoryzomydis Ferris

1921. Hoplopleura nesoryzomydis Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 2:90; figure 53A.

HOSTS AND DISTRIBUTION. Recorded as from Hesoryzomys narboroughi and
Mesoryzomys defessus in the Galapagos Islands. According to Ellender,
Nesoryzomys is merely a subgenus of Oryzomys and evidently defessus is an
error for indefessus.

Hoplopleura neumanni Fahrenholz

1901.

1902.

Haematopinus praecitus Neumann, Archives
(In part: error for praectsus)
Haematopinus praeclsus Neumann, Archives de
part)

138

de Parasitologie 5:600.
Parasitologie 6:144. (In

1919- Boplopleura neumannt Pahreuholz, Jahresbericht des niedersachsischen
Zoologischeu Vereins zu Hanuover 2-4 : 26.

1921. Hoplopleura neunuinnt Pahreuholz, Perris, Contributions Toward a Mon¬
ograph of the Sucking Lice, Part 2:101; figure 63.

HOSTS AND DISTRIBUTION . Based upon specimens recorded merely as "gros
rats," from Abyssinia. Perris lias recorded the species lrom Tatera ni£ri-
cauci a from British past Africa. Hopkins has noted that this species occurs
iu Abyssinia and it is probable that it was the type host.

HopLopleura ochotonae Ferris

1922. Hoplopleura ochotonae Perris, Contributions Toward a Monograph ot
the Sucking Lice, Part 3:142; figure 92.

HOSTS AND DISTRIBUTION. Described as from Ochotona cansus, which is a
synonym of Ochotona thibetana, from Taochao, China. Recorded also from
Ochotona danur ica, Tabool, Mongolia, and trom Ochotona roylei without iudi
cation of locality.

Hoplopleura oenomydis Ferris
Figures 59, 60

1921. Hoplopleura oenomydis Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 2:82; figures 47, 48.

1924. Hoplopleura pac if tea Ewing, Bishop Museum Bulletin 14:9; figure lb, e .
1932. Hoplopleura oenomydis Ferris, Ferris, Bishop Museum Bulletin 98:121;
figures 38a_k, 39-

1947. Hoplopleura oenomydis Ferris, Pritchard, The Journal of Parasitology
35:374.

HOSTS AND DISTRIBUTION. Type from Oenomys hypoxanthus at Molo, British
East Africa. Also recorded from Dasymys incomtus and Grammomys (as Tham-
nomys) surdaster from British East Africa; from Rattus exulans (as calc l s)
and Rattus mearnsl from the Philippine Islands. Described by Ewing as Hop¬
lopleura pacifica from Rattus exulans (as hawaiiensis) trom the Hawaiian
Islands and recorded from this same host from the Marquesas Islands. It
has recently been shown to be the most common louse on Rattus norveiicus in

southeastern United States. .

NOTES. Hopkins has recently expressed doubt concerning the synonymy in
dicated above, considering it to be "extremely improbable that all the
published records can refer to Hoplopleura oenomydis. _ A re-exam mat ion ol
all the available material reveals no reason for altering the opinions pre
viously expressed by Ferris in regard to the matter, whatever the iraproba

bilities" may be.

Hoplopleura oryzomydis Pratt and Lane

1951. Hoplopleura oryzomydis Pratt and Lane, Journal ol Parasitology 3< •

HOSTS AND’ DISTRIBUTION. Type from Oryzomys palustris, Oatland Island,
Chatham County, Georgia, and other specimens from the same host species in
Delaware, South Carolina, and Florida, United States.

Hoplopleura oxymycteri Ferris

1921 Hoplopleura oxymycteri Ferris, Contributions Toward a Monograph ol
the Sucking Lice, Part 2:122; figures 79, 80.

HOSTS .AND DISTRIBUTION. From Oxymycterus par omens is at Occabamba Pass,

Peru.

139

Hoplopleura oenomydis Ferris

Figure 59

140

v fed

para tergal plates

thoracic sternal plate

female genitalia
Hoplopleura oenomydis Ferris, details

male U genitalia

Figure 60

141

Hoplopleura pectinata Cummings

1913* Hoplopleura pectinata Cummings, Bulletin of Entomological Research

4:35-

1921. Hoplopleura pectinata Cummings, Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice. Part 2:99; figures 61, 62.

1929. Ctenura pectinata (Cummings), Ewing, A Manual of External Parasites,
page 199.

1932. Hoplopleura pectinata Cummings, Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 5:282.

HOSTS AND DISTRIBUTION. Described from Rattus (as Epimys) surifer at
Biserat, Jalor, Malay Peninsula, and later recorded from the same host from
Trong, lower Siam.

NOTES. This species has been designated as type of the genus Ctenura, a
genus which is here rejected.

Hoplopleura pelomydis Ferris

1921. Hoplopleura enormis pelomydis Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 2:96; figures 58A, 59 A.

HOSTS AND DISTRIBUTION. Described as from Pelomys fallax,* Summit Sagalla,
British East Africa. Also recorded from Lemniscomys striatus (in part as
pulchellus) from the Cameroons and from British East Africa. It is probable
that the species of Lemniscomys are th£ true hosts.

Hoplopleura phaiomydis Ferris

1921. Hoplopleura phaiomydis Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 2:120; figures 77,

HOSTS AND DISTRIBUTION. Recorded as from an undetermined species of
Phaiomys from East Ladak, Kashmir. Hopkins has ascertained that the host
species was Phaiomys blythi, which Ellender considers to be a synonym of
Phaiomys leucurus.

Hoplopleura quadridentata (Neumann)

1909. Haematopinus ( Polyplax ) quadridentatus (Neumann), Archives de Para^-
sitologie 13:5:3-15; figures 13-14.

1921. Hoplopleura quadridentata (Neumann), Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 2:87; figures 52, 53B, C, E.

HOSTS AND DISTRIBUTION. Recorded by Neumann from Holochilus squamipes,
which apparently is Nectomys squamipes, from Haut Peru. Later recorded by
Ferris from this same host at Sapucay , Paraguay, and from Nectomys palmipes,
which is perhaps the same as squamipes, from the island of Trinidad; from
Orizomys fulvescens from Orizaba, Mexico; and from Oryzomys rostratus from
Alta Mira, Tamaulipas, Mexico.

hoplopleura reducta Ferris

1921. Hoplopleura reducta Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 2:124; figure 81.

1935- Hoplopleura reducta Ferris, Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 8:615.

HOSTS AND Dl STRI 11! TION . Described as from phyllotis micropus from Todos
Santos, Guatemala. Later recorded from an undetermined host from unspeci¬
fied locality in South Anerica.

142

Hoplopleura reithrodontomydis Ferris, new species

HOSTS AND DISTRIUJTION . Type from a female Reithrodontonys dorsalis,
Todos Santos, Guatemala, U.S.N.M. skin number 76917. ParaLypes from Heith-
rodontomys australis, Volcan de Irazu, Costa Rica, U.S.N.M. skin number
116623, and Reithrodontomys chrysopsis, Ajusco, near Mexico City, Mexico.

CHARACTERS. Female in all respects identical with Hoplopleura hesper
omydis, except for the shape of the paratergites of abdominal segment seven,
which in hesperomydis have both dorsal and ventral lobes definitely acute
apically, while in reithrodontomydis the dorsal lobe is broad and apicaily
truncate or slightly emarginate. All of the rather numerous specimens at
hand from the three host species and three localities agree very closely.
An extended description and illustrations of the species will be presented
elsewhere.

Hoplopleura rukenyae Ferris

1921. Hoplopleura sukenyae Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 2:86; figure 51* (Misspelling)

HOSTS AND DISTRIBUTION. Known from a single male from Hus triton, Mount
Rukenya, British East Africa.

NOTES. Mr. G. h. E. Hopkins has called attention to the fact that the
type locality of this species is more commonly, and perhaps more correctly,
spelled Rukenya rather than Sukenya. It is probable that an error in read¬
ing a label was committed and under these circumstances it seems justifi¬
able to change the spelling of the specific name to rukenyae.

Hoplopleura sciuricola Ferris

1921. Hoplopleura sciuricola Ferris, Contributions Toward a Monograph of

the Sucking Lice, Part 2:110; figures 69, 70.

HOSTS AND DISTRIBUTION. Type from Sciurus carollnensis at Bayou St.
Louis, Mississippi. Also recorded from other species of Sciurus as follows:
hudsonicus iu Alaska, douglasi in California, ignitus in Peru, nesaeus in
Venezuela, variabllts in Colombia, and undetermined species in Bolivia and
Peru.

Hoplopleura somereni Waterston

1923. Hoplopleura somereni Waterston, Bulletin of Entomological Research
14:99; figure lb, c, d; figure 2c, d.

HOSTS AND DISTRIBUTION. From Dasymys helukus atWamia, Okedi Camp, Kenya.

Hoplopleura travassosi Werneck

1932. Hoplopleura travassosi Werneck, Revista Medico-Cirurgica do Brasil,
anno 40:345; figure.

1934. Hoplopleura travassosi Werneck, Memorias do Instituto Oswalno uruz

27:409; figures 1-6. D .

HOSTS AND DISTRIBUTION, type from Oryzomys flavescens at Angra dos Keis,
state of Rio de Janeiro, Brasil. Also recorded from Kannabateomys amblyonyx
and Oxymycterus judex at the same locality.

Hoplopleura trispinosa Kellogg and Ferris

191^. Hoplopleura trispinosa Kellogg and Ferris, Anoplura and Mailophaga
of North American Mammals, Stanford University Publications, Uni¬
versity Series (no volume number), page 22; text figure 8; Plate
4. figure 3. ^

1921. Hoplopleura trispinosa Kellogg and Ferris, Ferris, Contributions
Toward a Monograph of the Sucking Lice, Part 2:115; figures 73> 74.
1929. Euhoplopleura trispinosa (Kellogg and Ferris), Ewing, A Manual of
External Parasites, pages 135> 199*

HOSTS AND DISTRIBUTION. T>pe from Glaucomys sabrinus, Brownsville, Ore¬
gon. Also recorded from the same host at Yosemite National Park, Califor¬
nia, and from Glaucomys volans from Maryland.

NOTES. This species has been designated as type of the genus Euhoplo¬
pleura, which is here rejected.

Hoplopleura veprecula Ferris

1921. Hoplopleura veprecula Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 2:105; figures 64B, 66A, C, F.

HOSTS AND DISTRIBUTION. From Tatera boehmi at South Guaso Nyiro, Brit¬
ish East Africa.

Genus PTEROPHTHIRUS Ewing

1923* Pterophthirus Ewing, Journal of the Washington Academy of Sciences
13:147.

1932. Pterophthirus, Ferris, Contributions Toward a Monograph of the Suck¬
ing Lice, Part 5:280.

GENERIC TYPE. Hoplopleura alata Ferris. Three other species are here
referred to the genus.

CHARACTERS. Hoplopleurinae with f ive-segmenced antennae. Paratergal
plates of abdominal segment two produced into a long, tapering, blade-like
process which projects from the body. First sternal plate of abdominal
segment three not produced laterally to articulate with the corresponding
paratergal plates. Otherwise essentially as in Hoplopleura.

NOTES. This genus is very' close to Hoplopleura, differing from the lat¬
ter only in the form of the paratergal plates of abdominal segment two.
Even here it appears from the illustrations presented in connection with
the description of Pterophthirus imitans Werneck that this species is some¬
what of an intermediate between the two genera. As known at present the
genus is confined to South American rodents.

The four known species may be separated by the following key.

Key to Species of PTEROPHTHIRUS

1. Paratergal plates of abdominal segments 3-4 each with the ventral, api¬

cal angle produced into an acute point . 2

Paratergal plates of abdominal segments 3~4 with the ventral apical an¬
gle not at all produced . ’ . 3

2. Paratergal plates of segments 3-5 with both dorsal and ventral, apical

angles produced into an acute point . IMITANS

Paratergal plates of segments 3^5 with the ventral, apical angle pro¬
duced into an acute point, the dorsal angle broadly truncate .WERNECK1

3. Paratergal plates of segments 3~4 with the dorsal, apical angle pro¬

duced into a point . ALATA

Paratergal plates of segments 3-4 with the dorsal, apical lobe broadly
truncate . AUDAX

Pterophthirus alata (Ferris)

Figures 61, 62

1921. Hoplopleura alata Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 2:127; figures 84, 85-

144

Pterophthirus alata (Ferris)

Figure 61

145

146

1923- Pterophthirus alata (Ferris), Ewing, Journal of the Washington Acad
emy of Sciences 13:147.

1942. Pterophthirus alata (Ferris), Werneck, Kevista Brasil iera di Bio-
logia 2(3): 317.

HOSTS AND DISTRIBUTION. Recorded by Ferris l'rom Hicrocavia (as Kero Ion)
australis from the Upper Rio Chico, Patagonia, Argentina, and later record¬
ed by Werneck from the same host (as Cauiella) from the provinces ol Jujuy
and Catamarca, in Argentina.

Pterophthirus andax (Ferris)

1921. Hoplopleura audax Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 2:125; figures 82, 83.

I923. Pterophthirus audax (Ferris) , Ewing, Journal of the Washington Acad¬
emy of Sciences 13:148.

1923. Pterophthirus audax (Ferris), Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice. Pttrt 5 - 281 * <

1942. Pterophthirus audax (Ferris) , Werneck, Revista Brasiliera di Biolog-

ia 2(3):317.

HOSTS AND DISTRIBUTION. Types from Proechimys semispinosus and other
specimens from Proechimys (as Nelomys) mincae at San Javier, North Ecuador.
Recorded by Werneck from Proechimys oris at Abiete, Para, Brasil.

Pterophthirus imitans Werneck

1942. Pterophthirus imitans Werueck, Revista Brasiliera di Biologia 2(3):

317; figures. ,

HOSTS AND DISTRIBUTION. From Cavia aperea, Santo Amaro, state ol Sao

Paulo, Brasil.

Pterophthirus wernecki GuimarSLes

1950. Pterophthirus wernecki GuimarS.es, Papeis Avulsos do Departamento de
Zoologia, Secretaria da . Agricultura, SSo Paulo, Brasil 9:8:83;

figures. . f

HOSTS AND DISTRIBUTION. From Proechimys iherinti at Boraceia, state 01

SSo Paulo, Brasil.

Genus SCHIZOPHTHIRUS Ferris

1922. S chizophthirus Ferris, Contributions Toward a Monograph of the Suck
ing Lice, Part 3:143.

1932. Hasellus Jancke, Zeitschrift tiir Paras itenkunde 4: 33^. .

GENERIC TYPE. Pediculus pleurophaeus Burmeister, by original designa¬
tion. One other species is included in the genus.

GENERIC SYNONY’M. Hasellus Jancke, by community of type.

CHARACTERS. Hoplopleurinae with five-segmented antennae. Abdominal
segments , exclusive of the usual terminal and genital segments, without
tergal or sternal plates in the female except for plates belonging appar¬
ently to segments one to three. Male with such plates on all segments.
Female with three rows of setae on most of the segments both dorsalh .a <
ventrallv, the male with one row on each segment both dor sally and ventral^
lv. Sternal plate of segment two divided longitudinally into two much ex
nanded Dlates, each of which articulates by means of a process with the
corresponding paratergal plate and each of which bears on its posterior

border 2-3 stout, thorn-like setae. f -i aiiridae

The members of this genus occur on rodents ot the lamily Giindae.

147

Key to Species of SCHIZOPHTHIRUS

Dorsal and ventral lobes of the paratergal plates of abdominal segments 3~6
deeply divided into 2 very unequal lobes, the dorsal lobe being much nar¬
rower than the ventral lobe; known from the European genera Muscardinus

and Eliomys... . PLEUROPHAEUS

Paratergal plates of these abdominal segments not thus deeply divided, and
with the lobes equal; known from the genus Graphiurus in Africa. GRAPHIURI

Schizophthirus graphiuri Ferris

1922. Schizophthirus graphiuri Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 3:147; figures 93A, 96, 97.

HOSTS AND DISTRIBUTION. Type from Graphiurus murinus from British East
Africa and recorded also as from Graphiurus murinus (as raptor) from the
same region; from Graphiurus nanus from Natal, South Africa; from Graphiurus
alticola, locality not specified in available reference.

Schizophthirus pleurophaeus (Burmeister)

Figures 63, 64

1839- Pediculus pleurophaeus Burmeister, Genera Insectorum, Rhynchota,
Number 7.

1922. Schizophthirus pleurophaeus (Burmeister), Ferris, Contributions
Toward a Monograph of the Sucking Lice, Part 3:145; figures 94, 95-

HOSTS AND DISTRIBUTION. Type from Dryomys nitedula (as Myoxys nitella)
from somewhere in Europe. Also recorded from Eliomys quercinus (as pall idus)
from Italy and from Muscardinus avellanar ius from Germany.

Subfamily HYBOPHTHIRINAE Ferris, new subfamily

DESCRIPTION OF THE SUBFAMILY. Hoplopleuridae in which there is no ex¬
ternal indication of eyes. The most distinctive character is the presence
of a short, claw-like structure which arises beside the true claw on the
anterior legs. Paratergal plates present on abdominal segments 2-8 or 3-8,
with at least one of the posterior apical angles forming a lobe or point
which is free from the body wall. Abdomen with but a single row of setae
on any segment, both dorsally and ventrally, in either sex and without
sclerotized tergal and sternal plates other than those normally present on
the terminal and genitalic segments, except at times in the male. Antennae
five-segmented, sexually dimorphic only to the extent that the male may
bear a single enlarged seta near its apex.

Occurring on African mammals of the rodent family Echimyidae, subfamilies
Thryonomyinae and Petromyinae, and the family Orycteropodidae of the Order
Tubulidentata.

NOTES. A considerable amount of doubt is felt concerning the validity
of this subfamily. The presence of the peculiar claw-like structure along¬
side the true claw on the front tarsi seems to link the included species and
there are no other characters which specifically deny such an association.

Webb referred the two included genera to the Haematopinidae, but the
present writer is quite unable to agree with this assignment.

Key to the Genera of HYBOPHTHIRINAE

Hindhead almost triangular, the lateral margins strongly convergent; occur¬

ring on the genus Orycteropus of the Order Tubulidentata . HYBOPHTHIRUS

Hindhead with the lateral margins approximately parallel; occurring on rod¬
ents of the family Echiiqyidae . SCIPIO

148

Schizophthirus pleurophaeus (Burmeister)

Figure 63

149

2nd stcrnite

3rd claw

paratergal pl<de

nude genitalia

Schizophthirus pleurophaeus (Burmeister), details

Figure 64

Genus HYBOPHTHIRUS Enderlein

1909. Hybophthirus Enderlein, Kenkschrift der medicinishen-naturwissen
schaft-ichen Gesellschaft zu Jena 14:79*

1922. Hybophthirus , Ferris, Contributions Toward a Monograph of the Suck¬
ing Lice, Fart 3-175*

GENERIC TYPE* Hybophthirus orycteropodi Enderleiu, which is a synonym
of the earlier described Haematopinus notophallus Neumann.

CHARACTERS* llybophthirinae in which the head is short and broad, sharp¬
ly expanded posterior to the antennae and then constricting sharply, the
hindhead being almost triangular. Thorax without any sternal plate; pro-
thoracic sternal apophyses present, forming a pair of pits on the ventral
side; with the metanotum forming a distinct, apically tree lobe at each
posterior angle. First pair of legs slender, with slender claw; second and
third legs equal to each other, large and stout, with stout claw. Abdomen
with paratergal plates present on segments 2-8, each with the posterior,
dorsal angle prolonged into an apicaily rounded, free lobe. Abdomen with
but one row of small setae across each segment, both dorsally and veutrally,
in both sexes.

Hybophthirus notophallus (Neumann)
Figures 65, 66

1909. Haematopinus notophallus Neumann, Jahresbericht des Nassauische Ver-
eins fur Naturkunde in Wiesbaden, page 2.

1909. Hybophthirus notophallus Enderlein, Denkschrift der medicinischen-
naturwissenschaft-ichen Gesellschaft zu Jena 14:79-80; Plate 8,
figures 1-3*

1922. Hybophthirus notophallus (Neumann), Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 3:176; figures 117-118.

HOSTS AN ^DISTRIBUTION. Neumann's types were from Orycteropus afer
{■=-capensls) , the "Cape ant bear, "at "Gochas, Afrique occidentale allemande,"
and Enderlein's types were from the same host in "Klein— Namaland, Umgebung
von Steinkopf." The species has later been recorded from the same host in
South Africa.

NOTES* The priority of Neumann's specific name over that applied by
Enderlein has been pointed out by Cummings. The accompanying illustrations
are from specimens from the type host in the Zoological Garden at Pretoria,
South Africa.

Genus SCIPIO Cummings

1913* Sclpio Cummings, Bulletin of Entomological Research 3:393-

1916. Neumannel lus Fahrenholz, Archiv fur Naturgeschichte , Abteilung A,

81:11:31*

1922. Scipio, Ferris, Contributions Toward a Monograph of the Sucking

Lice, Part 3:170. ^ n

1936* Bed ford i a Fahrenholz, Zeitschrilt iiir Parasitenkunde 9. bp.

"GENERIC TYPE. Haematopinus aulacodi Neumann, the only included species
at the time of the naming of the genus. Heumannellus Fahrenholz has the
same type. Bedfordia Fahrenholz, type Scipio tripedatus Ferns, is here

considered to be a synonym. ... , ,

CHARACTERS. Head with the posterior lateral margins more or less nearly

parallel. Antennae of the male with the third segment somewhat modified by
the presence of a stout, subapical, dorsal seta. Thorax at the most with a
verv small sternal plate which is not apically or marginally tree. Poste
rior- lateral angle of the metathorax dorsally with a pronounced lobe, ex
cept that this is weakly developed in one species. Abdomen with definite

151

paratergal plates on segments 3-8* Female in all the species with the ab¬
domen membranous except for the usual ninth tergite and the genital plate;
male in two species with a single, very small, tergal plate on each of most
of the segments. Anterior legs small and with slender claw with a short,
claw-like process arising beside it. Middle and posterior legs enlarged
and with stout claw or middle legs larger and stouter than the posterior
pair.

NOTES. Four species, one probably invalid, are here referred to this
genus. Of these, one — Scipio tripedatus Ferris — is a rather peculiar form,
having the middle legs enlarged and with stout claw, hut the posterior legs
definitely smaller and with slender claw. Fahrenholz has considered this
sufficient to justify the erection of a new genus, Bedfordia, but this is
not accepted here, as the relationships of the species seem very definitely
to he with Scipio. All the species occur upon hosts of the rodent family
Echimy idae.

Key to Species of SCIPIO

One supposed species, longiceps Ewing, is omitted from this key.

1. Middle legs larger than posterior legs and with stouter claw. TRIPEDATUS

Middle and posterior legs of approximately equal size . 2

2. Head very noticeably elongate . AULACODI

Head but slightly longer than broad . BREVICEPS

152

male fjenitalia

Hybophthirus notophallus (Neumann), details

female genitalia

Figure 66

tst claw

antenna

Scipio aulacodi (Neumann)

Figures 67, 68

1911. Haematopinus aulacodi Neumann, Archives de Parasitologie 14:403-
fi -.mres 5-7.

1913. Scipio aulacodi (Neumann), Cummings, Bulletin ol Entomological Re¬
search 3:393- _ , M .

1922 Scipio aulacodi (Neumann), Ferris, Contributions Toward aMonogiap
of the Sucking Lice, Part 3:170; figures 113-114-
HOSTS AND DISTRIBUTION. Originally described from Thryonomys (- Aulacodus )
suinderianus from Dahomey, Africa. Later recorded from Thryonomys sp-,
Mfon.ros, Zululand, and from Thryonomys suinderianus ( aulacodus) vaneta.J.
from= Rust enburg, Transvaal District, South Africa. The hosts are mem! pis
of the rodent family Echimyidae.

153

Scipio aulacodi (Neumann)

Figure 67

Scipio breviceps Ferris

1922. Scipio breviceps Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 3:173> figures 114, 115, 116.

HOSTS AND DISTRIBUTION. Originally described from Thryonomys sp., Zulu-
land, and later recorded by Bedford from Thryonomys swinderianus variegatus.
The hosts are members of the rodent family Echimyidae.

Scipio longiceps Ewing

1937- Scipio longiceps Ewing, Proceedings of the Helminthological Society
of Washington, page 81; figure 29.

HOSTS AND DISTRIBUTION. From Thryonomys gregor pus Ulus from Majiya-
cgumvi , British East Africa. The host is a member of the rodent family
Echimyidae.

NOTES. The description and very inadequate accompanying illustration,

154

female genitalia male genitalia

Scipio aulacodi (Neumann), details Figure 68

166

both based upon the male alone, offer no convincing evidence that this
species is distinct from Scipio aulacodi.

Scipio tripedatus Ferris

1932. Scipio tripedatus Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 5:285; figures 173-175*

1936* Bedfordia tripedata (Ferris), Fahrenholz, Zeitschrift fur Parasiten-
kunde 9:55*

HOSTS AND DISTRIBUTION* Type from a "rock rat," one of the species of
the genus Petromus {-Petromys ) without locality other than South Africa and
also recorded from Petromus typicus tropicalis , Windhoek, South Africa, and
from Petromus sp . at Khan River, Southwest Africa. The hosts are members
of the rodent family Echimyidae.

NOTES. This species has been designated as type of the genus Bedfordia
Fahrenholz, but this genus is here rejected.

Subfamily PEDICININAE Enderlein

1904. Enderlein, Zoologischer Anzeiger 28:136, 138.

DESCRIPTION OF THE SUBFAMILY. Hoplopleuridae in which distinct eyes are
present as external lenses accompanied in life by pigment spots. Antennae
five-segmented, the last three segments at times more or less fused togeth¬
er; sexually dimorphic, the males having a small, stout seta on the dorsal
side of each of the last three segments. Paratergal plates present on ab¬
dominal segments 4-6 or 5-6 in the form of distinct plates of which at
least one of the apical angles is free from the body. Abdomen otherwise
membranous except for the usual terminal and genitalic plates. Spiracles
present on abdominal segments 3-8. Legs variable in form, either all more
or less similar and relatively slender or the first pair slender and the
others larger and stouter. Gonopods of segment eight of the female always
obsolete, their position represented only by a row of setae. Gonopods of
segment nine likewise represented merely by a row of setae, the apex of the
body never with lobes or processes. Genitalia of the male always with par-
ameres which are fused basally to the base of the aedeagus.

Occurring on Old World monkeys of the superfamily Cercopithecoidea.

NOTES. The removal of the genus Pedicinus from the family Pediculidae,
with which it has long been associated, has been decided upon only after
much doubt. The morphological community with the Pediculidae has lain only
in the presence of distinct eyes, but it is now known that the eyes are
present in various forms and they presumably mean nothing more than the re¬
tention of a primitive character which was once common to all the lice. In
the remainder of its morphology Pedicinus departs as widely from Pediculus
as do most of the other genera. Even the spiracles, which have been in¬
sisted upon by Webb as indices to relationship, have not been claimed by
him to have any special resemblance to those of the Pediculidae. In other
respects the members of the genus approach quite closely various forms of
the Hoplopleuridae and, considering the morphological evidence, there have
been but two alternatives apparent. One of these is to mime a new family
for the genus Pedicinus . The other is to regard this genus as constituting
a subfamily of the Hoplopleuridae. The second alternative has been chosen.

The removal of Pedicinus from the Pediculidae and its assignment to the
Hoplopleuridae will doubtless be viewed with horror by those whose ideas of
the relationship of the various genera of lice are at least colored — if not
determined — by the relationships of the hosts. The genus Pedicinus, being
from a Primate, ought to be related to Pediculus, but the morphological ev¬
idence does not support such a relationship. Possibly future workers will
see some other solution of the difficulty.

156

Genus PEDICINUS Gervais

1844. Pedlclnus Gervais, In Walckenaer's Histoire naturelle des insectes
up teres 3 * 30 1 -

1912. Phthirpedicinus Fahrenholz, Zoologischer Anzeiger 39:64.

1916. Neopedictrius Fahrenholz, Archiv ftir Naturgeschichte, Abteilung A,

81:11:7.

1934. Pedicinus , Ferris, Contributions Toward a Monograph of the Sucking
Lice, Fart 7 :5U2.

GENERIC TYPE. The question of just what name the type of this genus
shouLd bear is open to a difference of opinion, the genus having been based
upon a niisidenti f ied species. The genus was based by Gervais upon speci¬
mens which he identified as being the Pediculus euryiaster of Burmeister,
but which on the basis of his description and illustrations was clearly
misideutified. The opinion is here held that the generic name belongs with
the species which he actually had before him and upon which he based his
generic concept, not with the species which belongs with the name that he
mistakenly employed. The questiou then remains as to what species Gervais
actually had.

Ferris has maintained that in all probability it was the species later
described by Piaget as Pedicinus loniiceps. Hopkins, however, has main¬
tained the opinion that longiceps should be regarded as a synonym of the
earlier name Haematopinus obtusus Rudow. This opinion is here reluctantly
accepted and the type of the genus Pedicinus thus may be given as Haemato¬
pinus obtusus Rudow.

GENERIC SYNONYMS. Neopedicinus Fahrenholz, type Heopediclnus patas Fah¬
renholz; Phthirpedicinus Fahrenholz, type Phthirpedicinus micropilosus Fah¬
renholz, which is here considered to be a synonym of Pedicinus euryiaster
(Burmeister) .

CHARACTERS. With the same characters as the subfamily, of which it is
the only included genus.

NOTES. In the opinion of the writer there is no justification for the
two genera named by Fahrenholz, although the names are available should
future workers desire to employ them.

Key to Species of PEDICINUS

With but 2 pairs of paratergal plates on the abdomen . EURYGASTER

With 3 pairs of free paratergal plates on the abdomen . 2

All legs of essentially the same size and form . . . 3

Secondhand third pairs of legs definitely stouter and with heavier claw

than the first pair . * . . .

|p,rc verv Ion ^ and slender; penis of the male apically acutelj pointed.
“ . HAMADRYAS

Le'rs not thus long and slender . . . . . ;4

Female with the genital plate trapezoidal and with the posterior margin
deeply emarginate ; male with the penis apically flattened and pro¬
duced into two slight points . . . .ALBIDUS

Female with the genital plate transversely narrow; male with the penis

merely slightly swollen at the apex . OBTUSUS

With small paratergal sclerotization on segments t-S, in addition to
the 3 pairs of paratergal plates in both male and female. ..... .PICTI 5

Without such sclerotizations in addition to the three pairs ol parater¬
gal plates . . .

Penis of the male with a tooth on each side just anterior to the apex..

. . ANCORATUS

Penis of the male without such preapical teeth . PATAS

Pedicinus albidus (Rudow)

1869- Haematopinus albidus Rudow, Zeitschrift fur die gesamten Naturwis-
senschaften 34:168.

1934. Pedicinus albidus (Rudow), Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 7:511; figure 296.

HOSTS AND DISTRIBUTION. Type from the "barbary ape," Macaca s’jlvanus.
Ferris has recorded the species from the same host from Morocco and in the
London Zoological Garden.

Pedicinus ancoratus Ferris

1934. Pedicinus ancoratus Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 7:518; figures 299, 300A, F, H, I.

HOSTS AND DISTRIBUTION. Type from Presbytis pullata, Pulo Sebang, East
Sumatra. Also recorded from Presbytis cristata and Presbytis germaini from
the Malayan region, from Presbytis schistacea from Kashmir, from Presbytis
rubicunda from Borneo, doubtfully from Pygathrix priamus from Ceylon.

Pedicinus eurygaster (Burmeister)

1838. Pediculus eurygaster Burmeister, Genera Insectorum, Rhynchota, Spe¬
cies 21.

1864. Pediculus microps Nitzsch, Giebel, Zeitschrift fiir die gesamten Na-
turwissenschaften 23:32.

1880. Pedicinus eurygaster (Burmeister), Piaget, Les Pediculines, page 630.
(In part)

1880. Pedicinus longiceps Piaget, Les Pediculines, page 632. (In part)

1880. Pedicinus breviceps Piaget, Les Pediculines, page 632. (In part)

1881. Pedicinus piageti Stroebelt, -Jahresbericht der zoologischen Sektion

des Westfalischen provincial-Vereins fiir Wissenschaft und Kunst
9:82; Plate 1, figure 3*

1912. Phthirpedicinus micropilosus Fahrenholz, Zoologischer Anzeiger 39 : 55.
1932. Phthirpedicinus microps (Nitzsch) , Werneck, Annales da Academia
Brasiliera de Sciencias 4:163; figures 1-5-
1934. Pedicinus eurygaster (Burmeister), Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 7:521; figures 303— 30 5 *

HOSTS AND DISTRIBUTION. Recorded by Burmeister from Innuus sinicus,
without locality data, presumably from a captive animal. Recorded by Piaget
under three specific names from Macacus cynomolgus, Cercopithecus cynomol-
cus, and Cercopithecus mona ; by Stroebelt from Macacus erythraeus; by
Mj oberg from Macacus silenus ; by Fahrenholz from Macacus rhesus and silenus
and from Cercopithecus sp.; and by Ferris from a long series of hosts of
the genera Macacus, Pithecus, and Rhinopithecus; from skins of wild animals
from Kashmir, the Malayan area, and the Philippine Islands. The full list
will be found in the "Host List" at the end of this volume.

NOTES. The original description given by Burmeister fortunately men¬
tions specifically the distinguishing character of this species, which is
the presence of but two pairs of abdominal paratergal plates. Since in the
long series of specimens examined by Ferris no other species having this
character appears, and since the one species that does appear is frequently
encountered on captive monkeys, the identification may be accepted as prac¬
tically certain. Ferris was able to examine the specimens of the three
species — eurygaster, breviceps, and longiceps — recorded by Piaget and found
all of these to be compounded of eurygaster and obtusus. It is probable
that viirious published records under the name of eurygaster are erroneous,
as was the identification employed by Gervais when he founded the genus
Pedicinus.

158

This species has been designated as type of the genus Phthirpedlc inus,
which is not here accepted.

Pedicinus hamadryas Mjbberg

1910. Pedicinus hamadryas Mjbberg, Arkiv for Zoologi 6:13:172; figs. 86-87-
1939. Pedicinus hamadryas Mjoberg, Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 7:613; figures 86, 87-
HOSTS .‘AND DISTRIBUTION- Type recorded as from Hamadryas sp. from the
ZooLogical Garden at Hamburg, Germany. Ferris recorded the species- from a
single specimen from "monkey," bearing no indication of place of origin.

Pedicinus obtusus (Rudow)

Figures 69, 70

1844. Pedicinus eurygaster ( Burmeister) , Gervais, In Walckenaer1 s Histoire
naturelle des insectes apteres 3:301; Plate 48, figures 1, lb.
(Mis identification)

1869- Haematopinus obtusus Rudow, Zeitschrift fur die gesamten Naturwis-
senschaften 34:169-

1880. Pedicinus eurygaster (Burmeister), Piaget, Les Pediculines, page 6 30 -
(Part; misidenti f ication)

1880. Pedicinus longiceps Piaget, Les Pediculines, page 632. (In part)
1880. Pedicinus breviceps Piaget, Les Pediculines, page 632. (In part)
1886- Pedicinus graciliceps Piaget, Les Pediculines, Supplement, page 141.
1910. Pedicinus paralleliceps Mjoberg, Arkiv for Zoologi 6:13:174; fig. 88.
1912. Pedicinus rhesi Fahrenholz, Zoologischer Anzeiger 39:54.

1916. Pedicinus vulgaris Fahrenholz, Archiv far Naturgeschichte, Abteilung

A, 81:11:32-

1917. Pedicinus parallel iceps , variety colobi Fahrenholz, -Jahrbuch der

Hamburgischen wissenschaftlichen Anstalten 34:2:3, 8.

1934. Pedicinus long iceps Piaget, Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 7:506; figures 293-296D.

1946. Pedicinus obtusus (Rudow), Hopkins, Annals and Magazine of Natural
History (Series 11) 12:564.

HOSTS AND DISTRIBUTION. The types of Haematopinus obtusus Rudow were
said to be from Semnopithecus maurus, without indication of locality. The
lectotype selected from among the material of Piaget is from Semnopithecus
pruinosus. The types of Pedicinus graciliceps Piaget were recorded merely
as from monkey. The types of Pedicinus paralleliceps Mjoberg were said to
be from Hacacus rhesus. Pedicinus vulgaris Fahrenholz was named, on the
basis of the literature only, for the specimens recorded by Piaget from
Innuus nemestrinus, these here being considered actually to represent
Piaget's Pedicinus longiceps, and therefore being obtusus. All of the
specimens noted above were apparently taken from captive animals.

The list of hosts, other than as recorded above, is very long and is
.’iven in full with all available corrections in the host list at the end of
this volume. Some ol these records are of specimens taken from captive an¬
imals, but others are from wild animals or their skins, and the indications
are that the species may occur on almost any Cercopithecoid monkey. It is
apparently the species which is most likely to be found on captive animals.

NOTES." Ferris was able to examine the types of the four supposed spe¬
cies recorded by Piaget, of which three were described by Piaget as new.
He reports (1934) that there were but two actual species recorded under
four names and that in some of the preparations both were included on the
same slides. He selected the lectotypes for longiceps as indicated above.

In the same work Ferris placed Haematopinus obtusus Rudow as unrecognizr-
able other than supposedly as a species of Pedicinus, although he recorded

159

Pedicinus obtusus (Rudow)

Figure B9

the examination of specimens from the Hamburg Museum which might conceiv¬
ably contain Rudow' s types. Hopkins (1946) has adopted a different point
of view, advocating that a neotype be selected from among the specimens in
the Hamburg Museum. By doing so no name would be left as belonging to an
unrecognizable species and as obtusus antedates lonHceps the latter would
become a synonym.

We need not go into all the arguments concerning this matter. In prin¬
ciple the writer is opposed to the replacement of any name, the application
of which is certain, by any name which is at all uncertain or clouded. Nor
is the opinion here held that every name which has been proposed and of
which the type is lost or apparently lost should have a neotype named for
it. To adopt such a procedure is to open endless vistas for abuse and
nomenclator ial instability. In this particular case, however, there is
some legitimate argument for following the suggestion made by Hopkins and
for selecting a neotype from among the specimens in the Hamburg Museum,
thus getting rid of an unattached name that has been cluttering the litera¬
ture for more than seventy-five years. The point is here somewhat reluc¬
tantly conceded and the mime obtusus is employed.

160

Pedicinus obtusus (Rudow), details Figure 70

161

Pedicinus patas (Fahrenholz)

1916. Neopedicinus patas Fahrenholz, Archiv fur Naturgeschichte, Abteilung
A, 81:11:6; Plate f, figure 2; text figure 7.

1934. Pedicinus patas (Fahrenholz), Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 7:615; figure 300G.

HOSTS AND DISTRIBUTION. Type recorded as from Cercopithecus patas with¬
out indication of locality. Recorded by Ferris from Erythrocebus whitei,
Lasiopyia kolbi, and Lasiopyia albogularis from East Africa.

Pedicinus pictus Ferris

1934. Pedicinus pictus Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 7:518; figures 301> 302.

HOSTS AND DISTRIBUTION. Type from Colobus caudatus from Mount Kenya,
British East Africa. Also recorded from Pygathrix entellus from the London
Zoological Garden.

Subfamily P0LYPLAC1NAE Ferris, new subfamily

DESCRIPTION OF THE SUBFAMILY. Hoplopleuridae in which the antennae are
always five-segmented (with the exception of one poorly described species
in which they are said to be three-segmented) and are very commonly sexual¬
ly dimorphic, the male having the apical, preaxial angle of segment three
more or less prolonged and terminating in a sclerotized point or bearing
one or two short, stout, retrorse setae dorsally. Anterior legs (except in
one species) small and with slender claw; middle legs larger than the first
and with stouter claw; third legs (with the exception of a few species )
distinctly larger than the second and with stouter claw, but not flattened.
Paratergal plates always present on at least one abdominal segment (with
the exception of one species in which they are entirely lacking) and always
with at least one of the posterior angles forming a point which is free
from the body; never forming the apices of spiracle-bearing abdominal tuber¬
cles; never overlapping. Abdomen most commonly with well-developed tergal
and sternal plates; although in a few cases without such plates other than
on the terminal and genitalic segments. Sternal plate of segment two never
extended laterally to articulate with the corresponding paratergites. Tho¬
racic sternal plate well developed, except in a few species in which it is
lacking.

NOTES. This subfamily is a bit difficult to define definitely because
of the existence of a few species which in one respect or another depart
from the normal form. Thus the genus Lemurphthirus, with one included
species, has paratergal plates only on the second abdominal segment, al¬
though in other respects it is a typical member of the group. One species
assigned to the genus Haemodipsus entirely lacks paratergites and other ab¬
dominal plates, although an apparently closely related species has parater¬
gal plates, even though they are very small. In a few species the sternal
plate of the thorax is lacking, although in most species it is well devel¬
oped.

These departures from the normal form, however, are limited to a few
species and do not disturb the general homogeneity of the group.

The hosts of the subfami ly are found among various groups of the rodents,
although a few occur on insectivores and the Lagomorpha, and one doubtful
member of the group occurs even upon an ungulate.

Sixteen genera are here recognized as belonging to the subfamily.

162

Key to the Genera of POLYPLACINAE

1. Antennae described as 3- se^rmen t ed ; ascribed to a lemur in Borneo .

. HAM0PHTU1R1 S

Antennae definitely 5-segmented . 2

With no trace of paratergal plates on any abdominal segment; from Euro¬
pean hare; one species of the genus . HAEM0D1PSUS

With paratergal plates definitely present on at least one segment of the

abdomen . 3

Paratergal plates present only on the second abdominal segment; as¬
cribed to a lemuroid in Africa . LEMURPHTHI RUS

Paratergal plates present on more than the second abdominal segment... 4
With the paratergal plates present only on abdominal segments 4-6; as¬
cribed to donkey and zebra from Africa . RATO1IA

Paratergal plates present on at least four abdominal segments . 5

Paratergal plates of abdominal segments 2-7 consisting merely of narrow,
longitudinal, sclerotized strips; ascribed to a South American rodent

. GALE0PHTH1KUS

Paratergal plates otherwise . 6

Paratergal plates of the abdominal segments very small, consisting of a
single point which projects from a slight base; ascribed to hares and

rabbits in Europe, Africa, and North America . HAEM0D1PSUS

Otherwise . 7

Paratergal plates of abdominal segments 3~6 each with the basal, ventral
angle produced, the angle bearing 2 slender setae; ascribed to a

lemur in Madagascar . PHTHIRPEDICULUS

Paratergal plates of abdominal segments 3~6 otherwise . 8

Both sexes with one of the transverse rows of setae on the abdominal
tergites and sternites with setae which are flattened and leaf-like

or cuneiform; ascribed to a South American rodent . CTENOPHTHIRUS

Setae of the abdomen otherwise . •••••9

Paratergal plates of abdominal segment 2 definitely divided longitudi-
nallyinto 2 plates, one of which lies on the dorsum and one on the
venter, the ventral portion with a flat, raised, apically free point;
occurring on the New World rodent family Heteromyidae . . . .FAHRENHOLZIA
Paratergal plates of segment 2 of the abdomen with at the most slight
evidence of being thus divided, the ventral part never independent of

the dorsal part . . .

Abdominal segments with not more than 2 median setae and a single seta
on each side near the lateral margin on any segment, either dorsally

or ventrally ; occurring on the South American genus Lagidium .

. LAGIODIOPHTHIRUS

Abdominal segments with more numerous setae . 11

Antennae with both basal and distal anterior angles of the basal seg¬
ment proloncred into a distant hook; known from Anathema in India....
. . . 7 . DOCOPHTHIRUS

Antennae not thus . • . . . 12

Paratergal plates of abdominal segment 2 with evidence of a distinct

longitudinal division into 2 plates . . . . . . . • 13

Paratergal plates of abdominal segment 2 with no evidence ol such lon¬
gitudinal division . I4

Abdomen in both sexes with distinct transverse tergal and sternal

plates . POLYPLAX

Abdomen in both sexes without such transverse plates, there being

merely a small, tubercle-like, sclerotized area about the base of
each seta: occurring on African rodents of the genus Cricetomys .

. . . . 7T . proenderleinellus

14. Paratergal plates present on a variable number of abdominal segments

10.

11.

12.

13.

163

but never on segments 7-8 . EULINOGNATHUS

Paratergal plates present on abdominal segments 2-8 . 15

15. Second plate of the second abdominal tergite in the male always at
least slightly modified, having its posterior border emarginate and
with a group of setae set in an aster-like fashion at each end of
this emargination; if the tergite is not sclerotized some modifica¬
tion of the row of setae still appears; occurring chiefly on the

rodent family Sciuridae . NEOHAEMATOPINUS

Second plate of the second abdominal tergite in the male not thus mod¬
ified; occurring on the Murid genus Acomys in Africa . SYMOCA

Genus CTENOPHTHIRUS Ferris

1922. Ctenophthirus Ferris, Contributions Toward a Monograph of the Suck¬
ing Lice, Part 3: 153-

GENERIC TYPE. Ctenophthirus cercomydis Ferris, at present the only known
species.

CHARACTERS. Hoplopleuridae of the subfamily Polyplacinae in which the
antennae are five-segmented and not sexually dimorphic. Paratergal plates
present on abdominal segments 2-8; those of segment two very small; those
of segments 3-6 narrow and with each posterior angle produced into a pro¬
nounced point. Female with a distinct tergal plate on abdominal segment
one, two tergal plates which are more or less fused together on segment two
and two tergal plates on segment eight, the remaining segments with three
tergal plates. The posteriormost plate on segments 3~8 bears a single row
of flattened, almost foliate setae. On the ventral side the arrangement of
the sternal plates is similar except that segment three bears three plates
which are fused together. Male with two tergal plates on segment three,
these more or less fused, and two sternal plates on segments 3-7, the pos¬
teriormost plate with a row of mingled, simple, and flattened setae. Spir¬
acles present on segments 3-8. Sternal plate of the thorax present.

Ctenophthirus cercotqydis Ferris
Figures 71, 72

1922. Ctenophthirus cercomydis Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 3:153; figures 100-101.

HOSTS AND DISTRIBUTION. Known only from the original record, from Cer-
comys cunicularis (recorded as fosteri) from Sapucay, Paraguay. The host
belongs to the rodent family Echimyidae.

Genus DOCOPHTHIRUS Waterston

1923- Docophthirus Waterston, Bulletin of Entomological Research 14:101.
1932. Docophthirus, Ferris, Contributions Toward a Monograph of the Suck¬
ing Lice, Part 5:303-

GENERIC TYPE. Docophthirus acinetus Waterston, the only known species.

CHARACTERS. Polyplacinae with five-segmented antennae which are not
sexually dimorphic; with the first segment having the anterior margin pro¬
duced basally and ventrally into sclerotized hooks. Second and third legs
both enlarged and practically equal in size. Paratergal plates of the ab¬
domen present on at least segments 2-6, those of segment three not longitu¬
dinally divided, those of all the segments with each posterior angle pro¬
duced into a distinct point. Abdomen of the female with two rows of tergal
setae on segments 3-7, and one row on segments one and eight, with a small
tergal plate present in connection with the anteriormost row of setae on
segments 2-7; with two rows of setae on the sternites of segments 2-7, but
with a distinct plate present only on segment two. Male with a single row

164

Ctenophthirus cercomydis Ferris

Figure 71

of tergal setae on all tergites except segment two, which has two rows,
with one row and one plate on segments one and 4-7 and two plates on seg¬
ment two; ventral ly with but one plate and one row of setae on segment two
and with but one row of setae on any other segment and no plates except on
segments seven and eight. Spiracles present on segments 2-8. Thoracic

sternal plate not developed. . . . , .... . -Klr

NOTES. None of the known material is in very good condition and possibly

the above description will require some modification.

165

Ctenophthirus cercomydis Ferris, details

Figure 72

1()6

167

Docophthirus acinetus VVaterston , details Figure 74

Docophthirus acinetus Waterston
Figures 73, 74

1923. Docophthirus acinetus Waterston, Bulletin of Entomological Research
14:101; figures 21a and 21b.

1932. Docophthirus acinetus Waterston, Ferris, Contributions Toward a Mon¬
ograph of the Sucking Lice, Part 5:304; figures 185, 186.

HOSTIS AND DISTRIBUTION. Known only from the original record in which it
was ascribed to Anathana ellloti, which belongs to the family Tupaiidae
the tree shrews.

168

Genus KULINOGNATHUS Cummings

1916. Eul Inognathua Cummings, Annals and Magazine of Natural History (ser
ies 8) 17:90.

1929. Bathyergicola Bedford, Annual Report of the Director of Veterinary
Services, Union of South Africa 15:5^5*

1932. Bathyergicola, Ferris, Contributions Toward a Monograph of the Suck¬
ing Lice, Part 5:311*

1932. Eul inognathus, Ferris, Contributions Toward a Monograph of the Suck¬
ing Lice, Part 5:318.

SYNONYM. Bathyergicola Bedford.

GENERIC TYPE. Eul inognathus dent iculatus Cummings, by original designa¬
tion. The type of Bathyergicola is Bathyergicola hilli Bedford, b^ original
designation.

CHARACTERS. Polyrplaciuae with five-segmented antennae which are not
sexually dimorphic. Anterior legs small, with slender claw. Middle legs
larger than the first, with stouter claw and at times as large as the post¬
erior legs. Paratergal plates present on abdominal segments 2-6 at least,
except that they are lacking on segment two in one species. Abdomen always
membranous throughout except for the ninth tergite and the usual genital
plates. Abdominal segments with either one or two transverse rows of setae
on most of the segments, both dorsally and veutrally, in the female and al¬
ways with but one row in the male. Spiracles present on segments 3 -7 or 3-8.

NOTES. A most unsatisfactory situation exists in regard to this genus.
Ferris (1932) recognized Eul inognathus and Bathyergicola as distinct genera,
although it was indicated that neither of these genera was entirely homo¬
geneous or sharply defined. A re-study of the question in connection with
the present work does not support the separation there accepted. In 1932,
it was indicated that the two genera could be separated by the number of
abdominal spiracles, Eul inognathus having spiracles only on segments 3-7
while Bathyergicola has them also on segment eight. But this seems to sep¬
arate species which are actually more or less alike on the basis of other
characters. A separation on the basis of the rows of setae on the abdominal
segments of the female will not permit a separation of the males. A separa¬
tion of one species called Bathyergicola laii’rensis Bedford, on the basis of
the absence of paratergal plates on segment two of the abdomen would remove
this species from association with others which resemble it in other re¬
spects. A separation of the species which have tubercles on the head brings
together certain forms which seem to be connected by host aissociations, but
is^not supported by other characters. A separation on the basis of the
presence or absence" of the thoracic sternal plate leads to an evidently ar¬
tificial grouping. Any arrangement that may be made seems to receive no
support from host or geographical distribution, yet all the species seem to
share a general similarity. The soLution of naming several genera suggests
itself but seems to offer no especially sensible arrangement, although this
mav be the eventual solution.

"This solution is in part here accepted by recognizing the two genera
Lagidiophthirus and Galeophthirus which have previously been named for
species referred to Eul i nognat hus.

Key to Species of EULINOGNATHUS

1. Head with at least 1-2 stout, sclerotized, hook-like processes or tuber¬
cles on the ventral side near the bases of the antennae . 2

Head without such hooks or tubercles . 4

2. Head with both dorsal and ventral hooks or tubercles . DENTICULAT1 S

Head with hooks or tubercles only on the ventral side . •••••• *3

3. Ventral side of first antennal segment with hooks . ACuLEATi 'S

169

Ventral side of first antennal segment without such hooks . RmNCAlUS

4. Paratergal plates present only on segments 3-6 . LAWRENSIS

Paratergal plates present on at least segments 2-6 . 5

5- Abdominal segment 7 with distinct, apically free paratergal plates .

. LOPHIOMYDIS

Abdominal segment 7 without distinct, apically free paratergal plates 6

6. Sternal plate of thorax well developed . AMERICANUS

Sternal plate of thorax not at all developed . HILLI

Eulinognathus aculeatus (Neumann)

1912. Haematopinus aculeatus Neumann, Bulletin de la Societe Zoologique de
France 37:143; figures 5> 6.

1916. Eulinognathus aculeatus (Neumann), Ferris, Proceedings of the Cali¬
fornia Academy of Sciences (Series 4) 6:168.

1932. Eulinognathus aculeatus (Neumann), Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 5:321; figures 196, 197-
HOSTS AND DISTRIBUTION. Originally recorded from " Dipus sp. , " at Dj erha,
Tunis. According to Ellerman's list this is probably a species of Jaculus.
Recorded by Ferris from Allactaga mongolica which, according to Ellerman,
is Allactaga siberica. These hosts are members of the family Dipodidae.

NOTES. There is a possibility of a misidentification in the record by
Ferris, since the types of the species were not seen.

Eulinognathus americanus Ewing

1923. Eulinognathus americanus Ewing, Journal of the Washington Academy of
Sciences 13:148.

1932. Eulinognathus americanus Ewing, Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 5:325; figure 200.

HOSTS AND DISTRIBUTION. Type from Ctenomys brasiliensis on the Salade
River, Paraguay. Recorded by Ferris from Ctenomys serlcus from the Upper
Rio Chico, Paraguay. The hosts belong to the family Echimyidae.

NOTES. The curiously modified setae on the paratergal plates of seg¬
ments 5-5 offer a basis for the generic separation of this species if this
should prove desirable. It is at present known only from the female.

Eulinognathus biuncatus Ferris

1932. Eulinognathus biuncatus Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 5:324; figures 198, 199.

HOSTS AND DISTRIBUTION. From Dipodipus sowerbyi in Shensi, China. Ac¬
cording to Ellender this is Dipus sagitta.

Eulinognathus denticulatus Cummings
Figures 75. 76

1916. Eulinognathus denticulatus Cummings, Annals and Magazine of Natural
History (Series 8) 17:90; figure.

1932. Eulinognathus denticulatus Cummings, Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 5:3 19 ; figures 194-195-
1940. Eulinognathus denticulatus surdasteri Werneck, Revista de Entomologia
11:724; figure.

HOSTS AND DISTRIBUTION. Recorded by Cummings from Pedetes coffer with¬
out indication of locality, and by Bedford from the same host in South
Africa. Ferris has recorded it from Pedetes larvalis and Pedetes sp. at
Nairobi and Machakos, British East Africa, and from Mastomys coucha and
Rattus rattus at Nairobi. Werneck has described the supposed subspecies or

170

Eulinognathus denticuiatus Cummings

Figure 75

171

head

female genitalia

Eulinognathus denticulatus Cummings, details

Figure 76

variety surdasteri from Pedetes surdaster larvalis at Nairobi, basing it
upon slight differences in the form of the pseudopenis of the male. Males
at hand from the type host at Nairobi show definitely that the supposed
difference depends merely upon whether or not the pseudopenis happens to be
turned upward. The subspecies surdasteri is consequently rejected. Ac¬
cording to Ellerman's list there are but two species of Pedetes, these be¬
ing caffer and surdaster , larvalis being a subspecies of the latter. The
hosts belong to the rodent family Pedetidae.

Eulinognathus hilli (Bedford)

1929. Bathyeriiicola hilli Bedford, Report of the Director of Veterinary
Services, Union of South Africa, 1^:506; figures 6, 7, 7a, 8.

1932. Bathyerticola hilli Bedford, Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 5:312; figures 190, 191.

1932. Proenderleinellus hilli (Bedford), Bedford, Report of the Director
of Veterinary Services and Animal Industry, Union of South Africa,
18:401.

172

HOSTS AND DISTRIBUTION. Described from Georhychus hottentotus at Pieter¬
maritzburg, Natal, South Africa, and recorded only from this host and lo¬
cality. According to Ellender the generic name of the host is Cryptonys.
It belongs to the liunily Dathyergidae.

Eulinognathus lawi-ensis (Bedford)

1929. Bathyer£icola lawrensis Bedford, Annual Report of the Director of
Veterinary Services, Unionof South Africa, 15: 5U6 ; ligs. 7b, 9. ID.
1932. Bathyergicola lawrensis Bedford, Ferris, Contributions Toward a Mon¬
ograph of the Sucking Lice, Part 5:314; figure 192.

1932. Proemierleinellus lawrensis (Bedford), Bedford, Annual Report of the
Director of Veterinary Services and Animal Industry, Union ol
South Africa, 18:401.

HOSTS AND DISTRIBUTION. Described from Bathyerius suillus ( =maritimus )
from unspecified locality in Cape Colony, South Alrica. The host belongs
to the family Bathy ergi dae .

Eulinognathus lophiomydis (Ferris)

1932. Bathyergicola lophiomydis Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 5:315; figure 193 •

1940. Bathyerticola lophiomydis Ferris, Werneck, Revista de Entomologia
11:728; figures.

HOSTS AND DISTRIBUTION. Type, the female, described as from Lophiomys
thomasi from Mount Garguez, and recorded also from Lophiomys ibeanus at
Nakroru, British East Africa. The male was described by Werneck from
Lophiomys sp., "probably testudo," from Kenya, British East Alrica. Ac¬
cording to Ellender all these names represent subspecies of Lophiomys
tmhausi. The host genus belongs to the family Lophiomyidae.

Genus FAHRENHOLZIA Kellogg and Ferris

1915. Fahrenholzia Kellogg and Ferris, Anoplura and Mallophaga of North
American Mammals, Stanford University Publications, University
Series (no volume number), page 32-

1922. Fahrenholzia, Ferris, Contributions Toward a Monograph of the Suck¬
ing Lice, Part 3:158.

GENERIC TYPE. Fahrenholzia pinnata Kellogg and Ferris.

CHARACTERS. Polyplacinae with five-segmented antennae which are not
sexually dimorphic. Anterior legs small, with slender claw. Middle and
posterior legs equal in size, with very large tibiotarsus and stout claw,
the tarsus with a sclerotized, retrorse point at the outer basal angle.
Parater'ral plates of the abdomen present on a variable number ot segments,
alwaysAowever, with at least three pairs present. The paratergal plates
of what is apparently segment two consist each of two plates, one lying on
the dorsum, the other on the venter, distinctly separated from each other,
the ventral piece provided with a flat, apically free process which arises
somewhat anterior to the apex of the plate. It is possible thai we have to
do with the paratergal plates of segments one and two, one or the other/,t
which has been somewhat displaced posteriorly. Following these plates
there are always plates on segments three and tour, these having the apical
an rles free from the body. Plates which do not have the apex thus free may
occur as far posteriorly as segment eight. Abdomen entire 1> membranous ex
cept for the usual dorsal and ventral plates of the terminal and genital
segments, each segment with but a single row of setae, both d ^or sally and
vent rally, these setae strikingly stout. Spiracles present on segments j-S.
Thoracic" sternal plate strongly developed.

173

The members of this genus occur exclusively on members of the rodent fam¬
ily Heteromyidae, which occurs in North America and northern South America.

NOTES. The status of some of the species included in this genus is not
clear. The author has in the past named certain "subspecies, " a practice
which he would not now approve, for some of these forms. These are here
considered as species. It is probable that a considerable number of forms
remain still to be discovered and until more is known about the group it is
hopeless to talk about subspecies.

Key to Species of FAHRENHOLZIA

1. Para tergal plates present only on abdominal segments 2-4 . 2

Paratergal plates present on more than these segments . 4

2. Paratergal plates of segment 3 with both apical lobes acute . 3

Paratergal plates of segment 3 with the dorsal lobe apically broad and

truncate or slightly emarginate . MICROCEPHALA

3. Paratergal plates very small and slight; male with free, elongated par-

ameres and with a distinct pseudopenis . REDUCTA

Paratergal plates well developed; parameres of the male broadly expand¬
ed; with no distinct pseudopenis . PINNATA

4. Paratergal plates of segment 3 with but a single lobe . TRIBULOSA

Paratergal plates of segment 3 bilobed . ZACATECAE

Fahrenholzia microcephala Ferris

1922. Fahrenholzia microcephala Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 3:161; figures 106, 107.

HOSTS AND DISTRIBUTION. Holotype recorded as from Heteromys pictus at
San Carlos, Vera Cruz, Mexico, which, according to Ellender, belongs to the
genus Liomys. Also recorded from Heteromys goldmani at Achotal, state of
Vera Cruz, Mexico, and from Liomys irroratus in Texas and Mexico.

Fahrenholzia pinnata Kellogg and Ferris
Figures 77, 78

1916. Fahrenholzia pinnata Kellogg and Ferris, Anoplura and Mallophaga of
North American Mammals, Stanford University Publications, Univer¬
sity Series (no volume number), page 32; text figure 13; Plate 3,
figure 2; Plate 5, figure 5; Plate 6, figure 10.

1922. Fahrenholzia pinnata Kellogg and Ferris, Ferris, Contributions Toward
a Monograph of the Sucking Lice, Part 3:159; figures 104, 106.

HOSTS AND DISTRIBUTION. Type from Dipodomys cali fornicus at Covelo,
Mendocino County, California. Recorded also from Dipodomys merriami at
Independence, California, and Dipodomys desert i at Mecca, California; from
Dipodomys ornatus at Valparaiso, state of Zacatecas, and Dipodomys phillipsii
at Amecameca, Mexico; from Perodipus sp . at Coulterville , California; from
Perognathus parvus in the Pine Forest Mountains, Nevada.

Fahrenholzia reducta Ferris

1922. Fahrenholzia tribulosa reducta Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 3:165; figure 109B.

HOSTS AND DISTRIBUTION. Recorded only from Perognathus formosus at Vic¬
torville, California.

Fahrenholzia tribulosa Ferris

1922. Fahrenholzia tribulosa tribulosa Ferris, Contributions Toward a Mon-

174

o^raph of the Sucking Lice, Part 3:163, figures 108, 109 A, D. E.
HOSTS AND DISTRIBUTION. Known only from Perofnathus californicus at
Pleasant Valley, Merced County, California.

t Fahrenholzia zacatecae Ferris

1922. Fahrenholzia tribulosa zacatecae Ferris, Contributions Toward a Mon¬
ograph of the Sucking Lice, Part 3:166; figure 109 C.

HOSTS AND DISTRIBUTION. Type trora Perofnathus hispidus at Valparaiso,
state of Zacatecas, Mexico. Specimens are at hand from the same host
species in Savala County and at Somerset, Atasco County, Texas.

175

i=^

male genitalia

thoracic sternal plate

Fahrenholzia pinnata Kellogg and Ferris, details

Figure 78

176

Genus GALEDPHTULRUS Eichler

1950. Galeophthirus Eichler, Bolletino della Societa Entomolo rica Italiaua

79: 12.

G 01 ERIC TYPE- Hul i nognat hus caulae Werueck, the only included species.
CHARACTERS. Antennae t'i ve-seguieuted , not sexually dimorphic. First
legs small and weak, with slender claw. Second and third legs about equal
to each other, large and stout, with stout claw. Paratergal plates present
ou abdominal segments 3~7, each consisting of an elongated, narrow, sclero-
tized area which bears two long setae at the apex, the apex not free from
the boify wall. Abdomen membranous throughout in both sexes except for the
usual sclerotizations of the terminal and genital segments. Female with
two rows or partial rows of setae ou each of most of the segments, the male
with but oue row. Abdominal spiracles present ou segments "1-8. Thoracic
sternal plate present.

NOTES. About the only basis for recognizing this genus appears in the
form of the paratergal plates. The writer is somewhat dubious concerning
it, but it is here accepted.

Galeophthirus caviae Werueck
Figure 79

1934. Eulinognathus caviae Werueck, Meinorias do lustituto Oswaldo Cruz 29:
183; figures 6-11.

19^0. Galeophthirus caviae (Werueck). Eichler, Bolletino della Societa En-
tomologica Italiana 79:12.

HOSTS AND DISTRIBUTION. Known only from the original record, from Galea
leucoblephara, which, according to Ellender, is a subspecies of Galea
musteloides at Jujuy, Republic of Argentina. The host belongs to the
family Caviidae.

Genus HAEMODIPSUS Enderlein

1904. Haemodipsus Enderlein, Zoologischer Anzeiger 28:139, 143-
1932. Haemodipsus, Ferris, Contributions Toward a Monograph of the Suck¬
ing Lice, Part 5 : 59 -

GENERIC TYPE. Pediculus lyriocephalus Burmeister. Two other species
are included in this genus.

CHARACTERS. Anoplura referable to the Polyplacinae . Antennae five-
segmented, not sexually dimorphic. First pair of legs small and with slen¬
der claw; second and third legs moderately stout and with stout claw, about
equal to each other. Type species with the abdomen membranous throughout
and without trace of paratergal plates; in other species of the genus with
small paratergal plates on segments 3-6, these being merely a slight, scle-
rotized point which projects from the body wall and is supported by a
slight, expanded sclerotization at the base. Abdominal segments in both
sexes with a single row of setae on each, both dorsallv and ventrally .
Thoracic sternal opiate present, but very weakly developed and at no point
free from the body. Spiracles present on abdominal segments 3-8.

The members of this genus occur on hares and rabbits of the family
Leporidae of the Order Lagomorpha.

NOTES. The situation concerning this genus is very unsatisfactory • The
type species is unfortunately but little known and the information concern¬
ing it quite incomplete. It is possible that the other species which are
referred to this genus are so placed chiefly because of their hosts. As
based upon the type species the genus can scarcely be referred to the Hop-
lopleuridae, but the other species seem to belong to this family and to the
subfamily Polyplacinae. A thorough redescription oi both sexes of the type
species is much needed.

p^r-

f (

Galeophthirus caviae (Werneck)

Figure 79

178

Key to Species oi HAfMODI PSUS

1. Paratergul plates of the abdomen entirely lacking . LYRIOCEPH ALLIS

Paratergal plates present on the abdomen . 2

2. Head slender . AFRICANUS

Head strongly widened posterior to the antennae . 3

3. Sternal plate of the thorax forming a rather narrow, transverse bar....

Sternal plate of the thorax filling the space among the coxae and more
or less hexagonal . SETONI

Haemodipsus africanus Bedford

1934. Haemodipsus africanus Bedford, Onderstepoort Journal of Veterinary
Science and Animal Industry 2:48; figure 10.

HOSTS AND DISTRIBUTION . Recorded from Lepus zuluensis at Jericho, Trans¬
vaal, South Africa.

Haemodipsus lyriocephalus (Burmeister)

Figure 80

•

1839 - Pediculus lyriocephalus Burmeister, Genera lnsectorum, Rhynchota,
Species 11.

1842. Haematopinus lyriocephalus (Burmeister), Denny, Monograph i a Anoplur-
orum Britanniae, page 27; Plate 24, figure 4.

1904. Haemodipsus lyriocephalus (Burmeister), Enderlein, Zoologischer An-
zeiger 28:143.

1932. Haemodipsus lyriocephalus (Burmeister), Ferris, Contributions Toward
a Monograph of the Sucking Lice, Part 5:330; figures 202, 203.

1935- Haemodipsus lyriocephalus (Burmeister), Freund, Die Tierwelt Mitte-
leuropas, Band 4, Lieferung 3 : 20 ; figures 87-89-
HOSTS AND DISTRIBUTION. Described from Lepus timidus in Europe. Ferris
has recorded it from Lepus ilacialis without further data. Hopkins records
it from Lepus europaeus in Europe.

Haemodipsus setoni Ewing

1924. Haemodipsus setoni Ewing, American Journal of Tropical Medicine 3: 548.
1932. Haemodipsus setoni Ewing, Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 5:335; figures 205B, E.

HOSTS .AND DISTRIBUTION. Described from Lepus californicus at Wichita,
Kansas, and recorded from the same host at San Diego, California. Recorded
by Kellogg and Ferris from the same host in California and Arizona. Speci¬
mens from an undetermined species of "cotton tail rabbit," presumably a
species of Sylvilaius , are at hand from the state of Montana.

Haemodipsus ventricosus (Denny)

Figures 81, 82

1842. Haematopinus ventricosus Denny, Monographia Anoplurorum Britanniae,
paj/e 30 ; Plate 25, figure 6.

1904. Haemodipsus ventricosus (Denny), Enderlein, Zoologischer Anzeiger
28: 143-

1932. Haemodipsus ventricosus (Denny), Ferris, Contributions Toward a Mon¬
ograph of the Sucking Lice, Part 5:332; figures 204, 205.

HOSTS AND DISTRIBUTION. Described from the European rabbit, Oryctolaius
(- Lepus ) cuniculus, from England and many times recorded from this host and
from domestic rabbits which are supposed to have been derived from this
species in many parts of the world.

179

180

Haemodipsus ventricosus <Denny> Figure 81

181

Haemodipsus ventricosus (Denny), details

Figure 82

182

Geuus HAMOPHTHIRUS MjOberg

192S. Hamophthirus Mjoberg, Psyche 32:283.

1932. Hamoph tht rus, Ferris, Contributions Toward a Monograph of the Suck¬
ing Lice, Part 5:3%.

G 01 ERIC TYPE. Hamophthirus galeopi thee i Mjoberg, the only included
species.

CHARACTERS. Unfortunately this genus is known oni) from the original,
veiy inadequate description and the accompanying crude figures. The fol¬
lowing abstract of the important characters which can be gained from these
is given.

Polyplacinae with three-segmented antenuae, the basal segment much en¬
larged and with an apical hook at the anterior distal angle. Head very
broad, its posterior angles produced each into a prominent point. Parater-
gal plates present on abdominal segments 3-7* Legs said to be "fairly
equally developed."

There is in this description and the accompanying fibres a suggestion
that this genus is rather closely related to Docophtht rus , although the
three-se gnented antenuae would immediately suffice to separate it from all
the other members of the subfamily.

Hamophthirus gadeopitheci Mjoberg

192*1. Hamophthirus £aleopitheci Mjoberg, Psyche 32:283: figure.

1932. Hamophthirus galeopitheci Mjoberg, Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 5:3^7; figure 187.

HOSTS AND DISTRIBUTION. From a Dermopteran, Galeopithecus sp., at Fes-
seltan, British North Borneo. This is a Cynocephalus variegatus.

NOTES. In spite of the inadequate description and poor illustrations it
should be possible to recognize this species if it is recovered. Ferris
(1932) merely reproduces the description and figures given by Mjoberg.

Genus LAGIDIOPH'ffllRUS Eichler

1950. La^idlophthirus Eichler, bolletino della Societa Entomologica Itali-
ana 79:12.

GENERIC TYPE- Haemodipsus parvus Kellogg and Ferris, the only included
species.

CHARACTERS. Polyplacinae with five-segmented antennae, which are not
sexually dimorphic. Anterior legs small and weak; second and third legs
large and stout, about equal to each other. Paratergal plates of the ab¬
domen present on segments 2-6, these quite small and with each posterior
angle produced into a strong point. Abdomen otherwise membranous except
for the usual plates of the terminal and genitalic segments. Body setae
very few, there being a single row, both dorsally and ventrally, on each
abdominal segment, each row having a median group of 2-4 setae and there
being a single seta, both dorsally and ventrally, close to the lateral mar¬
gin. Spiracles present only on abdominal segments 3 • Prothoracic ster¬
nal plate well developed. . .

NOTES. Recognizing a new genus for the single species included in this
renus is the only way of escaping from the problems which it presents in
regard to its generic assignment, in spite of the weakness of the charac¬
ters on which the genus is based.

Lacidiophthirus parvus (Kellogg and Ferris)

Figure 83

1915. Haemodipsus parvus Kellogg and Ferris, Anoplura and Mallophaga ol

183

Lagidiophthirus parvus (Kellogg and Ferris)

Figure 83

North American Mammals, Stanford University Publications, Univer¬
sity Series (no volume number), page 30; text figure 12; Plate 2,
figure 4; Plate 4, figure 6.

1932. Eulinognathus parvus (Kellogg and Ferris), Ferris, Contributions
Toward a Monograph of the Sucking Lice, Part 5:327; figure 201.
1940. Eulinognathus parvus (Kellogg and Ferris), Werneck, Revista de En-
tomologia 11;726; figure.

1950. Lagidiophthirus parvus (Kellogg and Ferris) , Eichler, Bo lletino della
Societa Entomologica Italiana 79:12.

HOSTS AND DISTRIBUTION. Type from Lagidium peruanum from an unspecified
locality in Peru. Werneck has recorded the species from Lagidium inca —
which, according to Ellender, is the same as peruanum — from the Cordilheira
Songo, Province of Murillo, Department of La Paz, Bolivia.

NOTES. This species was originally described from the female alone but
Werneck has described the male. It is possible here to illustrate only the
female.

184

Genus LhMIlkPWTHIKIJS Bedford

1927. Lemurphthirus Bedford, Parasitology 19:263-

1932. Lemurphthirus, Ferris, Contributions Toward a Monograph of the Suck¬
ing Lice, Part 5:299-

GENERIC TYPE. L> murphthirus galut*us Bedford.

CHARACTERS. Polyplaciuae with f i ve-segmented antennae which ai‘£ sexual¬
ly dimorphic, the male having the distal, preaxial angle of the third seg¬
ment strongly produced and curved posteriorly and bearing two short, stout
setae. Anterior legs small and weak, with weak claw; middle and posterior
legs definitely larger, but not greatly so, about equal to each other.
Paratergal plates present only on the second abdominal segment, consisting
of a simple, flat, somewhat cuniefoim, sclerotized piece, the extreme apex
of which is free from the body. Abdomen, in both sexes, with a single
plate and a single row of hairs on each segment, both dorsally and ventral-
ly, except that the female appears to have two plates and two rows ol setae
on segment two. Thorax unusually elongate, with a very large, marginally
free, sternal plate. Spiracles present on abdominal segments 3~8-

NOTES. The head and thorax of the species of this genus might very well
belong to a species of Neohaematopinus, hut the abdomen is quite diffei*ent.
The genus contains two known species.

Key to Species of LfcMl’RPHTHI KUS

Sternal plate of the thorax with a slender, median, anterior prolongation..

. VERRUCULOSUS

Sternal plate of the thorax without such anterior prolongation . GALAGUS

Lemurphthi rus galagus Bedford
Figures 84, 85

1927. Lemirphthirus galagus Bedford, Parasitology 19:263; figures.

1932. Lemurphthirus galagus Bedford, Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 6;30U; figures 183, 184.

HOSTS AND DISTRIBUTION. type from Galagus moholi (which, according to
Hopkins, is senegalensis) , a Lemuroid, from Onderstepoort, Transvaal, South
Africa and recorded at the same time from Southwest Africa. Ferris has re¬
corded the species from the same host, without data, in the British Museum.
Hopkins records it without locality from Galago demidovil .

Lemurphthirus verruculosus Ward

1951- Lemurphthirus verruculosus Ward, Entomological News 62:190; figures.

HOSTS AND DISTRIBUTION. Known only from "mouse lemur" from Bemangidy,
Fort Dauphin District, Tulear Province, Madagascar. Presumably this refers
to some species of the genus Cheirogaleus.

Genus NEOHAEMATOPINUS Mjoberg

1910. Neohaematopinus Mjdberg, Arkiv for Zoologi 6:13; 160.

1910. Acanthopinus Mjoberg, Arkiv for Zoologi tv. 13; 160.

1914. Linognathoides Cummings, Bulletin of Entomological Research 3:393-
1916. Lutegus Fahrenholz, Archiv fur Naturgeschichte , Abteilung A, 81: 11: 31-
1923. Neohaematopinus, Ferris, Contributions loward a Monograph ol the
Sucking Lice, Part 4:237- ,

1929. Ahaematopinus Ewing, A Manual of External Parasites, page 19(-
1949. Petauristophthirus Eichler, Bolletino della Societa Entomologica
Italiana 79: 12.

Lemurphthirus galagus Bedford

Figure 84

186

thoracic dorsum

thoracic sternal plate

female genitalia
Lemurphthirus galagus Bedford, details

male genitalia

Figure 85

antenna

2nd or 3rd claw

187

GENERIC TYPE. Haematopinus sciuropteri Osborn.

GENERIC SYNONYMS. Acanthopinus Mjoberg, type Haematopinus antennatus
Osborn; this specific name being preoccupied was later changed to Acantho¬
pinus sciurinus Mjoberg. Linognathoides Cummings, type Linognatho ides
citelli Cummings. Lutegus Fahrenholz, type Haematopinus ( Polyplax ) pectin-
ifer Neumann. Ahaematopinus Ewing, type Heohaematopinus inornatus Kellogg
and Ferris. Petauristophthirus Eichler, type Heohaematopinus petauristae
Ferris.

CHARACTERS. Polyplacinae in which the antennae are usually at least
slightly sexually dimorphic, the third segment in the male having the dis¬
tal preaxial angle slightly prolonged and bearing dorsally 1-2 small, stout,
recurved setae. Head usually abruptly broadened posterior to the antennae.
Legs with the first pair small and with slender claw; second and third
pairs larger and with stout claw, usually almost equal but the third pair
at times larger than the second, although never flattened and expanded.
Thoracic sternal plate usually well developed, although lacking in one
species. Abdomen always with paratergal plates on segments 3-8> and occa¬
sionally with a vestige of such plates on segment one, the plates of seg¬
ment two never divided longitudinally. Abdomen always with some develop¬
ment of tergal plates in the male and usually so in the female, but in some
species with the plates very' weakly developed or present only on segment
two or 2-3- Female normally with two transverse rows of setae on segments
2-7 dorsally and segments 3-6 ventrally, but in a few species with three
rows on these segments. Male normally with but one row of setae on any
segment dorsally, except that segment two has two rows; normally with two
rows of setae on segments 2-6 ventrally. In all known species the row of
dorsal setae which is probably the second row of segment two is associated
with an at least slightly developed tergal plate which is posteriorly emargi-
nate, with some of the setae at the lateral ends of the plate somewhat dif¬
ferentiated in size or form from the others and more or less radiately ar¬
ranged. In some species this character is weakly developed but in all it
is to some degree indicated.

NOTES. In spite of some departures by certain species from the charac¬
teristic pattern of the genus this group, as here understood, seems to be
quite natural and relatively homogeneous. The group of species for which
the name Linognathoides might be employed departs most widely from the typ¬
ical form, especially in its type species, but it would be very difficult
to advance any very cogent reason for its separation. The genus Lutegus
would, in any event, be a synonym of Linognathoides. The genus Ahaemato¬
pinus is utterly without justification and there is no satisfactory reason
for the naming of the genus Petauristophthirus.

The genus Heohaematopinus is characteristically associated with members
of the rodent family Sciuridae, but two species occur on North American
species of the genus Heotoma and perhaps some other closely related genera,
these being members of the Murid subfamily Cricetinae. One species record¬
ed from a South American rodent of the family Octodontidae probably does
not belong to Heohaematopinus.

Key to Species of NEOHAEMATOPINUS

1. Sternal plate of the thorax entirely lacking; known from Spermophilus

leptodactylus in the Caspian Sea area in Asia . CITELLI

Sternal plate of the thorax present and sclerotized . 2

2 (1). Thoracic sternal plate always with the posterior angles each pro¬

duced into a distinct point . 3

Thoracic sternal plate not with its posterior angles thus produced.
. 13

3 (2). Paratergites of abdominal segments 4-6 with not more than 2 setae

188

4 (3).

5 (4).

6 (4).

7 (6).

8 (7).

9 (3).

10 (9).

11 (9).

12 (11)

13 (2).

14 (13)

15 (14)

on the posterior border . 4

Paratergal plates ot‘ segments 4-6 each with 3 or more setae . 9

With an enlarged thorn-like seta at the extreme apex of the postax-

ial angle of the first antennal segment . 5

Without suchaseta, or if an enlarged, thorn-like seta is developed
it is not at the apex of the postaxial distal angle, being more

or less removed therefrom . 6

Abdomen of the female with a definitely developed tergal plate as¬
sociated with the anterior row of setae on segments 3— 7 ; attribu¬
ted to many species of squirrels . SQUR1NUS

Abdomen of the female with no tergal plate on segments 3— 7 ; known
only from Sciurus griseus in western United States. .. .GRISEIOOLUS
Head very broad, definitely broader than long; known from North

American flying squirrels of the genus Glauconys . SClliRUPTERI

Head definitely longer than broad . 7

Known from Sciurus vulgaris in Europe (see notes under species)....

. SCIURJ

Known from New World Cricetinae of the genus Neotoma and closely

related genera . 8

Female with no trace of abdominal tergal and sternal plates other
than those normally present on the teimiiual and genital segments
and extremely small tergal, plates on abdominal segment 2; known

from Neotoma cinerea in western United States . INORNATIJS

Female with definitely developed tergal and sternal plates associ¬

ated with the first row of setae on each abdominal segment; known
from Neotoma albigula, Neotoma fuscipes, and Neotoma micropus in
southwestern United States and Hodomys alleni in Mexico . .NED TOMAE
Paratergal plates of abdominal segments 3-6 each with 3 setae on

the posterior margin . 10

Paratergal plates of abdominal segments 3“6 each with 5-6 setae on

the posterior margin . 11

First antennal segment with a stout, thorn-like seta borne at the
apex of the distal postaxial angle; known from members of the

genus Neotamias in North America . PACIFICUS

First antennal segment with no trace of such a seta; known trom
Citellus tereticaudus in southwestern Unuted States. . .Cl TELLINUS
Paratergal plates of abdominal segments 3-6 each with 6 setae on
the posterior border, these arranged in 2 groups of 3 setae

each; known from Sciurus anomalus in §yria . SYRIACUS

Paratergal plates with the setae otherwise arranged. ...... . 12

. Paratergal plates of abdominal segments 3_6 each with 5 setae
which are arranged with a single seta near the ventral angle
and a group of 4 near the dorsal angle; known from Funambulus

in Ceylon . . • • • •CEY1DNICUS

Paratergal. plates of abdominal segments 4-5 each with their setae
arranged in 2 groups, the dorsal group with 4 and the ventral^
group "with 3; known from Funambulus palmarum in India ECHINATUS
Paratergal plates of abdominal segments 3~6 each with one or both
of their posterior angles produced into a short, slender, some¬
what finger-like process; known from Abrocoma cinerea (a Crice-

tine) in Peru . .

Paratergal plates not so . . • • . . •••••>•* *

. Female with 3 rows of tergal setae on abdominal segments J-b...lb

Female with 2 rows of setae on these segments . 18

. Abdomen of the female with sclerotized tergal and sternal plates
present only on the terminal and genital segments; male with
the pseudopenis joined to the apices of the parameres. known
from Atlantoxerus getulus in Africa . PECTINIFER

189

16 (15).

17 (16).

18 (14).

19 (18).

20 (18).

21 (20).

22 (21).

23 (22).

Abdomen of the female with well developed tergal and sternal
plates in both sexes; male with the pseudopenis enclosed be¬
tween the parameres; a group of species from African squirrels

of the genera Heliosciurus and Paraxerus . 16

Antennae with the distal postaxial angle somewhat produced and

bearing a stout, apically blunt seta at its extreme apex .

. HELIOSCIURI

Antennae with the distal postaxial angle not at all produced and

if it bears a seta this is small and apically acute . 17

Genitalia of the male with the parameres having their lateral

margin strongly arcuate . SUAHELICUS

Genitalia of the male with the outer margin almost straight .

. EENYAE

Thoracic sternal plate in the shape of a 7-sided polygon, all the

sides of which are almost straight . 19

Thoracic sternal plate otherwise shaped . 20

Tergal and sternal plates in both sexes entirely lacking except
for those present on the terminal and genital segments; para-
tergal plates extremely small; known from Petaurista petaurista

in the Malayan area . BATUANAE

Tergal and sternal plates definitely developed on all abdominal
segments in both sexes; paratergal plates strongly developed;

known from Petaurista inornatus in Kashmir . PETAURISTAE

Thoracic sternal plate longer than wide, relatively narrow, some¬
what irregular in shape; known from Xerus inauris in Africa....

. FAUREI

Thoracic sternal plate as wide as long or wider, usually more or
less transversely oval; a group of forms occurring on the Mar-
mota section of the Sciuridae (see notes under laeviusculus ) .21
Thoracic spiracles notably large, their diameter equaling about
one-half the length of the second coxae; known from Marmota and
perhaps occurring on some species of Citellus in North America.

. MARMOTAE

Thoracic spiracles smaller, scarcely exceeding one-fourth the

length of the second coxae and usually smaller . 22

Rows of tergal and sternal setae on the abdomen continuous across
each segment,, not interrupted by bare areas . 23

Rows of tergal and sternal setae across the abdomen interrupted
by bare areas which divide them into lateral and median groups

. MATHESONI

Setae of the ventral rows of segments all noticeably stout (ac¬
cording to the original description) ; described as occurring on

Citellus adocetus in Mexico . TRAUBI

Setae of all the ventral rows slender; 2 described species of
which one, PATIKI, is probably a synonym of . LAEVIUSCULUS

Neohaematopinus batuanae Ferris

1923. Neohaematopinus batuanae Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 4:261; figure 167B.

HOSTS AND DISTRIBUTION. From Petaurista batuana from the Batu Islands,
Malaysia.

Neohaematopinus ceylonicus Ferris, new species

1922. Neohaematopinus echinatus (Neumann) , Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 4:250; figure 161. (Misiden-
ti fication)

190

HOSTS AND DISTR1 Lil'T ION . Type a male, upon which the illustration cited
was based, from Funambulus palmar m at Colombo, Ceylon. Allotype and para-
types from the s;une species at Kandesanturai , Ceylon, received through the
kindness of Mr. Gordon 15. Thompson.

ChARACTERS. Male as described and illustrated in the reference cited.
Female about ihiun . Ion . As in the male the paratergites of abdominal seg¬
ments each bear four stout setae of varying length on the dorsal half
and a single seta on the ventral half, this being well separated from the
others.

NOTES. It was indicated by Ferris that the single specimen upon which
his description was based did not agree entirely with the description given
by Neumann and it is now clear that it does not represent Neumann's species,
since specimens of the latter are now available which agree with the types.
It is undoubtedly close to the true echinatus.

Neohaematopinus citelli (Cummings)

1914. Linoftnathoides spermophili Cummings, bulletin of Entomological Re¬
search 5:160; figure 3* (Specific name preoccupied)

1916. L ino£nathoides citelli Cummings, Annals and Magazine of Natural His¬
tory (Series 8) 17:107.

1923. Neohaematopinus citelli (Cummings) , Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 4:261; figures 168, 169 •

HOSTS AND DISTRIBUTION . Originally recorded from Citellus leptodactylus
and from Cr icetulus phaeus from Transcaspia. The record from Cr icetulus is
almost certainly an error and can probably safely be disregarded. The
probable true host is now known as Spermophilopsis leptodactylus.

Neohaematopinus citellinus Ferris

1942. Neohaematopinus citellinus Ferris, Microentomology 7:85; figure 41.

HOSTS .AND DISTRIBUTION. Type from Citellus tereticaudus at Tucson,
Arizona, and other specimens attributed to Ammospermophilus harrissi from
the Santa Rita Mountains, Arizona, United States.

Neohaematopinus echinatus (Neumann)

1909. Raematopinus (Polyplax) echinatus Neumann, Archives de Parasitologie
13:517; figures 19, 20.

1912. Neohaematopinus echinatus (Neumann), Cummings, Bulletin of Entomo¬
logical Research 3:393-

HOSTS AND DISTRIBUTION. Recorded from Funambulus palmarum from Rajkote,
India, this being some other species of the genus. Specimens are at hand
from the same host genus at Agra, India, which agree with the original de¬
scription, these received through the kindness of Mr. Gordon B. Thompson.

NOTES. The specimens at hand from Agra indicate clearly that this
species was misidentif ied by Ferris. The species attributed by him to
echinatus is here described as Neohaematopinus ceylonicus .

Neohaematopinus faurei (Bedford)

1920 LlnoQnatholdes faurei Bedford, Report of the Director of Veterinary'
Research, Union of South Africa 7-9:710; Plate 1, figure 2; Plate

7, figure 3- . _

1932. Neohaematopinus faurei (Bedford), Ferris, Contra but 10ns Toward a
Monograph of the Sucking Lice, Part 5:292; figures 1<8, 179 -
HOSTS AND DISTRIBUTION. Recorded by Bedford from Geosc iurus capensis,
which is now known as Xerus inauris, Bloemfontein, Orange Free State. It

191

was later recorded by Bedford from this host in other localities in South
Africa, and from Tatera and Rattus coucha. Specimens at hand received from
Bedford are indicated as being from Mongoose. It is practically certain
that the true host is the species of Xerus.

NOTES. This is an extreme member of the genus, even of the section to
which the name Li nognat ho ides has been applied. The male has no sclero-
tized tergites and the characteristic form of tergite two, with its associ¬
ated grouping of setae, is here represented only by a slight irregularity.

Neohaematopinus griseicolus Ferris

1923. Neohaematopinus sciurinus griseicolus Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 4:248.

HOSTS AND DISTRIBUTION. Type from Sciurus griseus at Inverness, Marin
County, California, U. S. A. Also recorded from the same host at other lo¬
calities in California.

NOTES. It is perhaps inconsistent to recognize this form as a species,
in view of the wide range of forms retained in sciurinus, but all the spec¬
imens at hand agree in the almost complete absence of abdominal tergal and
sternal plates in the female, while in all other specimens included in
sciurinus no such departure occurs.

Neohaematopinus heliosciuri Cummings

1913* Neohaematopinus heliosciuri Cummings, Bulletin of Entomological Re¬
search 3:393; figure 1.

1923. Neohaematopinus heliosciuri Cummings, Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 4:255; figures 164, 16 5A, C,
E, H.

HOSTS AND DISTRIBUTION. Type from Paraxerus (as Heliosciurus) palliatus
from Uchweni Forest, Witu, British East Africa. Also recorded from Paraxer¬
us ochraceus (as Paraxerus jacksoni and as Parasc iurus animosus) from vari¬
ous localities in British East Africa.

Neohaematopinus inornatus (Kellogg and Ferris)

19 15 - Linognathoides inornatus Kellogg and Ferris, Anoplura and Mallophaga
of North American Mammals, Stanford University Publications, Uni¬
versity Series (no volume number), page 25; text figure 10; Plate
4, figure 7; Plate 5> figure 5; Plate 6, figure 3*

1923. Neohaematopinus inornatus (Kellogg and Ferris), Ferris, Contribu¬
tions Toward a Monograph of the Sucking Lice, Part 4:252; figures

162, 163-

1929. Ahaematopinus inornatus (Kellogg and Ferris), Ewing, A Manual of Ex¬
ternal Parasites, page 198.

1942. Neohaematopinus inornatus (Kellogg and Ferris), Ferris, Microento¬
mology 7:84; figure 39*

HOSTS AND DISTRIBUTION. Type from Neotoma cinerea, South Yolla Bolly
Mountain, Tehama County, California. Recorded also from the same host from
Yosemite Valley, California, and from "mountain rat," which was almost cer¬
tainly the same host, from Colorado.

NOTES. This species has been designated as type of the genus Ahaemato-
pinus tVing, a genus for which there is, in the opinion here held, no ex¬
cuse whatsoever.

Neohaematopinus kenyae Ferris

1923. Neohaematopinus kenyae Ferris, Contributions Toward a Monograph of

192

the Sucking Lice, Part 4:258; figures 16 f)IJ , F, G.

HOSTS AND DISTRIBUTION. From He l lose iurus gambtanus (as keniae) , Mount
Kenya, British East Africa.

Neohaematop iuus laeviuscuius (Grube)

1851. Perticulus laeviuscuius Grube, In M iddendorff ' s Reise 2:498; PLate
32, figure 7. (Figure labelled s permophili)

1896. Haematopinus montanus Osborn, United States Department of Agricul¬
ture, Division of Entomolor^y , Bulletin (new series) 5:184; lig. 107.
1900. Haematopinus columbianus Osborn, Canadian Entomologist 32:215-
1904. Polyplax laeviuscula (Grube) , Enderlein, Zoologischer Anzeiger 28: 142.
1923. Neohaematopinus laeviuscuius (Grube), Ferris, Contri hut ions Toward a
Monograph of the Sucking Lice, part 4:264; tigs. 170, 171A, L, D, G.

HOSTS AND DISTRIBUTION. Described from Cite llu s (as Spermophi lus)
eversmanni, .Jakutsk, Siberia. Later recorded from this host 1 rom Altai,
Siberia, and from a long list of other species ol Cite l lus 1 1 om North
America ranging from Point Barrow through westeni United States into Mexico,
and from Cynomys leucurus from Colorado in the United States.

NOTES. A re-examination of all the available material shows no satis¬
factory basis for breaking this species up, although a considerable degree
of variation exists. Rubin has named three species of this group, which
will here be listed as distinct although the opinion is held that they can¬
not be definitely recognized. Even the species marmotae , which is probably
the most definitely differentiated form in the group, presents no very pre¬
cise limits turd some specimens have been seen which are dubiously referable
either to this or to laeviuscuius.

The prayer may be voiced that future students will attempt to learn some-
thing about the group before engaging in the indiscriminate naming of new
species.

Neohaematopinus longus Werneck

1948. neohaematopinus longus Werneck, Revista brasiliera de Biologia (8)2:
175; figure.

HOSTS AND DISTRIBUTION. Recorded from Abrocoma cinerea at Caccachara
near Llave, Peru. The host is a member of the subfamily Abrocominae of the

family Echimyidae. . , , ... „ . , .

NOTES This species is known from but a single female which was evident
ly very imperfectly prepared for study. It is highly probable that it does
not belong to this genus.

Neohaematopinus marmotae Ferris

192S Neohaematopinus marmotae Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 4:268; figures 171C, E, F.

HOSTS AND DISTRIBUTION. Type from Ha rmot a flaviventns, iosemite Nation
al Park, California. Recorded also from M armota species at Florence,
Montana! Specimens which agree very closely with the type are at hand 1 rom
unspecified Harmota at Bannock, Idaho, and from La Manga Pass, Colorado.
Fe??is has previously doubtfully referred a specimen from Harmota aurea

from Pamir, Asia, to this species. . 11 +

NOTES. While in its typical form this species seems clearly to be sep

arable from laeviuscuius, specimens have been examined from Armota and
from species of Cttellus which raise some question as to the validity ol
Ke species, or at Least as to its Limits and definition. The study ol
more material will be required lor any satisfactory solution ol the problem.

193

Neohaematopinus mathesoni Rubin

1946. Aeohaematopinus mathesoni Rubin, Proceedings of the Entomological
Society of Washington 48:121; figure 1.

HOSTS AND DISTRIBUTION. Described from "citellus v. couchi , " which is
presumably Citellus uar iegatus couchi from the state of Nuevo Leon, Mexico.

NOTES. A specimen from the type lot of this and other specimens from
the same host species (as subspecies grammurus) are at hand from Arizona.
It is very doubtful that the species can be distinguished from laev iusculus.

Neohaematopinus neotomae Ferris

1942. Neohaematopinus neotomae Ferris, Microentomology 7:84; figure 40.

HOSTS AND DISTRIBUTION. T^pe from Neotoma albigula at Tucson, Arizona,
and other specimens from the same host near Tucson. Also recorded from
Neotoma streatori from the Hastings Reservation near Monterey, California,
and from Hodomys alleni from Manzanillo, Mexico. Specimens from Neotoma
micropus from White Sands, New Mexico, have been somewhat doubtfully re¬
ferred to the species.

Neohaematopinus pacificus Kellogg and Ferris

1915. Neohaematopinus pacificus Kellogg and Ferris, Anoplura and Mallophaga
of North American Mammals, Stanford University Publications, Uni¬
versity Series (no volume number), page 28; text figure 14C, D;
Plate 5, figures 3, 7a, b.

1923. Neohaematopinus pacificus Kellogg and Ferris, Ferris, Contributions
Toward a Monograph of the Sucking Lice, Part 4:249; figure 160A-D.
HOSTS AND DISTRIBUTION. TyPe from Neotamias townsendii at Freestone,
Sonoma County, California. Recorded also from Neotamias hindsi, merriami,
alpinus, and speciosus (as Eutamias) from various localities in California,
U. S. A.

Neohaematopinus patiki Rubin

1946. Neohaematopinus patiki Rubin, Proceedings of the Entomological Soci¬
ety of Washington 48:121; figures 2, 5.

HOSTS AND DISTRIBUTION. Recorded as from an undetermined species of the
subgenus Ammospermophilus of the genus Citellus at Delta, Utah.

NOTES. Unfortunately no material from any species of Ammospermophilus
is available and consequently it is not possible to offer any observations
in regard to this supposed species of Neohaematopinus, other than to ex¬
press extreme doubt that it can be recognized.

Neohaematopinus pectinifer (Neumann)

1885- Haematopinus setosus Piaget, Les Pediculines, Supplement, page 143;
Plate 15, figure 6 (preoccupied) .

1909. Haematopinus ( Polyplax ) pectinifer Neumann, Archives de Parasitologie
15:528; figures 28 , 29.

1914. Ltnognathoides pectinifer (Neumann), Cummings, Bulletin of Entomo¬
logical Research 5:160.

1916. Luteius pectinifer (Neumann), Fahrenholz, Archiv fur Naturgeschichte.
Abteilung A, 81:11: 31 -

1923. Neohaematopinus pectinifer (Neumann), Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 4:268; figure 172. (Descrip¬
tion of male)

1932. Neohaematopinus pectinifer (Neumann), Ferris, Contributions Toward a

194

Monograph of the Sucking Lice, Part 5:291; figure 177. (Descrip¬
tion of female)

HOSTS AND DISTRIBUTION. Recorded from Xerus tetulus (Now referred to
Atlantoxerus) in South Africa.

Neohaematopinus petauristae Kerris

1923. Neohaematopinus petauristae Kerris, Contributions Toward a Monograph
of the Sucking Lice, Part 4:258; figures 166, 167A, C, E.

1949. Petaur istophthirus petauristae (Kerris) , Eichler, Boiletino della So-
cieta Entomologica Italiana 79:12.

HOSTS AND DISTRIBUTION. From Petaurista inornate in Kashmir.

NOTES. This species has been designated as type of the genus Petaur is¬
tophthirus Eichler, a genus for which no very cogent reason can be offered.

Neohaematopinus sciuri Jancke

1931. Neohaematopinus sciuri .Jancke, Zeitschrift fur Paras i ten kunde 4:241;
figures.

HOSTS .AND DISTRIBUTION. Described from Sciurus vulgaris in Germany and
later recorded from the same host in that country. North American and
Asiatic material that might be referred to this species is as follows: from
Sciurus aberti in Arizona, Sciurus carolinensis from Mississippi, Sciurus
poliopus from Oaxaca, Mexico — all these Irom North America; 1 rom Malayan
squirrel" in the Zoological Gardens of London, from Callosciurus finlaysoni,
caniceps, and procerus in the Malayan area. Other specimens approach the
condition seen in these with various degrees of closeness.

See notes under Neohaematopinus sciurinus.

Neohaematopinus sciurinus Mjoberg

1891. Raematopinus antennatus Osborn, United States Department of Agricul¬
ture, Division of Entomology, Bulletin (old series) 7:25; figure
13. (Not Raematopinus antennatus Piaget)

1910. Acanthopinus antennatus (Osborn), Mjoberg, Arkiv for Zoologi 6:161.
1910. Acanthopinus sciurinus Mjoberg, Arkiv for Zoologi 6:161.

1915. Neohaematopinus antennatus (Osborn), Kellogg and Ferris, Anoplura

and Mallophaga of North American Mammals, Stanford University Pub¬
lications, University Series (no volume number) , page 36; text
figure 14A, B; Plate 5, figure 10; Plate 6, figure 5.

1916. Neohaematopinus antennatus semifasciatus Ferns, Psyche 23:1UU.

1919. Neohaematopinus macrospinosus Fahrenholz, Jahresbericht des nieder-

sachsischen zoologischen Vereins zu Hannover 5~ 10. 24.^

1923. Neohaematopinus sciurinus (Mjoberg), Ferris, Contributions Toward a
Mono Taph of the Sucking Lice, Part 4:243; figures 155> l-®> ^59-
HOSTS AND DISTRIBUTION. The types of Osborn's Raematopinus antennatus
were from Sciurus niter rufiventer at Ames, Iowa. The types of Mjoberg s
Acanthopinus sciurinus were recorded as from Sciurus vulpinus, which is a
synonym of niter, in the Hamburg Zoological Garden, Germany. Later record¬
ed by Ferris from a long list of species of Sciurus and related genera from
North .America, Central America and South America and from the Malayan
Region. The list will not here be repeated, since it will be necessary for
future workers to reconsider the entire situation in any case before the
problem of the extent of the species can be settled. This is considered in

theN0TESOW1Tbenven’ considerable amount of material at hand has been care¬
fully renewed in connection with this work, with results but little more
satisfactory than those recorded by Ferris in 1922. Within this material

195

the only character that seems to offer any basis for a separation into
species is that of the enlarged seta on the first antennal segment. In
typical sciurinus this seta is quite large and is borne upon a prolongation
0 t the posterior apical angle of the segment, the whole structure forming a
pronounced hook. Every degree of variation in this structure is present in
the material at hand, down to specimens in which the seta is scarcely pres¬
ent at all. Attempts at arranging this material in groups according to
hosts and according to geography have revealed no logical pattern. Thus,
specimens from squirrels of the genus Callosciurus in the Malayan area are
practically identical with specimens from Sciurus aberti from Arizona in
the United States.

Certain of the material at hand would apparently be referable to Neo¬
haematopinus sciuri Jancke, described from the European Sciurus vulgaris,
on the basis of the development of this seta on the first antennal segment.
But if we attempt to group specimens on this basis, again no logical pat¬
tern appears. While the species sciuri is here listed, this is done solely
in order not to prejudice any development of later studies and for the
present all other material of this type is referred to sciurinus.

The problem must be left to future workers who may be able to accumulate
a great mass of material from a long series of squirrel species.

Neohaematopinus sciuropteri (Osborn)

Figures 86, 87

1891. Haematopinus sciuropteri Osborn, United States Department of Agri¬
culture, Division of Entomology', Bulletin (old series) 7:23; fig¬
ure 12.

1910. Neohaematopinus sciuropteri (Osborn), Miobers, Arkiv for Zoolo^i 6*
160; figure 79.

1923. Neohaematopinus sciuropteri (Osborn), Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 4:241; figures 156, 157.

HOSTS AND DISTRIBUTION. Described as from Sciuropterus volucella, which
is a synonym of Glaucomys volans, at Ames, Iowa. Later recorded from
Glaucomys sabrinus at Yosemite National Park, California.

Neohaematopinus suahelicus Ferris

1923. Neohaematopinus suahelicus Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 4:258; figures 165B, F, G.

HOSTS AND DISTRIBUTION. Described as from Paraxerus palliatus from
British East Africa, and recorded from Paraxerus ochraceus (as jacksoni and
as Parasciurus animosus) from the same area.

Neohaematopinus syriacus Ferris

1923. Neohaematopinus syriacus Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 4:250; figure 160E.

HOSTS AND DISTRIBUTION . From Sciurus anomalus (as syriacus) from Sjyria.

Neohaematopinus traubi Rubin

1946. Neohaematopinus traubi Rubin, Proceedings of the Entomological Soci¬
ety of Washington 48:120; figures 3. 4.

HOSTS AND DISTRIBUTION. Described as from Citellus adocetus from the
state of Michaocan, Mexico.

NOTES. This species is supposed to be separable from laeviusculus and
other members of that group by having the abdomen with three rows of setae
on each "typical segment." However, the accompanying illustration does not

196

iuropteri (Osborn)

Figure 86

197

female genitalia

male genitalia

Neohaematopinus sciuropteri (Osborn), details

Figure 87

198

agree with the description. It provides only fourteen rows of setae to be
distributed among six segments, this allowing but two rows per segment with
two left over, which does not indicate that a "typical segment" can have
three rows. The discrepancy is probably due to an error in counting and
the surmise may be hazarded that the species is not distinguished as its
author supposed. However, according to the illustration given, the stout
ness of the ventral setae may possibly afford a basis for its recognition.

Genus PHTHIRPEDICULUS Ewing

1922. Phthirpediculus Ewing, Journal of the Washington Academy of Sciences
13:148.

1932. Phthtrpedtculus, Ferris, Contributions Toward a Monograph ol the
Sucking Lice, Part 5:295.

GENERIC TYPE. Phthtrpedtculus propithect Ewing, the only included spe¬
cies.

CHARACTERS. Polyplacinae with five-segmented antennae which are sexual¬
ly dimorphic, the male having the distal, preaxial angle of the third seg¬
ment produced and bearing dorsally two, stout, retrorse setae. Anterior
legs small, with weak claw; middle and posterior legs about equal to each
other, enlarged and stout, with stout claw. Paratergal plates present on
abdominal segments 3-6, distinctly developed and with free apical angles;
marked by having the basal, inesal angle produced into a distinct sclerotiza-
tion which extends somewhat toward the mid-line ol the body and bears two
slender setae. Otherwise the abdomen in both sexes is membranous except
for the usual terminal and genitalic plates. Each abdominal segment, both
dorsally and ventrally, with one row ot setae except that in the female the
dorsum of segment two apparently has two rows. Abdominal spiracles present
on segments 3-8. Prothoracic sternal plate distinctly developed, of a
pecaliar type, being divided longitudinally into two plates, each ol which
bears a pair of slender setae at its posterior end.

Phthirpediculus propitheci Ewing
Figures 88, 89

1922. Phthirpediculus propitheci Ewing, Journal of the Washington Academy

of Sciences 13: 149 • A T .

1932. phthirpediculus propitheci Ewing, Ferris, Contributions lowara a
Monograph of the Sucking Lice, Part 5:296; figures 180, 181.

HOSTS AND DISTRIBUTION. From Propithecus edwardsii, a lemur, from Mada¬
gascar.

Genus POLYPLAX Enderlein

1904.

1907.

1909.

1923.

1929.

1935-

1838.

1938.

>olyp lax Enderlein, Zoologischer Anzeiger 28:142, 223.

Iremophthirius Glinkiewicz, Sitzungsberichte der mathematischnatur-
w is sensch aft lichen Class der kaiserlichen Akademie der Wissen-

schaften zu Wien 116:381. • * 1 • 19. £90

laematopinus {Polyplax) , Neumann, Archives de Parasitologie 13: >"•
'olyplax, Ferris, Contributions Toward a Monograph of the Sucking

Lice, Part 4:184.

^olyplax, Ewing, A Manual of External Parasites, page 1J/.
n olyplax , Ewing, Proceedings of the Biological Society of Washington

Irem'ophthirius, Ewing, Proceedings of the Biological Society of Wash-

30 iyp i ax , ^Fahrenho lz , Zeitschrift fur Parasitenkunde 10 : 239 -
Iremophthirius, Fahrenholz, Zeitschrift fur Parasitenkunde 10:„4Z.

199

Figure 88

k i

\ mf I-tiir.

female genitalia

Phthirpediculus propitheci Ewing

201

GENERIC TYPE. Pediculus spinulosus Burmeister.

GENERIC SYNONYM. Eremophthirius Glinkiewicz, of which the type by mono-
typy is Eremophthirius werneri Glinkiewicz.

CHARACTERS. Polyplacidae with five- segmented antennae which are usually
at least slightly dimorphic, the males commonly having the distal, preaxial
angle of the third segment somewhat produced and bearing at its apex a re-
curved spine or short recurved, stout seta. Head usually abruptly widened
behind the antennae, but not always so. Thorax usually with a well devel¬
oped sternal plate which around its margins is free from the body wall.
Anterior legs small and weak, with slender claw. Middle leers definitely
somewhat larger, with stouter claw. Posterior legs similar to the second
but still larger, with stouter claw, but never strongly flattened or ex-
panded. Abdomen with paratergal plates always developed on segments 2-8
the plates rarely if ever overlapping each other in an expanded specimen
the plates of segment two having the appearance of being divided longitudi-

+ ir +y+i-t°/WO Ple?es> one of which lies upon the dorsum. It is possible
that this dorsal piece actually represents the paratergite of segment one
although occasionally the plates of segment three are similarly divided’
Paratergites of segments 4-6 usually with each posterior angle produced in-
to a siight point, or occasionally into a slender process^ the posterior
margm between these points being entire; at times with only the ventral
angle produced. Tergal and sternal plates always developed and sclerotized,
the female having two plates and two rows of setae on segments 4-7 dorsallv
and on sepients ventrally; the male with not more than one plate and
one row of setae on any segment dorsally and with either one or two plates
on the abdominal segments ventrally. The ventral plates on segments 2-2
are never produced laterally to articulate with or approximate the parater¬
gal plates of these segments. v

NOTES. The opinion previously expressed by Ferris that Eremophthirius
should not be separated from Polyplax is here still maintained. Fahrenholz
has attempted to revive Eremophthirius on the basis of the presence in the
male of but one ventral plate on any abdominal segment, but any such divi-
sion results merely m quite meaningless groupings of the species.

The members of the genus Polyplax occur almost exclusively upon members

,'ly althouSh at leasl ■>"<= species seems to occur

normal upon members of the msectivore family Soricidae.

Key to Species of POLYPLAX

in AVS fn^deraUe ' °f difficulty has been eucouutered ip prepar-

JS ™ y fpecles °flh;S genus' The characters separating some of

the species are clear enough when specimens are compared directly, but are

y treno.hant to be “f •“=" use in a key, or are of such a
nature that they cannot be expressed with the conciseness which is desir¬
able in a key. Furthermore, of some species only one sex is known In
some other instances the male only may present especially distinctive char¬
acters. Because of these facts the presentation of separate kevs to the
sexes is no more practicable than it is to include both in the same key.
This introduces serious difficulties, but at the present time there seems
to be no way to avoid them.

It should be remembered, also, that some of the species are known from
but very few specimens and consequently the possible range of variation is
unknown Jnder these circumstances the key must be used with caution.

Ihe following species, known only from their original descriotions arp

(Bilmeiste^r. lhC ^ dentattc0rnts Ewin*r> eriopepll (Ewing ), spini£era

202

1. Posterior legs with a distinct, sclerotized, retrorse tooth at the oub-
er basal angle of the tibia; known from Rattus sabanus iu the Malayan

. INSULSA

dl Uli* . . . . ^

Posterior legs without such a tooth . • . . . "

2 (1). head with a long seta at or close to the apex of the lateral, pos¬

terior angle . •••••**' . ,*.*.* * 'i* v

head with this seta borne on a small, apically free, ear- like lobe
which is somewhat removed from the apex of the posterior lateral
angle; occurring on New World species of the genera Peromjscus,
Onychomys, and Retthrodontomys . AUK1CULAR1S

3 (2). Abdomen with tergal and sternal plates undeveloped in the feuutle,

except for those associated with the genital segments and one or
two very small tergal plates at the base of the abdomen; known

from various hosts in India . •• . ASIATIC A

Abdomen of the female always with distinct, even it attenuated, tei -
tral and sternal plates on all segments . ...4

4 (3). Thoracic sternal plate with a distinct, narrow, hanlle-like proton

.ration extending forward between the anterior coxa, this prolon¬
gation being one-fourth as long as the plate itselt or longer...*)
Thoracic sternal plate at the most with nothing more than a slight

median, anterior point . . . -***::* - - •••]“

5 (4). With a pair of long setae on each abdominal paratergite, these se ae

longer than the plate which bears them . . . '.V *.*’.***,' . b

With not more than 1 seta on any paratergal plate which is longer
than the plate which bears it, and such setae not present at all

6 (5). Lateral margins of the hindhead in the femaie short and almost semi-

circularly convex (known only from the female); from undetermined

rodents from Abyssinia . . . . . . .PKj^J.lbA

Lateral margins of the hindhead straight and almost parallel in
both sexes; known from Tatera indica in India . oihrnhNol

7 (f)). Male with a row of very short, almost thorn-like, setae on the ter

cral plates of segments 4-7, in addition to the usuai slender

setae; known from Tatera bohwi in Africa . . . . .ttlbhKl A1A

Male with only the usual row of slender setae on any abdominal ter^

8 (7) Setae on paratergites of segments 3~6 almost or quite equaling in

8 (,)’ "length the plate that bea^rs them except for one longer seta on

se.ment 3; known from Tatera vicina in Africa. . ......... • • 1 AILKAh

Se2? on paratergites of segments 3-6 shorter than the plate which
"bears them except for 1 long seta on segment 3 or se^nnents 3 • 9

9 (8). Paratergal plates of segment 3 only with 1 long seta which^is^much

longer than the plate that bears it . . . . .

Paratergal plates of segments 3-4 each with 1 long seta which is

much longer than the plate that bears it. . • . .

10 (0) Ventral apical angle of paratergites ol abdominal segments J-b

10 (9). Ventra^ ^ that .g about twice as wlde at its base and

twice as long as the tooth at the dorsal angle; known iron Ger-

billus pyramidum in Egypt. . ........ .WXBILLi

Ventral apical angle of paratergites of segments 3-6 forming mere
ly a slight tooth which is not larger than that of the doi sa ^

11 (10) Lateral margins* oV "the "hindhead" in "the male (female unsown)

11 (10). La^tr* Convergent ; occurring on Pachyuromys in

Lateral margins of the hindhead in the male ivergen 0 P

n),llv parallel ; from Meriones auceps m China .

19 (4) ferater^'i teso f abdominal segments 2-6 with both posterior angles

12 (4K produced into a slender, apically acute process which is heset

203

13 (12)

14 (13).

15 (13).

16 (15).

17 (15).

18 (17).

19 (18).

20 (18).

with minute squamations . WATERSTONI

Paratergites not so . \\\ w

. Paratergal plates of abdominal segments 3-6 posteriorly emargi-
nate, the setae borne close to the apices of the points formed

by this emargination . .

Paratergal plates of these segments not so . 15

, Paratergites of abdominal segments 3~6 only shallowly emarginate;
tergal plates of abdominal segments 3-6 occupying scarcely more
than half the width of their respective segments; occurring on

Arvicanthis in Africa . 1 . ABYSSINICA

Paratergites of segments 3-6 deeply emarginate; tergal plates of
segments 3-6 occupying at least three- fourths of the width of
their respective segments; occurring on Arvicanthis in Africa..

Tergal plates of abdominal segments 3-6 each with the ventral pos¬
terior angle only forming a tooth and each with a pair of setae

which are about as long as the plate which bears them . 16

Tergal plates of abdominal segments 3-6 not presenting this com¬
bination of characters; if the setae are as long as the plates
each angle forms a tooth, if one angle does not form a tooth

Head truncate anteriorly immediately 'in 'front* of’ the’ antennae ;

occurring on Lophuromys in Africa . PHTHISICA

Head acutely pointed in front of the antennae; . OXYRRHYNCHUS

Genitalia of the male with the pseudopenis entirely enclosed with¬
in the apices of the parameres; tergal plates in the female all
ot almost uniform length in the longitudinal axis of the body-

occurring on Saccostomus in Africa . JONESI

Genitalia of the male with the pseudopenis art icuiating to the
apices of the parameres; anterior tergal plate of abdominal
segments 4-/ in the female distinctly longer in the longitudi¬
nal axis of the body than the posterior plate of tho same seg¬
ment .

Paratergites of abdominal segments 4-6 each with 1 seta which is
as long as or longer than the plate which bears it, the other
seta somewhat variable but never so short as to appear thorn-

^1*<e • . . . IQ

Paratergites of abdominal segments 4-6 with seta otherwise 20
Paratergites of abdominal segments 3-6 with well developed points
at the posterior angles, these points set somewhat anterior to
the posterior margin of their plate, this posterior margin pro¬
jecting posteriorly beyond the points; spiracles noticeably
iarge ; two supposed species which are probably identical- oc¬
curring on shrews in Europe, Africa, and Asia . _

........ - .... . RECLINATA and DELTOIDES

Paratergites of abdominal segments 3~6 with the points at their
posterior angles very small, or perhaps at times lackin'*; pos¬
terior margin of these paratergites not produced; spiracles ex-
tremel.) small and obscure; occurring on Rattus apoensis in the

Philippine Islands . TARSfiMYrn!

D°rsal seta of paratergites of segment 4 about m iin^as the

plate itself, the other seta of these plates and of the plates
of segments 2, 3, S and 6 being much shorter than the plate; a
slender-bodied species which, as far as known, occurs on the
bouse mouse and other closely related species of Mus and on

members of the genus Apodemus in Europe and Asia . SERRATA

U tnerwiSG • ^

. 21

204

21 (20). First abdominal sternite in both sexes quite strongly arcuate and

with its lateral angles somewhat prolonged; occurring on Microtus
and related forms in northern Europe and northern North America

. . ..ALASKENSIS

First abdominal sternite in both sexes not thus; its posterior
margin almost straight and the lateral angles not prolonged.

22 (21). Dorsal lobe of the pseudopenis very short, scarcely one-iourth

the length of the ventral lobe; parameres well developed, ex¬
tending' forward between the posterior arms of the basal plate;
occurring especially ou species ol Rattus throughout the world.

. . . . . SPINULOSA

Dorsal lobe of the pseudopenis equaling about half the length ol
the ventral lobe; parameres quite weakly developed and extend¬
ing forward only slightly past the apex of the arms of the bas¬
al plate; occurring on species of Microtus and related genera
in North America . ABSCISA

Polyplax abscisa Fahrenholz

1925. Polyplax spinulosa (Lurmeister) , Ferris, Contributions Toward a Mon¬
ograph of the Sucking Lice, Part 4:187. (In part; mi sidenti 1 lca-

1938. Polyplax abscisa Fahreuholz, Zeitschrilt liir Paras i tenkuude 10:277,

figures 13, 14. _ QC .0

1942. Polyplax abscisa Fahrenholz, Ferris, Microentomology 7:bb ; tigure 42.

HOSTS AND DISTRIBUTION. Described as from "Arvicola spec., Californien.
This was in all probability a species of Microtus, much less probably ol
Phenacomys. The species has been recorded by Ferris, under the name ol
polyplax spinulosa, from Microtus californicus and Microtus sp. in Caiitor-
nia', from Microtus intermedius from Nevada, from Microtus pennsylvan icus
from New York, United States.

NOTES. This species is exceedingly close to Polyplax spinulosa, appar
ently differing only in certain details of the genitalia ot the male as in¬
dicated in the accompanying key.

Polyplax abyss inica Ferris

1923 Polyplax abyss inica Ferris, Contributions Toward a Monograph ol the
Sucking Lice, Part 4:230; figure 150. „

1940. Polyplax abyssinica Ferris, Werneck, Revista de Entomologica 11:722,

HOSTS AND11 DISTRIBUTION, 'type from Arvicanthis abyssinicus at Bugondo
Te so Uganda, and recorded by Ferris from the same host lrom other local-
ities in U-anda. Recorded by Perris from Otomys tropical t« and Oeno»j/s
bacchante in Uganda. Recorded by Werneck from a subspecies of the type
host at various ^localities in Uganda: from Arvicanthis sp. , and from Mastomys

coucha from the West Nile District in Uganda.

NOTES. If is probable that the normal hosts of this species are species
of the genus Arvicanthis. Ferris described only the female. The male has
been described byr Werneck.

Polyplax alaskensis Ewing

Polyplax alaskensis Ewing, Proceedings of the Entomological Society

of Washington 29:118. , 0

Dniimlnx borealis Ferris, Parasitology 2b: 12/; tigures 1,

Polyplax alaskensis Ewing, Ewing, Proceedings of the Biological bo

ciety of Washington 29:118.

1927.

1933-

1935-

205

H0J1S,AN,D DISTRIBUTION. Described by Ewing from Microtus sp. from un¬
specified locality in Alaska. Described by Ferris from Evotomys sp.

ru%TanusS? lr0D1 Beskenj arrSa-> Finmark, Norway. This host is Clethrionomys
NOTES. It is here accepted that borealis is a synonym of alaskensis.

Polyplax arvicanthis Bedford

1919.

1923.

Polyplax arvicanthis Bedford, Report of the Division of Veterinary
Research, Department of Agriculture, Union of South Africa 5-6 • 716 •
Plate 1, figures. '

Polyplax arvicanthis Bedford, Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 4:227: fi/nires 148 14Q

HOSTS AND DISTRIBUTION. Described as from Arvicanthis pumilio at Onder-
stepoort, Pretoria, South Africa, and later recorded from a subspecies of
this host from Mount Kenya, British East Africa. The correct name for this
host is Rhabdomys pumilio.

Polyplax asiatica Ferris

1923. Polyplax asiatica Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 4:233; figure lfi2D.

HOSTS AND DISTRIBUTION. Type from Crocidura caerulea (a shrew) from
Rangoon, Burma. Also recorded from Nesokia hardwickii from Quetta, Baluch-
istan. Material not previously recorded is at hand from Bandicota bental-

mvvt Akivb’ aD>d fr0in Rattus ooncolor at Bellaiy, Madras, India.

NOTES. The additional material now at hand tends to substantiate the
suspicion, previously expressed by Ferris and by Hopkins, that this species
is normal to a rodent and not to shrews.

Polyplax auricularis Kellogg and Ferris

1915.

1923.

1933.

1938.

Polyp\lx aurlculaiMis Kellogg and Ferris, Anoplura and Mallopha<ra of
Noith American Mammals, Stanford University Publications, Univer-

4 fibre's D° Volume number) ' P^e 13; Plate 1, figure 4; Plate

Polyplax auricularis Kellogg and Ferris, Ferris, Contributions Toward
a Monograph of the Sucking Lice, Part 4:218; figures 140, 141.
Polyplax auricular -is, ^ variety californiae. Fahrenholz, Zeitschrift
fur Paras itenkunde 10:270; figures 19, 21.

Polyplax pa inei Fahrenholz, Zeitschrift fUr Parasitenkunde 10-270-
iitmres 20 22. 1 >

HOSTS AND DISTRIBUTION. Type from Peromyscus maniculatus at Inverness
Marin County California, and recorded from the same host at Yosemite

FoirestirPT^l4ndCalAV°r|IliaV AiSo° rehcorded froir Peromyscus sitchensis from
Forrester Island, Alaska, from Onychomys torridus at Victorville, Califor-

nwirV' m zhomy^lT°taSUr at Colorado Springs, Colorado, and at
Liberal, Kansas. All these records are from within the United States. Al¬
so recorded from Reithrodontomys mexicanus at "Tehontepec, Chiapas " which

liiioobCity, Mexico!’6'’'0’ *" ^

it came actually from this host there must have been

itTnnhp^ Lrood evidence that the members of this family harbor only
lice of the genus Fahrenholzla. There is a Peromyscus maniculatus streatori
and Hopkins suggests that there may have been an error in reading a label
No reason appears for naming the supposed variety. g 1

206

Fahrenheit has aLso named the species Polyplax painet i'rom Peromyscus
caltfornicus from San Mateo County, California, but in the light ol the
available material there seems to be no reason for recognizing this species.

Polyplax biseriata Ferris

1923. Polyplax biseriata Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 4:199; figures 125A, 126.

1938. Eremophthirius btserlatus (Ferris) , Rahreuholz, Zei tschri ft fur Para-
si tenkunde 10:243-

HOSTS AND DISTRIBUTION. From Tatera bohmi at South Guaso Nyiro, British
East Africa, and from Tatera lobengulae at Bothaville, Orange li'ee State,
South Africa.

Polyplax chinensis Ferris

1923. Polyplax chinensis Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 4:209; figures 132, 133-
HOSTS AND DISTRIBUTION. But one record, from Heriones auceps, Shensi,
China. According to Ellerman this is listed as a subspecies of Heriones
ner id ianus.

Polyplax deltoides Fahrenholz

1923. Polyplax reclinata (Nitzsch), Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 4:192; figures 120C, 120G. (Mis-
identification?)

1938. Polyplax deltoides Fahrenholz, Zeitschnft fur Paras 1 tenkunde 10:29b;
figure 12.

HOSTS AND DISTRIBUTION. This species was based by Fahrenholz upon the
specimens recorded by Ferris as from Crocidura coerulea at Rangoon, Burma;
from Crocidura sp . at Atchebal, Valley of Kashmir; Pachyura luzonensis at
Manila, Philippine Islands, and from Scut isorex sp. at "Medjie, this place
name being without other data and perhaps being a misspelling.

NOTES. Since this species name was based entirely upon the records pub¬
lished by Ferris, the type must be selected from among the material record¬
ed by fterris. The type is therefore designated as a female from Pachyura
luzonensis at Manila, Philippine Islands, which is in the Stanford Univer¬

sity collection. , ,

As far as any evidence given by Fahrenholz is concerned, no good reason

appears for regarding this species as anything more than Polyplax reclinata.
But since specimens from the type host of the latter are not available the
name is here accepted.

Polyplax dentaticornis Ewing

1935. Polyplax dentaticornis Ewing, Proceedings of the Biological Society
of Washington 48:20V; figure 2c. . . „

HOSTS AND DISTRIBUTION. Recorded, on the basis ot a single male, irom
Cricetulus andersoni, Shensi, China.

Polyplax eriopepli Ewing

1936. Eremophthirius eriopepli Ewing, Proceedings of the Biological
ety of Washington 48:209. , ,

HOSTS AND DISTRIBUTION. From Eriopeplus incanus Iron Celebes,
host is Cricetulus longicaudatus.

Soci-

This

207

Polyplax gerbilli Ferris

1923. Polyplax gerbilli Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 4:203; figures 128, 129.

1938. Eremophthirius gerbilli Ferris, Fahrenholz, Zeitschrift fur Para-
sitenkunde 10:243.

HOSTS AND DISTRIBUTION. From Gerbillus pyramidum at Khartoum, Egypt.

Polyplax gracilis Fahrenholz

1910. Polyplax gracilis Fahrenholz, Jahresbericht des Nieders&chsischen zo-
ologischen Vereins zu Hannover 2-4:42; text fi<mres 16. 17; Plate
1, figures 10, 11.

19-3 - Polyplax gracilis Fahrenholz, Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 4:234.

1938. Polyplax gracilis Fahrenholz, Zeitschrift fur Parasitenlomde 10:263-
figure 231. ’

HOSTS AND DISTRIBUTION. Recorded from Mus minutus, presumably in Ger¬
many. The host is now known as Micromys minutus.

Polyplax insulsa Ferris

I923. Polypi ax insulsa Ferris, Contributions Toward a Monograph of the
Sucking Lice’ Part 4:231; fig111*63 151, 152A, C.

1929. Ahaematopinus insulsus (Ferris), Ewing, A Manual of External Para¬
sites, page 198.

HOSTS AND DISTRIBUTION. Known only from the original record, from Epimys
\-Rattus) sabanus from Bunguran, Natuna Islands, Malaysia.

1915.

1923.

Polyplax jonesi Kellogg and Ferris

Polypi ax jonesi Kellogg and Ferris, Annals of the Durban Museum 1-
151 ; Plate 15, figures 3-3e.

Polyplax jonesi Kellogg and Ferris, Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 4:216; figures 138 139

HOSTS AND DISTRIBUTION. Known only’ from the origin^ record, fL Sac-
costomus campestris , at Mfongosi, Zululand, South Africa.

Polyplax otomydis Cummin<?s

1912.

1916.

1923.

POlJEl2X otomydis Cummings, Bulletin of Entomological Research 3-
396-7; figures.

P°iAa\al-ir^illL¥^enho^> ¥erris’ kimals of the Durban Museum 1:
240, figures 23C, 24. (Misidentification)

Polyplax cummingsi Ferris, Annals of the Durban Museum 1-240; fig¬
ures 25, 26A.

Polyplax otomydis Cummings, Contributions Toward a Monograph of the
Sucking Lice, Part 4:211; figures 134, 135.

Polyplax cummingsi Ferris, Ferris, Contributions Toward a Monograph
oi the Sucking Lice, Part 4:213; figures 136, 137.

HOSTS AND DISTRIBUTION. Described by Cummings from Otomys Irroratus
from Mount Kenya, British East Africa. Also recorded from this host from
Onderstepoort, Pretoria, and from Mfongosi, Zululand, South Africa. Re¬
corded from Otomys angoniensis from Naivasha, British East Africa, and from
Otomys bran t s i without indication of locality, this last host now bein<r re¬
ferred to Parotomys. Recorded by Ferris as Polyplax cummingsi from Datymys
Incomtus from Mfongosi, Zululand, and from Kaimosi, British East Africa and
from Rattus (now known as Aethomys) chrysophilus from Mfongosi, Zululand.

208

NOTES. A re-examination of the material at hand reveals no sound basis
for the recognition of the species cummtntsl, the characters given b> Ferris
for the separation of the latter species now appearing to have been quite
illusory .

Polyplax oxyrrhyuchus Cummings

1915- Polyplax oxyrrhynchus Cummings, Proceedings of the Zoological Soci¬
ety of London, page 251; text figures 4-6, 8, 9, 11. 13-

1923. Polyplax oxyrrhynchus Cummings, Ferris, Contributions Toward a Mono¬

graph of the Sucking Lice, Part 4:225; figures 146, 147.

1938. Polyplax oxyrrhynchus, variety hystrellae Fahrenholz, Zeitschnft
fQr Paras itenkunde 10:275*

HOSTS AND DISTRIBUTION. Type from Acomys cahirinus at Assiut, Egypt.
Recorded by Ferris from Acomys hystrella from Numule, Uganda, and irom

Acomys perctvalt , British East Africa. . .

NOTES. Fahrenholz, apparently without seeing any specimens and working
merely from the illustrations accompanying the original description ol this
species and the later record by Ferris, assumed to establish the supposed
,lvarietv" hystrellae for the specimens illustrated by Ferris. The type ol
this naine will therefore be among these specimens. The differences cited
by Fahrenholz for his "variety" are merely such as are inevitable in illus¬
trations made by different persons from specimens which were dillerently
prepared and this supposed variety is here reduced to synonymy.

Polyplax phthisica Ferris

1923. Polyplax phthisica Ferris, Contributions Toward a Monograph ol the
Sucking Lice, Part 4:223; figures 144, 145. .

HOSTS AND DISTRIBUTION. Type from Lophuromys aquilus trora Ngani Narok
River and from the same host species (recorded as zena ) from Molo, British
East Africa; from Lophuromys sikapusl, Rhino Camp, Uganda; from Lophuromys
sp. from Nyasaland. A record from Thamnomys ibeanus, British East Alnca,
is probably due to contamination.

Polyplax praecisa (Neumann)

1901. Haematopinus praecitus Neumann, Archives de Parasitologie ?-600. (In

part; also typographical error for praecisus, later corrected by

1902. Haematopinus praecisus Neumann, Archives de Parasitologie 6:144; fig.

1904. Polyplax praecisa (Neumann), Enderlein, Zoologischer Anzeiger . 3

1919. Polyplax praecisa (Neumann), Fahrenholz, Jahresbericht des Nieder-
s&chsischen Zoologischen Yereins zu Hannover 5-lU:Zh.

1923. Polyplax praecisa (Neumann), Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 4:196; figure 123. f f p

1938. Eremophthirius praecisus (Neumann), Fahrenholz, Zeitschnf u

HOSTS AND DISTRIBUTION. Recorded as from "gros rats, in Abyssinia. Hop-
kins^notes that T^tera nlgri cauda occurs in Abyssinia and assumes the spe-

^^OTES^^ Neumann included two distinct species in his praecisus. This was
• ' 1 vv Fahrenholz who in 1919 named one of the two as Hoplopl eura
th"Acted the Le to the other , -hic^is a »
Fprric; saw the type specimens and confirmed this procedure, r ye
praecisa is still known only from the female and without precise host desig

nation.

209

Polyplax reclinata (Nitzsch)

1864. Pediculus reclinatus Nitzsch, Zeitschrift fiir den gesamten Naturwis—
senschal’ten 25:23.

1874. Haematopinus reclinatus (Nitzsch), Giebel, Insecta Epizoa, page 37.
1904. Polyplax reclinata (Nitzsch), Enderlein, Zoologischer Anzeiger 28:
142 ■

1910. Haematopinus ( Polyplax ) reclinatus (Nitzsch), Neumann, Archives de
Parasitologie 13:524; text figure 24.

1912. Polyplax rec l inata (Nitzsch) , Fahrenholz, Jahresbericht des nieder-
sachsischen zoologischen Vereins zu Hannover 2-4:37; text figures
11, 12; Plate 1, figures 12, 13; Plate 2, figures 2, 4; Plate 3,
figure 7.

1923. Polyplax reclinata (Nitzsch), Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 4:192. (In part)

1932. Polyplax reclinata, variety leucodontis Jancke, Zeitschrift fiir Par—
asitenkunde 4:525; figure 2.

1938. Polyplax reclinata (Nitzsch), Fahrenholz, Zeitschrift fiir Parasiten-
kunde 10:254; figures 9, 10, 11.

HOSTS AND DISTRIBUTION. Originally recorded from the European shrew
Sorex araneus, and since recorded a few times from this host. Also record¬
ed, as variety leucodontis, from another European shrew, Crocidura leucodon .

NOTES . Ferris has previously referred to this species specimens from
various Asiatic shrews, but Fahrenholz has considered these to represent a
distinct species to which he has applied the name deltoides. Also -Jancke
has recorded material from a European shrew, Crocidura leucodon, as a vari¬
ety of reclinata. In the absence of specimens from the type host it is not
here possible to come to any deiinite decision regarding this variety. On
the basis of the published notes and descriptions by Jancke and Fahrenholz,
the present writer still believes all these to represent a single species
for which the name reclinata is correct. The supposed "variety" leucodontis
is here placed in synonymy with reclinata, but the species name deltoides
is admitted, in the absence of the material which is necessary before it is
definitely reduced to synonymy.

Polyplax serrata (Burmeister)

J§39. Pediculus serratus Burmeister, Genera Insectorum, Rhynchota, No. 6.
1842. Haematopinus serratus (Burmeister), Denny, Monographia Anoplurorum
Britanmae, page 36.

1904. Polyplax serrata (Burmeister), Enderlein, Zoologischer Anzeiger 28:

1912. Polyplax affin is (Burmeister) , Fahrenholz, Jahresbericht des nieder-
sachsischen zoologischen Vereins zu Hannover 2-4:39; figures 13-15.
(Misidentification) ’

1923. Polyplax serrata (Burmeister), Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 4:191; figure 120B, E.

1932. Polyplax serrata (Burmeister), Jancke, Zeitschrift fur Parasiten-
kunde 4:252.

1938. Polyplax affinis Fahrenholz, Zeitschrift f(ir Paras itenkunde 10-261.
(As a new species)

HOSTS AND DISTRIBUTION. Originally described from the house mouse, Mus
musculus, in Europe. Later recorded as Polyplax affinis (Burmeister) from
Apodemus syluaticus in Europe. Ferris has recorded it from this host from
Switzerland, Bohemia, and England; from Apodemus airarius from Manchuria
and China; from Apodemus speciosus from China; from Nus spicilegus, which
is merely a subspecies of musculus, from Spain; and from Nus musculus from
Scotland and England.

210

NOTES. The species described as Pediculus af finis by burmeister was at
one time placed in Polyplax but is now regarded by both Kali renholz and
Perris as be in;' a Hoplopleura. A species of Polyplax which has been misi-
dentified as afflnis is considered by Perris to be the same as Polyplax
serrata. Fahrenholz (193^), however, has maintained that this species is
distinct and has described it as new, giving affints as a new name lor it.
Under the revised Rules (bulletin of Zoological Nomenclature 4:97-125. 1950)
this name is available, since at the time of its proposal (193&) if was a
nonconcurrent secondary homonym.

In connection with the present work the material at hand lrom Hus muscu-
lus and the various species of Apodemus has been carefully re-examined.
The opinion is here maintained that there is absolutely nothing in these
specimens which will justify a specific separation. Polyplax afflnis
Fahrenholz is therefore placed as a synonym of Polyplax serrata (burmeister).

Polyplax spinigera (burmeister)

1S39. Pediculus spiniger burmeister. Genera Insectorum, Rhynchota, No. 9-
1909. Haematopinus ( Polyplax ) splniger (burmeister) , Neumann, Archives de
Paras i to logie 13:524; figure 24.

HOSTS AND DISTRIBUTION. Described from a rodent cited by the older au¬
thors as Hypudaeus amphibius, which is now referred to the genus Arvicola.

NOTES. Since its original description only one author, Neumann, has
seen specimens from this host and the species cannot be definitely identi¬
fied from his notes and figures. It is apparently distinct from Polyplax
spinulosa and the suggestion that it is the same as, and there! ore ante¬
dates, either Polyplax alaskensis Ewing or Polyplax abscisa Fahrenholz will
need to be explored when specimens are obtained from its type host.

Polyplax spinulosa (Burmeister)

Figures 90, 91

1839. Pediculus sptnulosus Burmeister, Genera Insectorum, Rhynchota, No. 8.
1842. Haematopinus spinulosus (Burmeister) , Denny, Monographia Anoplurorum
britanniae, page 26; Plate 24, figure 5.

1864. Pediculus denticulatus Nitzsch, Zeitschrift fur den gesamten hatur-
wissenschaften 23:24. .

1905. Polyplax spinulosa (Burmeister), Enderlein, Zoologischer Anzeiger

28:142.

1923. Polyplax spinulosa (Burmeister), Ferris, Contributions Toward a Mon¬
ograph of the Sucking Lice, Part 4:187; figures 119, 120A, D, F, H.
1929 Polyplax praomydis Bedford, Annual Report of the Director ol Veteri¬
nary Services, Union of South Africa 15:503; figure3 2-5- .

1938. Polyplax spinulosa (Burmeister), Fahrenholz, Zeitschrift. fur Para
sitenkunde 10:249; figures 1-8, 23c. . .

1945. Polyplax campylopteri Zavaleta, Anales del Instituto de Biologia

HOSTS SdXDISTOIBUTION. Originally described from the brown rat, Rattus
(=Mus = f!pimi/s) norveiicus, in Europe. Later many times recorded from this
host and from Rattus rattus and some of its subspecies in many' parts ol the
world. It has been recorded by Ferris from Rattus calc is, which is consid¬
ered to be exulans, in the Philippine Islands and Rattus strtdens\n the
Malay Peninsula and from Bandicota (as Gunomys ) benialensis in Burma. De
scribed by Bedford as Polyplax praomydis from Praonys (now called .hallomys)

namaouensis at Onderstepoort, South Attica. . _ . .

Records by Ferris of the occurrence of this species upon various specie-
of Hicrotus Synaptomys, and Phenacomys are erroneous and are here trans¬
ferred to Polyplax abscisa Fahrenholz.

211

Polyplax spinulosa (Burmeister)

Figure 90

212

Polyplax spinulosa (Burmeister), details

Figure 91

213

NOTES. A male from the type lot of Polyplax praomydis Bedford is avail¬
able and is very precisely spinulosa. Polyplax campyloptera Zavaleta, de¬
scribed as from a bird, is, on the basis of illustrations presented by its
author, very obviously nothing more than spinulosa.

Polyplax stephensi (Christophers and Newstead)

1906. Haematopinus stephensi Christophers and Newstead, Thompson, Yates
and Johnston Laboratories Report (new series) 7:3; Plate 1.

1923. Polyplax stephensi (Christophers and Newstead) I, Perris, Contributions
Toward a Monograph of the Sucking Lice, Part 4:206; figures 130,
131 •

1938- Eremophthirius stephensi (Christophers and Newstead), Fahrenholz,
Zeitschrift fur Paras itenkunde 10:243.

HOSTS AND DISTRIBUTION. First described from Gerbillus (-Tatera) indica
from India, without precise indication of locality. Cotype specimens are
labeled as from Madras. Later recorded by Ferris from the same host from
various localities in India.

Polyplax tarsomydis Ewing

1935- Polyplax tarsomydis Ewing, Proceedings of the Biological Society of
Washington 48 : 206 .

1938. Eremophthirius tarsomydis (Ewing), Fahrenholz, Zeitschrift fur Para-
si tenkunde 10:243.

HOSTS AND DISTRIBUTION . Described by Ewing as from Tarsomys (a subgenus
of Rattus) apoensis from the island of Mindanao, Philippine Islands. = One
male taken from a skin in the United States National Museum. Two females
which may be considered to belong to this species are at hand frcm the same
host, taken from a skin, United States National Museum Number 144616, from
the summit of Mount Bliss, on the island of Mindanao.

NOTES. This species was described from a single male. It is not prac¬
ticable here to describe the female except as it is included in the key to
the species.

Polyplax taterae Ferris

1923. Polyplax taterae Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 4:198; figures 124, 125D.

1936. Polyplax subtaterae Bedford, Onderstepoort Journal of Veterinary
Science and Animal Industry- 7:63; figure 7.

1938. Eremophthirius taterae (Ferris), Fahrenholz, Zeitschrift fur Para-
si tenkunde 10:243.

1938. Eremophthirius subtaterae (Bedford), Fahrenholz, Zeitschrift fur
Parasitenkunde 10:243.

HOSTS AND DISTRIBUTION. TyPe of taterae from Tatera vicina from Mount

Rukenya, British East Africa. Type of subtaterae from Tatera liodon at

Kampala, Uganda.

NOTES. Specimens of subtaterae are at hand, through the kindness of the

late G. A. H. Bedford, and it is upon these that the above synonymy is based .

Polyplax waterstoni Bedford

1919. Polyplax waterstoni Bedford, Report of the Division of Veterinary
Research, Department ot Agriculture, Union of South Africa 6—7*
715: Plate 1, figures 1, 2, 4, 6.

1923. Polyplax waterstoni Bedford, Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 4:193; figures 121, 122.

214

1938. Polyplax emlnatus Fahrenholz, Zeitschrift t’iir Parasiteukunde 1U:266;
Figures 16, 17.

HOSTS AND DISTRIBUTE ON . First described as from "several rats," at On
derstepoort, Pretoria, South Africa. Recorded by Ferris from Eptmys pero-
myscus, for which the proper name is apparently Rattus tullbergi, at Moio,
British East Africa. Described as Polyplax emlnatus by Fahrenholz from
"Paderorycte s tadat, " from East Africa. No such host name appears in any
available Lists. Hopkins has hazarded the guess that this is a corruption
of Tachyorycte s audax, but this is purely a surmise unsupported by any spe¬
cial evidence.

NOTES. On the basis of the description and illustrations given by Eah-
renholz, there is not the slightest justification for the separation of
Polyplax emlnatus from waterstoni .

Polyplax werneri (Glinkiewicz)

1907. Eremophthirius werneri Glinkiewicz, Si tzuugsberichte der mathemat-
i sch-n atu r wi sse nschaftlichen CLasse der kaiserliche Akademie der
Wissenschaften zu Wien 116:381; figures.

1923. Polyplax werneri (Glinkiewicz), Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 4:202; figure 127.

1938. Eremophthirius werneri Glinkiewicz, Fahrenholz, Zeitschrilt liir Par-
asitenkunde 10:243*

HOSTS .AND DISTRIBUTION . Described from Pachyuromys duprasi. Natron Val¬
ley, Egypt, and recorded by Ferris from the same host and locality.

NOTES. This species is the type of the genus Eremophthirius.

Genus PROEN DERLEIN ELLUS Ewing

1923. Proenderleinellus Ewing, Journalot the Washington Academy ol Sciences

13* 147.

1938. Vaterstonla Fahrenholz, Zeitschrift fur Parasitenkunde 10:244. (Pre-

1939. Symysadus Fahrenholz, Mitteilungen aus dem entomologischen Verein

Bremen (no volume number), page 44. . .

GENERIC TYPE. The genus was based upon Proenderleinellus africanus
Ewing, which is a synonym of Polyplax calva Waterston. The generic names
proposed by Fahrenholz were based upon Polyplax calva Waterston. This is

the only included species. . , , . . .

CHARACTERS. Polyp lacinae in which the antennae are live segmented, not
sexually dimorphic. .Anterior legs moderately large, but with slender claw;
middle legs but slightly' larger than the first and with slender claw; third
legs much larger than the others and with stout claw. Paratergal plates ol
the abdomen present on segments 2-8, those of segment two divided longitu¬
dinally into two pieces. Abdomen entirely membranous in both sexes excep
for the usual sclerotizations of the terminal and genitalic segments and
even these are much reduced. Abdominal segments ot the female, both dor
sally and ventrally, with for the most part two rows ol setae, these setae
borne each on a small, sclerotized plate or tubercle which surrounds its
base. Abdomen of the male with one row ol such setae on all segment^ or-
sally except perhaps the second, which has two, and with two row., on most
of the segments ventrally. Sternal plate of the thorax well developed.

AM5"0Tb' S ll^'wrYteT',6 Ssuspec t, ing* th\t p"roe nderleinellus .frlcanue Ewing
might be a synonym of the earlier Polyplax calva Waterston enlisted the
aid of Mr. C. F. W. Muesebeck, who examined the single male type ol the

former and agreed.

215

Proenderleinellus calva (Waterston)

Figures 92, 93

1917. Polyplax calva Waterston, Parasitology 9: 199; figures.

1923. Polyplax calva Waterston, Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 4:234; figures 153, 154.

1923. Proenderleinellus africanus Ewing, Journal of the Washington Academy
of Sciences 13:147.

1938. Raterstonia calva (Waterston), Fahrenholz, Zeitschrift fur Parasiten-
kunde 10:244.

1938. Vaterstonia calva zanzibariensis Fahrenholz, Zeitschrift fur Para-

sitenkunde 10:244.

1939. Symysadus calva (Waterston), Fahrenholz, Mitteilungen aus dem ento-

mologischen Verein Bremen (no volume number) , page 44.

HOSTS AND DISTRIBUTION. Described by Waterston from Cricetomys iambianus
at Accra, French West Africa, and also recorded from Cricetomys sp. from
Zanzibar. The supposed subspecies zanzibariensis was based solely upon the
illustration given by Waterston of the male, this male having come from
Zanzibar. The supposed species africanus of Ewing was based upon a single
male which had been taken from a skin of Thryonomys £regorianus which Rad
come from British East Africa. There are at hand specimens from the type
host at Accra and also from Cricetomys from Zanzibar.

NOTES. The specimens at hand deny the slight differences offered by
Fahrenholz as a basis for the naming of his supposed form zanzibariensis.

Genus RATEMIA Fahrenholz

1916. Ratemia Fahrenholz, Archiv fur Naturgeschichte, Abteilung A, 81:11:31.
1922. Ratemia, Ferris, Contributions Toward a Monograph of°the Sucking
Lice, Part 3:156.

1949. Ratemia, Webb, Proceedings of the Zoological Society of London 119:
164.

GENERIC TYPE. Haematopinus (LinoPnathus) squamulatus Neumann, the only
included species.

CHARACTERS. Polyplacinae with five-segmented antennae which are not
sexually dimorphic. Anterior legs relatively small, with slender claw;
middle and posterior legs about equal to each other, larger than the first
and with somewhat stouter claw. Paratergal plates of the abdomen occurring
on segments 3-6 • Abdomen otherwise membranous in both sexes except for the
usual terminal and genital areas. Abdominal segments in both male and
female, both dorsally and vent rally, with a single transverse row of setae
which along the mid-line of the body becomes merged with a median cluster
of two or three rows of setae. Spiracles present on abdominal segments 3-9.
Thoracic sternal plate well developed.

The type species of the genus apparently occurs normally on hosts of the
Perissodactylous family Equidae.

NOTES. Because of the hosts of the type species it is difficult to
shake off a prejudgment that this genus should not be assigned to the Hop-
lopleuridae, which are primarily parasites of the rodents and there is a
pressure to find some excuse for placing it in the Haematopinidae or the
Linognathidae. However, no morphological reason for eithur of these latter
assignments appears and it is here placed in the Hoplopleuridae. Webb
(1949) has assumed the genus to be related to Linognathus because of simi¬
larities ol the spiracLes, but the present writer remains unimpressed by
this supposed resemblance, especially in the face of other morphological
features.

216

Ratemia squamulata (Neumann)
Figure 94

1911.

1916.

1922.

1946.

j ematopinus (Linot*nathus) squamulatus Neumann, Archives de Para-
sitologie 14:401; figures.

itemia squamulata (Neumann), Fahrenholz, Archiv fur Naturgeschichte,

Abteilung A, 81:11:31- . T , M

itemla squamulata (Neumann), Ferns, Contributions Toward a Mono

rrraph of the Sucking Lice. Part 3:167; figure 103-
-Hernia squamulata (Neumann), Hopkins, Annals and Magazine oi Natural
Historv (Series 11) 12:666. (Dated August, 1946, but available
copy bears the notation that it was published in May, 194b)

217

pai atei gal plates female genitalia

thoracic sternal plate

Proenderleinellus calva (Waterston), details

Figure 93

HOSTS AND DISTRIBUTION. Originally described from an undetermined host
f> Dl*je_I)a°Va» Abyssinia. Since that time Hopkins has recorded the species
from domestic donkey at Liro, Lango District, Uganda, and from Burchell's

gZlS ^U.rls) burchell l , taken on the Athi Plains near Nai robi!
K,. uh aPPar®n1;ly establishing the normality of the species to members
of the Perissodactylous family Equidae. It should, therefore, enter into
anj list of the parasites of domestic animals.

NOTES. Through the kindness of Mr. Hopkins it has been possible to see
IT fr°: b0fVhc hosts recorded by him. In the accompanying illus^
ha nihPh i° temaie 15 lr0m a sPecimen in the original type lot:

as a !!an Sr!SHoprHn“s!PeC1"Cn ‘'r°" ^ bur^“1’ ^ ™ received

218

male genitalia

female genitalia

spiracle

• * vl /

ii A ^

I* 1

Ratemia squamulata (Neumann)

Figure 94

219

Genus SYMOCA Fahrenholz

1938. Symoca Fahrenholz, Zeitschrift fiir Parasi tenkunde 10:245.

GENERIC TYPE. Polyplax brachyrrhynchus Cummings, the oniy included spe-
cies.

CHARACTERS. Polyplacidae with five-segmented antennae which are sexual¬
ly dimorphic, those of the malq having the distal, preaxial angle of the
third segment slightly prolonged and hearing a stout, recurved seta at its
apex. Head with the basal segment of the antennae set very close to the
truncate anterior margin. Thorax narrow and somewhat elongate, with no
distinct sternal plate. Anterior legs small and weak and°with slender
claw, middle legs distinctly larger and with somewhat larger claw; posteri¬
or legs with the tibio-tarsus much enlarged and with a stout, heavy claw.
Abdomen with paratergal plates present on segments 2-8, the plates on seg-
ment two apparently not divided longitudinally. Female with abdominal seg¬
ments 4r- 1 each with two plates and two transverse rows of setae both dor-
sally and ventrally . Male with but one row of setae on any segment both
dor sally and ventrally, except for the sternum of segment three which ap¬
parently has two rows. Sternal plates not developed in the male.

NOTES. While the relationships ot this genus are certainly with Polyplax
and the bases for a generic separation are not especially convincing the
genus is here accepted.

Symoca brachyrrhyncha (Cummings)

Figures 95, 96

1915. Polyplax brachyrrhynchus Cummings, Proceedings of the Zoological So¬
ciety of London, page 246; figures 13, 14.

1923. Polyplax brachyrrhynchus Cummings, Ferris, Contributions Toward a
Monograph of the SuckinS Lice, Part 4:220; figures 142, 143.

1938. Symoca brachyrrhyncha (Cummings), Fahrenholz. Zeitschrift f&r Para—

sitenkunde 10:245.

1939. Symoca brachyrrhyncha, variety minor Fahrenholz, Mitteilun.ren aus

dem entomologischen Yerein Bremen, page 32.

HOSTS AND DISTRIBUTION. Described as from Acomys cahirinus at Assiut
Egypt. Recorded by Ferris from Acomys hystrella in Uganda and from Acomus
perctvali in British East Africa.

NOTES. Fahrenholz based the supposed variety minor entirely upon a com¬
parison of the description and illustrations given by Cummings with those
|Jve° v Ferns, although Ferris had compared his specimens with the types,
pie differences cited by Fahrenholz are merely such as may readily appear
m work of two individuals, even utilizing the same material.

Family LINOGNATHIDAE Webb

1904. Trichaul inae Enderlein, Zoologischer Anzeiger 28:138.

1905. Linoinathinae Enderlein, Zoologischer Anzeiger 29-194.

1946. Linojnathidae Webb, Proceedings of the Zoological Society of London
116 : 107 .

DESCRIPTION OF THE FAMILY. Anoplura in which external evidence of eyes
m.iy or may not be present. Abdomen entirely membranous in both sexes ex¬
cept for plates associated with the terminal and genital segments and rare¬
ly (one species) tergal plates in the male. Abdominal segments each with
at least one row of hairs both dorsally and ventrally, usually much haired.
Second and third legs almost always equal to each other and much larger
than the first pair. Thorax usually without any trace of a sternal pllte
but ll present always without the apex free from the body. Spiracles as

220

Symoca brachyrrhyncha (Cummings'

Figure 95

far as known, with distinct internal ledges which show externally as rings,

or with the ledges forming internal points.

The members of the family are restricted to members of the Orders Artio
dactvla and Hyracoidea, except for two species from Carnivora.

NOTES. To" this family there are here assigned four genera— Lino$nat hus,
Microthoraclus, Solenopotes, and Prollnoinathus. A certain ajnoun^ of
is felt concerning the genus Microthoraclus since an •argument can be made
for its assignment to the Haematopinidae. One other question, that ol t
lelns H^mdlpsus, is troublesome. The type of this genus is unfortunately
an° imperfec t ly known species and, because of

SDecies which are assigned to the genus, it is here placed in the Hoplo
nleuridae Taken by itself it might very properly be assigned to the
Linognathidae and eventually it may be necessary’ to break j^P^-enus
Haemod ipsus, assigning its type species to a position near Linotnathus.

221

antenna

abdominal seta

f emale genitalia

Symoca brachyrrhyncha (Cummings), details

male genitalia

Figure 96

Key to the Genera of LINOGNATHIDAE

1. Eyes indicated externally by a definite lens; occurring on Camelidae

u . . . .

No external trace of eyes . o

2. Abdominal spiracles borne in a slightly sclerotized tubercle which’ pro¬

jects at least slightly from the body; occurring chiefly on Cervidae,

but 1 species known from domestic cattle . SOLENOPOTES

Spiracles not thus borne in sclerotized tubercles . 3

3. Setae of abdomen greatly reduced in number, there being normally but *2

setae in each median group, both dorsally and ventrally, on any ab¬
dominal segment, these in a single row; spiracles extremely small;
occurring as far as known only on members of the Order Hyracoidea _

. . prolinognathus

Setae ol abdomen more numerous, usually abundant, and in 2 or more rows
on each segment, spiracles not minute; occurring for the most part on
members of the Order Artiodactyla, but represented on the family
Canidae of the Order Carnivora . LINOGNATHUS

222

(k'lius L1N0GNATHUS Enderlein

1904. Tr Ichaulus Enderlein, ZooLogischer Auzeiger 28:139, 141. (Preoccu¬

pied)

1905. Ltno£nathus Enderlein, Zoologischer Anzeiger 29:194.

1909. Ltnotnathus (Haematopinus) , Neumann, Archives de Parasi to logie 13:529.
1932. L tnognathus , Ferris, Contributions Toward a Monograph of the Sucking
Lice, Part 5:336.

1949. Stobbella Kichier, Bolletino della Societa Entomologies Italians

79:13-

GENERIC TYPE. Pedlculus plllferus Burmeister, which is considered to be
a synonym of Pediculus setosus von Olters.

GENERIC SYNONYMS. Stobbella Eichler, type Linoinathodes plthodes Cum-

raings .

CHARACTERS. Linognathidae in which there is no external evidence ol
eyes. Antennae five-segmented, the fourth and fifth segments not lused.
Spiracles of the abdomen usually more or less spherical, the internal ledges
of the atrium appearing externally as partial rings; never elongated and
never borne in a sclerotized tubercle. Thoracic sternal plate lacking, or
it' present very weakly developed and divided longitudinally into two small
plates. Abdominal segments usually much haired both dorsal ly and ventral-
ly, with the hairs in at least two rows. Abdomen terminally with a pair ol
ventral lobes, but these never produced into flat processes. Genitalia oi
the male always with pararoeres well developed and enclosing the oedeagus.

NOTES. This isagenus of considerable size, 26 species now being known.
These are confined to the families Bovidae and Giralfidae of the Order
Artiodactyla, except for two species which occur on carnivores of the iam-

' Eichler has recently named the genus Stobbella, with Linotnathus plthodes
as its type, but this genus is here rejected.

Key to Species of LINOGNATHUS

1. Abdomen beset with short, stout, more or less fusiform setae . 2

Abdomen with no such fusiform setae . • . . • "

2 (1). Anterior legs of same form as the others and almost as large, ab¬

domen very thickly and uniformly beset with fusiform setae ; on

Gorton taurinus in Africa . . . . . . .bPILATUS

Anterior Legs definitely smaller than the others and with slender
claw; fusiform setae more or less numerous, but leaving bare

are as ..»••••••••••■••••••• •••**"**********#*********************~^

3 (2). Gonopods of female well separated mesally from each other ; on

ope cervicapra in India . _ . ;; . PITHUUEb

Gonopods of female very broad and leaf-like, meeting each other

4 (3). Median genital plate of female expanded at its posterior end, on

Gaze 1 1 a thomsoni in British East Africa . . . ...LLWlbl

Median genital plate of female not broadened at its posterior ex
tremity; on Antidorcas marsupialls in South Africa. ..... .BEDFORDI

6 (1). Female with no trace of a sclerotized, median, genital plate . .

Female with at least a small, median, genital plate . l‘’n

6 (5). Gonopods of the female apically truncate, emarginate or serrate.../

Gonopods of the female apically rounded or acute . . . -9

7 (6). Gonopods of the female with the posterior border somewhat emargi¬

nate and bearing a sclerotized hook at the mesal angle; torehead
elongate and apically acute, hindhead longer than wide, almost
rectangular; occurring on domestic cattle throughout the world^

223

8 (7).

9 (6).

10 (9).

11 (5).

12 (11).

13 (12).

14 (13).

15 (14).

16 (11).

17 (16).

Not with this combination of characters . 8

Head long and slender, the forehead acute, the hindhead with the
lateral margins slightly convex; gonopods of female with a slight
tooth close to the posterior border near the mesal angle; occur¬
ring especially on domestic goats, but also at times on sheep and

recorded from the European Capra ibex and Caprella rupicapra .

. STENOPSIS

Head very small and short, not prolonged in front of the antennae;
gonopods of the female with the posterior margin irregularly ser¬
rate; widely distributed on domestic sheep . PED.4LIS

Head elongate and slender; dorsum of metathorax with an apically
free lobe close to the margin just above the posterior coxal con¬
dyle; occurring on Cervicapra in Africa . FAHRENHOLZI

Head short and broad, the anterior margin broadly rounded, the fore¬
head not at all prolonged anteriorly . 10

Abdomen in both sexes with very few elongate setae, there being a
median pair and a single seta near the margin, both dorsally and
ventrally, in either sex, with a few extremely minute setae be¬
tween the median and lateral setae; occurring on Gorgon taurinus

in Africa . '.HOLOGASTRUS

Abdomen with numerous slender setae both dorsally and ventrally in
both sexes; occurring on Canis brasiliensis in South America....

. : . . . TAENIOTRICHUS

Genital plate of the female oval, triangular, or lozenge-shaped,
and never with a slender median, stalk-like prolongation con¬
necting with the lip of the vulva . ?, . 12

Genital plate of the female with various configurations, but not

as above . 16

Genital plate relatively very large and definitely lozenge-shaped;
head unusually elongate and narrow; occurring on "North African

antelope " . PETASMATUS

Not with this combination of characters . 13

Female with the genital plate transversely oval and with a slight
median, posterior point; gonopods elongate, apically somewhat
spatulate; male with the terminal sternite produced and with a
flattened, sclerotized lobe on each side; occurring on Conno-

chaetes and Gorgon in Africa . GNU

Not with this combination of characters . 14

Head elongate and slender, anteriorly acute; genital plate of the
female forming an almost equilateral triangle; occurring on

Hippotragus in Africa . HIPPOTRAGI

Head broad throughout, genital plate of the female forming merely

a small, slightly elongate, oval area . 7 . 15

Head very short and broad, scarcely longer than wide; occurring
on domestic dogs and on foxes; occasionally taken from other

carnivores . . SETOSUS

Head broad, but fully twice as long as wide; on domestic sheep in

various parts of the world . OVILLUS

Female with a pair of small, hook-like processes on the lip of
the vulva between the gonopods; male with a deeply pigmented,
transverse plate just in front of the anus; occurring on Tauro-

tragus in Africa . TAUROTRAGUS

Not with these characters . .

Female with the genital plate definitely spatulate (not merely
slightly expanded anteriorly) and with a slender, stalk-like
posterior prolongation nearly or quite connecting with the lip

of the vulva . .

Female with the genital plate not thus spatulate . 21

224

20 (19).

21 (17)
22 (21)

18 (17). Genital plate of the female described as more or less T-shaped,

the arms of the T uuite short; occurring on Peleus capreolus in

Africa . • . PELEUS

Genital plate of the female otherwise . 19

19 (18). Gonopods of the female curved, their apices directed toward the

mid-line of the body and with setae extending from the lateral
margin around the apex and well up the mesal margin; head rather
smoothly fusiform; occurring on Cephalophus maxwelli, an African

species . .

Gonopods of the female almost straight and parallel, their setae

confined to the mesal margin . . . ...20

Gonopods with mesal margin from apex to j unction with body straight,

occurring on Aepyceros melampus in Africa....... . AEPYCEKUS

Gonopods with mesal margin from apex to j unction with body convex,

occurring on giraffe . BREVICORNIS

Gonopods of female with setae confined to the apex . 22

Gonopods of female with setae extending well up the mesal margin

from the apex . ;:*••••*•* • * * :

Gonopods of the female apically convergent; genitalia ol male with
definite pseudopenis and with parameres not basally swollen.. 2J
Gonopods of female described as parallel; genitalia ol male de¬
scribed as having the parameres basally swollen and as lacking
pseudopenis; occurring on Antilope cervtcapra, an Indian spe-
. . CERVICAPRAE

22 (22). Gonopods of female extremely small, strongly convergent through¬
out their length; occurring on Antilope euchore and various

other species of Ant Hope in Africa . TIBIALIS

Gonopods of female with sc lerotization elongate and strongly
curved at the extreme apex; occurring on domestic sheep, widely

distributed, and perhaps on Oreotragus saltator in Africa . .

. AFRICANUS

24 (21). Forehead slender and apically acute; lateral margins of the hind-

head smoothly convex; gonopods of the female short and quite
broad; genital plate of the female elongate, somewhat expanded
anteriorly but scarcely to be described as spatulate; occurring

on Tragelaphus sylvatlcus in Africa . FRACTUS

Forehead truncate or at the most broadly rounded....... .

25 (24). Gonopods of the female with their mesal margins straight, parallel

and only these margins bearing setae; genital plate of the te
male shoe sole-shaped, projecting slightly from the vuLva api

oally ; known from Tragelaphus gratus in Africa . LIMNUIKAbi

Otherwise, occurring especially on species of Cephal°phus

Linognathus aepycerus Bedford

1936. Linognathus aepycerus Bedford, Onderstepoort Journal of Veterinary'
Science and Animal Industry 7:62; figures 5> o. ,

HOSTS AND DISTRIBUTION. From Aepycerus melampus, commonly called impaia,
between Pretoria and Johannesburg, South Africa.

Linognathus africanus Kellogg and Paine
Figures 97, 98

1911.

1916.

nognathus africanus Kellogg and Paine, Bulletin of Entomological

Rpc;parch 2-146; Plate 4, figures 1, 5* . „ , .

nognathus stenopsis (Burmeister) , Ferris Proceedings of the Cali
fornia Academy of Sciences (Senes 4) 6:155- (Part, misidentit

cation) 22f.

Figure 97

female penitalia

Linognathus africanus Kellogg and Paine, details

Figure 98

1927. Linognathus stenopsts (Burmeister) , Bedford, ReP°rt °f Director
•0f Veterinary Education and Research, Union ol South Africa 11 12.

737. (Part; misidentification) A ._ .. - .

1932. Linognathus africanus Kelloggand Paine, Ferris, Contri buti ons Tow

a Monograph of the Sucking Lice, Part 5:83 ; tigures 212B, G, E, 21J.
1932. Linognathus africanus Kellogg and Paine, Bed lord, Report ol the D
rector of Veterinary Services and Animal Industry, Inion ol South

1939. Linognathus africanus Kellogg and Paine, Fahrenholz, Mitteilungen
aus dem entomologischen Verein, Bremen, page 3.-
HOSTS ANT) DISTRIBUTION. Type from "sheep," Abeokuta, southern Nigeria,
AfrfS ulr SSriSfr- domestic sheep and domestic goats fr-jmj-
localities in South Africa, Abyssinia, India, and the K'

nnrded bv Fahrenholz from Ovis longipes from Algeria, North Africa.

NOTES There are apparently four species of I inognat hu$ namely, ov 1 1 1 us,
Dfidal is stenopsis, and africanus— which may be encountered on domestic
sheep and goats. They are quite distinct species and the probability is
that here has been some interchange of hosts.

227

Linognathus angulatus (Piaget)

1885* Haematopinus ungulatus Piaget, Les Pediculines, Supplement, page 144;

Plate 15, figure 7. (A misprint for angulatus, since that name
appears on the plate and on the label of the type specimens.)

1932. Linognathus angulatus (Piaget), Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 5:380; figures 231D, 232G, 233.
HOSTS AND DISTRIBUTION. Type from Cephalophus nigrifrons without indi¬
cation of locality.

NOTES. Since this species was described from the same host genus as was
breuiceps, it might be suspected that synonymy is involved, but apparently
the two species are distinct. There may have been some mixing of material
or some misidentification involved. Actually the species aepycerus ap¬
proaches it very closely.

Fahrenholz has maintained that the spelling ungulatus should be pre¬
served, hut it seems clear that this was a typographical error, the spell¬
ing angulatus being used in connection with the plate accompanying the o-
riginal description and on the label of the type specimens.

Linognathus bedfordi Ferris

1932. Linognathus bedfordi Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 5:387; figures 236 , 238B, C, 11, I, and 239A, D.
HOSTS AND DISTRIBUTION. From Antidorcas marsupialis at Anderstepoort,
Union of South Africa.

Linognathus breviceps (Piaget)

1885-

1910.

1932.

Haematopinus breviceps Piaget, Les Pediculines, Supplement, page 142;
Plate 15, figure 5*

Linognathus gazella Mjoberg, Arkiv f6r Zoologi 6:157; figure 78.
Linognathus gilvus Fahrenholz, Jahrbuch der Haraburgischen Wissen-
schaftlichen Anstalten 34:2:18; figure 5-
Linognathus breviceps (Piaget), Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 5:107; figures 216A, 231B, C, E.
F, H; 232A. * '

1932. Linognathus gazella Mjdberg, Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 5:111; figures ^2B, E, F, 234.

HOSTS AND DISTRIBUTION. First described fTom Cephalophus maxwelli with¬
out indication of origin. Described as Linognathus gazella Mjoberg from a
gazelle, in the Hamburg Zoological Gardens. Described as Linognathus
gilvus Fahrenholz from Cephalophus sp. from the same Gardens, these having
been some of the same specimens, apparently, as the lot that Mj5ber<' had.
Erroneously recorded by Ferris and by Bedford as Linognathus angulatus
(Piaget) from Cephalophus natalensis and Sylviacapra grimmi from Zululand
and Transvaal.

NOTES. Concerning this species, or perhaps closely related group of
species, there is very muoh of a problem. Ferris, in connection with the
reference cited above, was able to examine types of all of them and a con¬
siderable range ol this material is still at hand. This material has been
examined and re-examined in connection with the present work and no consis¬
tent basis for the separation of these species has appeared. Perhaps even
Linognathus l imnotragi cannot definitely be separated.

In specimens from the type lot of gazella the forehead is almost rectan¬
gular, but there is some variation in this and the character does not hold.

Specimens from Cephalophus grimmi and Cephalophus natalensis — over 36
specimens from the iirst being available — show enough variation in head
form to represent two or three species if these characters can be taken

228

seriously . Some specimens have the hiudhead definitely annulate laterally,
while in others it is parallel sided.

The typical "shoe-sole shaped" genital plate ol the female appears in
the types of limnotragt, hut does not hold in other specimens which are at
hand from the same host. There is also some range of variation in the form
of the gonopods, some being acute and others rounded. However, the species
l tmnotragi is here accepted as probably distinct.

The result has been that the species gazella and gilvus are here placed

in synonymy with breviceps. _ .

This species has sometimes been placed in synonymy with L inognat hus
angulatus (Piaget), which also is attributed to Cephalophus, but the two
are clearly distinct.

Linomathus brevicornis (Giebel)

1874. Haematopinus brevicornis Giebel, Insecta Epizoa, page 43.

1932. Linognathus brevicornis (Giebel), Perris, Contributions 'lowardaMon-
O'jraph of the Sucking Lice, Part 5:3^5; figures 216D, 219-
HOSTS AND DISTRIBUTION. Prom Camelopardalis giraffa in a zoological

NOTES. The redescription given by Ferris was based upon specimens in
the Piaget collection. Whether or not these belonged to the same lot as
did Giebel 's types is not known.

Linognathus cervicaprae (Lucas)

1847. Haematopinus cervicaprae Lucas, Annales de la Socidt£ entomologique
de France (2)5:534; Plate 8, figures II, l-lh.

1938. Linognathus cervicaprae (Lucas), Werneck, Libro jubilar do Prol . Ira
vassos, page 527; figures 1-5-

HOSTS AND DISTRIBUTION. Described by Lucas from Antilope cervicapr a,
from a menagerie in Paris. Not seen again until recorded by Werneck fr°™
specimens taken from the same host species in the Zoological Garden of

^NOTES In 1939 Fahrenholz recorded under this name and described speci¬
mens from the Museum d'Histoire Naturelle de Paris which he assumed to have
been left bv Lucas and therefore possibly to be the types of the ^cies.
However Werneck had already redescribed the species and later published
note in' which he indicated that Fahrenholz agreed that these specimens were
d rob ably not those recorded by Lucas and accepted the interpretation given
bv Werneck. The species involved was referred by Werneck to Linognat.us
tibialis (Piaget) and the pertinent reference will be found under that name.

Linognathus damaliscus Bedford

1936. Linognathus damaliscus Bedford, Onderstepoort Journal of Veterinary
Science and Animal Industry 7:61; figures 3, 4. -nnl m-iral

HOSTS AND DISTRIBUTION. IVpe from Damaliscus aibi ^rons^lhe n° %u th
gardens at Johannesburg; also from Damaliscus dorcas at Bredasdorp, South

AfrNOTE5. On the basis of the illustrations given by Bedford this appears
to be identical with Linognathus taurotragi, but disposal of it must await
a re— examination of its types.

Linognathus fahrenholzi Paine

linognathus forficulus Kellogg and Paine, Bulletin of Entomological
2i47; Plate 4. fibres 2. 4. (Not Sae.atoptnus forficulus

1911-

229

Rudow, which is supposed to be also a Linognathus)

1914. Linognathus fahrenholzi Paine, Psyche 21:117.

1932. Linognathus fahrenholzi Paine, Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 5:370; figures 224, 225.

HOSTS AND DISTRIBUTION. Described as from Cervicapra arundtnarum, Marim¬
ba District, Nyasaland, and later recorded from Cervicapra fulvorufula at
Mfongosi, Zululand.

Linognathus fractus Ferris

1932. Linognathus fractus Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 5:366; figures 220, 221.

1932. Linognathus sp., Bedford, Report of the Director of Veterinary Ser¬
vices and Animal Industry, Union of South Africa 18:409.

HOSTS AND DISTRIBUTION. From Tragelaphus sylvaticus at Onderstepoort,
South Africa.

Linognathus gnu Bedford

1927. Linognathus gnu Bedford, Transactions of the Royal Society of South
Africa 14:349; figures 3, 4.

1927. Linognathus ferrisi Bedford, Transactions of the Royal Society of
South Africa 14:351; figures 5, 7«

1929. Linognathus gorgonus Bedford, Report of the Director of Veterinary
Services, Union of South Africa 15:502.

1932. Linognathus gnu Bedford, Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 5:368; figures 222, 223.

HOSTS AND DISTRIBUTION. The type of Linognathus gnu was from Connochae-
tes gnu at Clocolan, Orange Free State, and the type of gorgonus was from
Gorgon taurinus, Zoutpansberg District, Northern Transvaal . The species
was later recorded from this host at Maastrom, northern Transvaal, and in
the Zoological Gardens at Pretoria.

NOTES. This species was described by Bedford as gnu. and as ferrisi be¬
cause of the fact that he had the two sexes separately from different
hosts. The name ferrisi, being preoccupied, was later changed by him to
gorgonus.' He later concurred that they represent the same species. Fah-
renholz, in 1939, without seeing any specimens, considered the two to be
distinct merely on the basis of illustrations given by Bedford and by
Ferns. There is no reason to accept this opinion.

Linognathus hippotragi Ferris

1932. Linognathus hippotragi Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 5:373; figures 226, 227.

HOSTS AND DISTRIBUTION. From Eippotragus niger, in the zoological war¬
den at Johannesburg, South Africa. 0 0

Linognathus hologastrus Werneck

i937. Linognathus hologastrus Werneck, Memorias do Institute Oswaldo Cruz
32:397; figures 7-10.

HOSTS AND DISTRIBUTION. From Gorgon taurinus at Grootfontein Southwest
Africa.

NOTES. Although this species occurs upon the same host as does Linog¬
nathus gnu, the two species are very distinct and no confusion between them
need arise.

230

Linoguathus lewisi Bedford

1934. Linognathus leulsi Bedford, Onderstepoort Journal of Veterinary Sci¬
ence and Animal Industry 2:48; figure 11.

HOSTS AND DISTRIBUTION. From Gazella thomsont at Naivasha, British East
Africa.

Linognathus limnotragi Cummings

1913. Limnognathus limnotragi Cummings, Bulletin of Entomological Research
4:36; figure.

1932. L inognathus limnotragi Cummings, Ferris, Contributions Toward a Mon¬
ograph of the Sucking Lice, Fart 5:333; figs* 231A, I, J, 232C, D.
HOSTS AN D DISTRIBUTION. Types from Llmnotragus (= Trage laphus) gratu s in
the zoological garden of London. Recorded also trom Tragelaphus s ylvaticus
at Onderstepoort, Pretoria, and from Tragelaphus scrlptus in the zoological
garden at London.

NOTES. See the discussion under Linognathus breulceps.

Linognathus oviformis (Rudow)

1869. Haematopinus oviformis Rudow, Zeitschrift fur die gesamten Naturwis
senschaften 34:170.

1916. Linognathus ovl form is (Rudow) , Fahrenholz, Archiv fur Naturgeschichte,

Abteilung A, 81:11:33* „ 4 , „

HOSTS AND DISTRIBUTION. Recorded by Rudow from Hircus manifrlclus, a
host name that appears in no available lists. The host was presumably a goal .

NOTES. This is an utterly unrecognizable species that Fahrenholz as¬
sumed to Linognathus purely as a guess.

Linognathus ovillus (Neumann)

Figures 99, 100

1907. Haematopinus ovillus Neumann, Revue veterinaire 32:520; ligure.

1932. Linognathus ovillus (Neumann), Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 5:346: figures 209, 210 A, B, C, E.
HOSTS AND DISTRIBUTION. Described by Neumann as from domestic sheep
from New Zealand and Scotland. There have been various records which are
dubious because three other species of this genus occur on domestic sheep .
The types were examined by Ferris and, on the basis ol this, records Irom
domestic sheep in the Falkland Islands and New South Wales are here accepted.

Linognathus pedalis (Osborn)
Figures 101, 102

1896. Haematopinus pedalis Osborn, United States Department ol Agriculture,
Division of Entomology, BuLletin (new series S.liO, l^ure .
1911. Haematopinus microcephalus Garnett, Journal of Comparative Patho ogj
and Therapeutics, page 2; figures 1, 2. M

1932. Linognathus pedalis (Osborn), Ferris,1 Contributions ^Toward a. Mono¬
graph of the Sucking Lice, Part 5:344; ligures 207B, D, 2DS.

HOSTS AND DISTRIBUTION. First described from domestic sheep in the
United States and recorded from this host in South America, New Zealan ,
Australia, and South Atrica.

Linognathus peleus Bedford

1936. Linognathus peleus Bedford, Onderstepoort Journal of Veterinary Sci

231

Linognathus ovillus (Neumann)

spiracles (above) compared with
same of stenopsis ( below)

female genitalia

male genitalia

Linognathus ovillus (Neumann), details

Figure 100

ence and Animal Industry 7:59; figures 1, 2.

HOSTS AND DISTRIBUTION. From Pelea capreolus at Onderstepoort, South

Africa.

Linognathus petasmatus Ferris, new species

HOSTS AND DISTRIBUTION. Holotype, a female, allotype, and several par ar-
tvpes, young and eggs, from "North African anti lope, presumably tron. a
zoological garden in Manchester, England. Types in the collections oi

Stanford University, California. . , u ,

FEMALE. About 2.25 mm. in length. Body form rather slender. Head e
Ion crate and relatively slender, ven smoothly fusiform, the forehead on y
slightly' shorter than the hindhead and apically acute, the widest point o
the hindhead being slightly posterior to the center of this p0^10"'.
rax and legs entirely normal. Abdomen with an uninterrupted row ot short
setae and a median cluster of similar setae on segments 3-6 both dorsJ^
j ventrallv Median genital plate of the female unusually large, very
definitely lozenge-shaped, the anterior extremity slightly Prol^ed* ^“-
opods of female slightly convergent posteriorly, narrowly rounded apically
and with setae around the apex and slightly up the mesal margin.

233

Linognathus pedalis (Osborn)

Figure 101

MALE. About 2 mm. long. Genitalia with basal plate long and slender,

®xpanded ^nd slightly bifid at posterior end and somewhat expanded
at the anterior end. Parameres apparently strongly curved dorsoventrally •
the pseudopenis very short and indistinct, the endomeral piece either lack¬
ing or very weaklj' developed.

NOTES. This species was received some years ago from a correspondent in
Birmingham, England, whose name has unfortunately been lost, and it is not
clear whether the specimens came directly from North Africa or from an ani¬
mal in a zoological garden. The species seems to be quite distinct from
anything else that has been described.

Linognathus pithodes Cummin ^s

1916. Linognathus pithodes Cummings, Proceedings of the Zoological Society
of London, page 260; figures 3-5.

1932. Linognathus pithodes Cummings, Ferris, Contributions Toward a Mono¬
graph ot the Sucking Lice, Part 5:385; figures 235, 238D, 239F, G.

234

Linognathus pedalis (Osborn), details

Figure 102

1949. Stobbella pithodes (Cummings), Eichler, Bolletino della Societa En-
tomologica Italiana 79:13*

HOSTS AND DISTRIBUTION. From Antilope cervicapra from India, in the zo¬
ological garden at London.

NOTES. Eichler has designated this species as type ot the genus Stob¬
bella, which is not here accepted.

Linognathus saccatus (Gervais)

1845* Haematopinus saccatus Gervais, In Walckenaer, Histoire Naturelle des

Insectes Apt^res 3:307* . .

1905. Linognathus saccatus (Gervais), Enderlein, Zoologischer Anzeiger 29:

194.

HOSTS .AND DISTRIBUTION. Known only from the original record from "un
Bouc d ' Egyp te . " An utterly unrecognizable species which has been placed in

Linognathus purely as a guess.

Linognathus setosus (von Oilers)

Figures 103, 104

1S16. Pediculus setosus von Olfers, De vegetativis et animatis corporibus
in corporibus animatis reperiundis commentarius, page 80.

235

Linognathus setosus (von Olfers) Figure 103

1838. Pedtculus piliferus Burmeister, Genera Insectorum, Rhynchota, Spe¬
cies 13.

1842. Haematopinus piliferus (Burmeister), Denny, Monographia Anoplurorum
Britanniae, page 28: Plate 25, figure 4.

184 1 . Haematopinus bicolor Lucas, Annales de la Soci6t4 Entomolo<rique de
France (2)5:538; Plate 9, figure 2a.

1861. Fed iculus isopus Nitzsch, Zeitschrift tiir die gesajnten Naturwissen—
schaften 18:290.

1864. Pediculus flavidus Nitzsch, Zeitschritt tiir die gesamten Naturwis—
sen schaften 23:2 7 .

1874. Haematopinus piliferus (Burmeister), Giebel, Insecta Epizoa, page 40.

1905. Linognathus piliferus (Burmeister), Enderlein, Zoolo^ischer Anzei<rer
29:194. 0

1919. Linognathus setosus (von Olfers), Fahrenholz, Jahresbericht des Nie-
dersachsischen zoologischen Yereins zu Hannover 5-10:23.

236

female genitalia

male genitalia

Linognathus setosus (von Olfers)

Figure 104

1932. Linognathus setosus (von Olfers), Ferris, Contributions Toward a Mon¬
ograph of the Sucking Lice, Part 5:70; figures 206, 207A, C, F, G

and 216E. . , „ j

HOSTS AND DISTRIBUTION. Described from the domestic dog in Europe and
reported from this host throughout the world. Also recorded from the white
fox " or "Arctic fox," presumably Alopex lagopus, from various localities
in Canada and Alaska; from fox in Manchuria; from Canis cupus in Croatia,
from unspecified captive fox in the United States; from coyote, ( Can
sp.) in the United States and even recorded from ferret and from rabbit.
NOTES. This is the type of the genus Linognathus .

Linognathus spicatus Ferris

1932. Linognathus spicatus Ferris, Contributions Toward a Monograph of the
Sucking Lice, Part 5:388; figures 237, 238A, G, and _39E.

HOSTS AND DISTRIBUTION. From Gorgon taurinus, Maastrom, northern irans-

^NOTES. This species need not be confused with either of the other two
members of this genus which occur on Gorgon. They are very distinct fonrs,
as will be seen by reference to the key.

237

Linognathus stenopsis (Bunneister)
Figures 105, 106

1838.

1842.

1864.

1869.

1905.

1916.

1932.

Pediculus stenopsis Bunneister, Genera Insectorum, Rhynchota, Spe¬
cies j. ’ r

Haenatoptnus stenopsis (Bunneister), Denny, Monographia Anoplurorum
ontanniae, page 36.

PetlCsUJu!ohln!u°23-S0 NUZSOh’ ZeitSohrift f“r die eesmtea Natur-

“Sfi°nr» RUd°W’ ZeitS°hrift f“r die «— *«

"ZllZTZnT^m. RUd°W’ ZeHs0hrift di° Natur-

L%‘m'!US stenopsts (Bunneister) , Enderlein, Zoologischer Anzeiger

linognathus forflculus (Rudow), Fahrenhoiz, Archiv fur Naturge-
schichte, Abteilung A. 81:11:24; figures 19, 20.

Linoinathus stenopsis (Bunneister), ferris. Contributions Toward a
212ASCaPD °F the Sucklng Llce> Part 5:349; figures 210D, 211,

HOSTS AND DISTRIBUTION. Originally described from domestic goats in
Europe and since recorded many times from this host in various parts of the
world. Some o i the numerous records are perhaps erroneous because of the
presence of other species of this genus on this host. Described by Rudow

N0t£C FerrTs”1 MqS) ^ *S ru^ica^ae from Caprella rupicapra.
with thf; Ipip'j 9 ? has discussed at length the problems connected

W) ^ d ^ various names that have been employed for it. He

had at hand specimens from Capra ibex and Caprella rupicapra which he con-

H1 erte+Kt0 lde!ltlcal with those from domestic goats and consequently re-

review^of “1 ruPic^ to synonymy with s tenopsil A

™ oLni! Tk11 connection with the present work indicated no rea-

1D SPlte °f the faCt ^ attempted

Linognathus taeniotrichus Werneck

1937' Ll^hUlt"nl0t:iChU3 Werneck- Memories do Institute Oswaldo Cruz
3^:3yi; iigures 1-5.

HOSTS AND DISTRIBUTION. Described from Canis brasiliensis at Sao Ber¬
nardo das Russas, state of Ceard and recorded also from Canis azarae from
Lassance, state of Minas Geraes, Brasil. azarae t rom

NOTES. This species seems clearly to be distinct from Pediculus setosus
which occurs on the domestic dog and also on arctic foxes. *

Linognathus taurotragus Bedford

1927‘ “ZSSTJSZ'gfiflSSSi r“TCtiM,s 0f the ^

193;- iKSS gr ‘ MOD-

HOSTS AND DISTRIBUTION. Described from TaurotmZ! 07ux \ t CloeoLn
Orange Free State. Also recorded from the sume host inNaJX South A, A*”’.

Linognathus tibialis (Piaget)

K80' '“^8.“ ttbta“S Piaeet’ US Pedic“lines> page 646; Plate 52,

188°. Saenotoptnus tibialis, variety antennatus Piaget, Les Pediculines,

238

Linognathus stenopsis (Burmeister)

Figure 105

239

female genitalia

Linognathus stenopsis (Burmeister), details

male genitalia
Figure 106

1880. Haematopinus tibialis, variety append iculatus Piaget, Les Pedicu-
lines, page 647.

1916. Linognathus tibialis, variety euchore Waterston, Annals of the South
African Museum 10:275; figure.

1932. Linognathus tibialis (Piaget), Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 5:360; figures 217, 218.

HOSTS AND DISTRIBUTION. Type from "Antilope maori " from the zoological
garden at Rotterdam and recorded also by Piaget from Antilope subgutturosa
and Ant Hope sp. from the same place. The type of Waterston's variety
euchore was from Anti lope euchore in South Africa and the species has been
recorded by Bedford from Aepyceros melampus from South Africa.

NOTES, ieiris, in connection with the reference cited, examined the
types of the forms recorded above.

Specimens from Antidorcas marsupial is from Onderstepoort , Pretoria
South Africa, which were recorded by Ferris as tibialis have been re-exam¬
ined and raise some doubt as to their identification. They are certainly
very close to tibialis, but differ in the form of the genital plate of the
female. Other specimens from Raphicerus campestris from the Rustenburg
District in the Transvaal, South Africa, are extremely close to* tibialis
but are somewhat different. Questions concerning these may be allowed to
rest until someone with more material can approach the problem.

240

Linognathus vituli (Linnaeus)

Figure 107

Linognathus vituli (Linnaeus)
Figures 107, 108

1758. Pediculus vituli Linnaeus, 3ystema Naturae, ^Edition X, page 611.
1829. Haematopinus vituli (Linnaeus), Stephens, Catalogue of British In¬
sects 2: 329 .

1838. Pediculus tenuirostris Burmeister, Genera Insectorum, Rhynchota,

Species 17. M .

1864. Pediculus oxyrrhynchus Nitzsch, Zeitschrift fur die gesamten Natur

wissenschaften 23:21.

1932. Linognathus vituli (Linnaeus), Ferris, Contributions Toward a Mono
Taph of the Sucking Lice, Part 5:356; figures 214, 2l5, -16C.
HOSTS AND DISTRIBUTION. Originally described from domestic cattle in
Europe. Since reported from domestic cattle in all parts of the world.
There are certain unusual records from wild boar in Europe, t rom sheep, and
even from domestic dog, which may result from erroneous information as to

the host.

241

thoracic dorsum

head

Linognathus vituli (Linnaeus), details

Figure 108

Genus MICROTUORACIUS Fahrenhoiz

1915. H icrothoracius Fahrenhoiz, Archiv fur Naturgeschichte, Abteilung A,

81: 11:89.

1932. Hicrothoracius , Ferris, Contributions Toward a Monograph of the Suck¬
ing Lice, Fart *1:390.

1933* Hicrothoracius, Werneck, Memorias do Instituto Oswaldo Cruz 27:21.

GDi ERIC TYPE. Haematopinus (Linoinathus) praeloniiceps Neumann.

CHARACTERS. Linognathidae in which clearly evident eyes are present,
these being represented by a lens just posterior to the base ol' each anten-
na. Head greatly elongated and more or less fusiform, at times being al¬
most as long as the abdomen. Thorax very small, with no indication of a
sternal plate but with the sternal apophyses present and with a definite
notal pit. Antennae five- segmented but with segments 4-5 at times more or
less fused. Spiracles beset internally with points. Derm of the abdomen
tending to be minutely wrinkled.

NOTES. The assignment of this geuus to the Linognathidae is open to
question, for an argument can certainly be made supporting its assignment
rather to the Haematopinidae. The chief reason for the present assignment
is the absence of the paratergal plates, but the presence of the thoracic
sternal apophyses, the definite eyes, the thoracic notal pit, and possibly
the spiracles, argue for its placement with the Haematopinidae.

Such understanding of this genus as we have we owe chiefly to the work

of Dr. Fa bio Werneck.

The history of this genus and especially of its type species is ol in¬
terest and is worth recital. In 1688 Redi, in his famous work on the gen¬
eration of insects, illustrated certain ectoparasites of birds and mammals,
among these being one which he designated merely' as the louse of the camel.
There was apparently no discussion of any of these species, identity being
indicated merely by the legend on the illustration.

In his "Systema Naturae," Linnaeus gave this louse a binomial name, call¬
ing it Pediculus cameli, purely on the basis of the illustration given by
Redi. From the time of Redi until as recently as 1934 the insect was never
again reported. In fact the illustration given by Redi seemed so fantastic
that its faithfulness was doubted and the peculiarities ol the species were
ascribed merely to bad drawing of some other species, perhaps Haematopinus
tuberculatus, which was the only sucking louse known from camels. However,
in 1909. Neumann described a species from a llama which was sufficiently
similar ’to cameli to indicate that the latter was not imaginary. In 1915
Fahrenhoiz assigned Neumann's species to the genus Hicrothoracius.

In 1932 Ferris discussed this genus. He had examined a female from the
type material of praeloniiceps and had also at hand specimens from a llama
from the zoological park at Washington. His conclusions concerning the
species were thus based upon what seemed to be quite authentic material.

However, in 1932, Werneck described a new species, Hicrothoracius mazzal,
and in 1933 he discussed these two species at length, the result of his
work being to indicate that Neumann actually had two quite distinct species
in his type lot of praeloniiceps . Neumann had indeed indicated that one ol
his specimens differed from the others in the form of the head and it seems
clear that this was the specimen examined by Ferris.

Then in 1934 Werneck reported the rediscovery of the long-lost Pediculus
cameli on the basis of specimens taken in Algeria and he described another
species, Hicrothoracius minor, from another llama. In 1935, Ferris dis¬
cussed the crenus again but his work had gone to press before kerneck s
paper of 1934 had become available and full account had not been taken ol

Werneck 's conclusions. . .

It is now evident that there are at least four species ol this genus on
members of the family Camelidae, a family that according to Itfdekker s work

243

•1,,t ® a,s contains but tw0 genera, Camelus with one species and

Lama with but two. Whether or not we even yet have the full complement of
Anoplura occurring on this family remains to be determined.

Key to Species of MICR01H0RACIUS

a,HkreCk4haf4dKiS4CUSfd the problem of separating the species of this genus
and has noted that it is difficult to do so upon the basis of any single
character but that, m the totality of their characters, they seem to°be
readily recognizable. Unfortunately, specimens of minor are not available
and this species must be included in the key solely upon the basis of
Werneck s description, while available material of the other species is ex¬
tremely scanty. Furthermore, differences in the treatment of the specimens
may cause apparent differences in the form of the head and lead to confu-
sion thus in mazzai a species in which the head is extremely slender, a
slight collapse of the specimen will exaggerate this slenderness. In the
light of this the following key must be used with discretion.

1. Head very narrow throughout, but with a quite definite swelling on each

side just posterior to the antenna, this swelling bearing the eyes:

it P°s^erior this swelling the head narrowing aorain . MAZ7AT

Head broader, definitely fusiform, expanding into a moderately Tharp
angle just posterior to each antenna, narrowing again immediately
posterior to the swelling, the margins of the hindhead being smoothly

2. Anterior legs almost identical in size and shape with the others: a iar-

?®r t01rm' attaining a length of 4 mm. for the female and 2.5 mm. for
the male; 4th and 5th antennal segments quite closely fused. .. .CAMELI
Waller forms the adult female not exceeding 3 mm. in length; first

-*-egs definitely more slender than the others . 9

J. Female attaining 3 niro- in length . PRAFT/lNrTnrpQ

Female recorded as less than 2 mm. long . 5^77. N^MiSr

Microthoracius cameli (Linnaeus)

Figures 109, 110

Pediculus cameli Linnaeus, Sy sterna Naturae, Edition X, page 611
18/4. Haematopinus cameli (Linnaeus), Giebel, Insecta Epizoa, p^ge 47.’

1909. Haematopinus tuberculatus (Burraeister) , Neumann, Archives de Para-
sitologie 13:499. (Misidentification)

1916. Microthoracius cameli (Linnaeus), Fahrenholz, Archiv fur Naturcre-
schichte, Abteilung A, 81:11:30. " °

1932. Microthoracius cameli (Linnaeus), Ferris, Contributions Toward a Mon-
int), ograph of the Sucking Lice, Part 5:394; figure 242.

1934. Microthoracius cameli (Linnaeus), Werneck, Memorias do Instituto Os-
waldo Cruz 29:179; figures 1-5.

HOSTS AND DISTRIBUTION. The only actual record of the species is that
given by Werneck, from Camelus dromedarius in Algeria. P 13 thdt

Microthoracius mazzai Werneck

1909. Haematopinus (Llnoinathus) praeloni iceps Neumann, Archives de Para-
1014; stogie 13:509. (In part; specimen recorded in a footnote)

1916. Microthoracius praeloni iceps (Neumann), Fahrenholz, Archiv fur Na-
ioo9 turgeschichte, Abteilung A, 81:11:30. (In part)

93 • Microthoracius praeloni iceps (Neumann), Ferris, Contributions Toward

(MisYdenUftcation* Dg LlC6’ Part 5:S9: fi6ures 24°- 2«-

244

Microthoracius cameli (Linnaeus)

245

female genitalia

Microthoracius cameli (Linnaeus), details

Figure 110

246

1932. Mtcrot horac lus mazzal Werneck, Revista medico-cirurgica do Brasil
40:346; figure.

1933* Mlcrothoraclus mazzal Werneck, Werueck, Meaorias do lnstituto Oswal do
Cruz 27:26; figures 9-15*

1935* Mlcrothoraclus mazzal Werueck, Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Fart 9:612; figures 338B-E.

HOSTS AND DISTRIBUTION. Types from Auchenia liana at Santa Catalina,
province de Jujuy in Argentina. Presumably also the specimen recorded by
Neumann in the footnote to his description of praeloniiceps, this from
Auchenia llama from Choquecomato, Bolivia, is this species, and the speci¬
mens recorded by Ferris from the same host, froman animal in the Zoological
Park of Washington, are the same.

Microthoracius minor Werueck

1935. Mlcrothoraclus minor Werneck, Revista medico-cirurgica do Brasil 43:

112.

1935. Microthoracius minor Werneck, Werneck, Revista de Entomologia 5:168;
figures 1-4, 5-

HOSTS AND DISTRIBUTION. Types from Lama pacos, Abra Pampa, province de
Jujuy, Argentina, and other specimens from Lama ilama at the same place.

Microthoracius praelongiceps (Neumann)

1909. Haematoplnus ( Linoinathus ) praeloniiceps Neumann, Archives de Para-
sitologie 13:508; figures 10-12. (In part)

1916. Microthoracius praeloniiceps (Neumann), Fahrenholz, Archiv fur Na-
turge schichte, Abteilung A, 81:11:30. (In part)

1932. Microthoracius praeloniiceps (Neumann), Werneck, Revista medico-

cirurgica do Brasil 40:346; figure.

1933. Microthoracius praeloniiceps (Neumann), Werneck, Memorias do Insti¬

tute Oswaldo Cruz 27:21; figures 1-8.

1935. Microthoracius praeloniiceps (Neumann), Ferris, Contributions Toward
a Monograph of the Sucking Lice, Part 8:610; figure 338A.

HOSTS AND DISTRIBUTION. Types recorded as from Auchenia huanaca, Choque-
comato, Bolivia. Also recorded by Werneck from Auchenia llama. A specimen
at hand from "llama" in the zoological garden at Onde rstepoort , South
Africa, received from the late Mr. G. A. H. Bedford, seems definitely to be
this species.

Genus PR0LIN0GNATHUS Ewing

1929. Prolinoinathus Ewing, Manual of External Parasites, pages 136, 201.
1932. Prolinoinathus, Ferris, Contributions Toward a Monograph of the Suck¬
ing Lice, Part 5:408. _ , ...... i .

1939. Prolinoinathus, Fahrenholz, Zeitschrift fur Parasitenkunde 111:1.1.
GENERIC TYPE. Pediculus caviae-capensis Pallas.

CHARACTERS. Linognathidae in which there is no external indication of
eyes. Antennae apparently' four-segmented because of a close fusion of the
fourth and fifth segments, although the sensoria of these segments remain.
Thorax without a sclerotized sternal plate and the ventral thoracic apophy¬
ses lacking. Spiracles of the abdomen extremely small. Abdominal setae
much reduced in numbers, there being usually not more than two setae in the
median group on each segment, either dorsally or ventrally, and never any
indication of more than one row of setae on any segment. Gonopods ol the

female well developed and produced apically. . „ r 4.

NOTES- The members of this genus occur exclusively on members ol the

family Procaviidae of the Order Hyracoidea.

247

The material at hand representing this genus is quite scanty and does
not permit an extended treatment.

Key to Species of PROLINOGNATHUS

This key is based largely upon that given by Fahrenholz in the reference
cited above, with some modifications suggested by such material as is avail¬
able.

1. Head definitely more than twice as long as wide . 2

Head not more than twice as long as wide . 3

2. Sclerotized areas at the lateral margins of the forehead produced some¬

what posteriorly on the dorsal side about the bases of the antennae..

^ . •••••: . CAVI AE- CAPEN SI S

These sclerotized areas produced posteriorly on the dorsal side of the

head until they meet at the mid-line . ARCUATUS

3- Abdomen with a single long seta on the lateral margin of each of the
segments anterior to the seventh, in addition to the usual, long,

paired setae at the margins of segments 7 and 8 . LEPTOCEPHALUS

Abdomen with such setae absent only on segments 5 and 6 . 4

4. These setae present only on segments 2 and 3 . AEftilOPICUS

These setae present on segments 2-4 . pj

5* Forehead rather short and the sclerotized transverse area so broad that

it involves at least half its length . FERRISI

Forehead with this transverse area involving at the most one-fourth of
its length . FOLEYI

Prolinognathus aethiopicus Fahrenholz

1939- Prolinoinathus aethiopicus Fahrenholz, Zeitschrift fur Parasiten-
kunde 11:1:13; figures 8 A, 9.

HOSTS AND DISTRIBUTION. Described as taken from Procauia shoana in the
zoological garden at Copenhagen. According to Hopkins the proper name of
this host is Procavia habessinica sciona. A specimen that seems to be this
species is at hand from a host identified merely as Procauia capensis from
Rooi Krans, Transvaal.

Prolinognathus arcuatus Fahrenholz

1939. Prolinognathus arcuatus Fahrenholz, Zeitschrift fur Paras it enkunde
11:1:5; figures lb, 2-4.

HOSTS AND DISTRIBUTION. Described as from the same animal as recorded
under aethiopicus . Certain specimens at hand from a host identified as
Procavia coombsi at Onderstepoort, Pretoria, South Africa, seem to be this.
This host seems to be regarded as a subspecies of Procavia capensis.

Prolinognathus caviae-capensis (Pallas)

Figures 111, 112

1767. Pediculus caviae-capensis Pallas, Spicilegia Zoologica 2:32: Plate
3, figure 12.

1816. Pediculus collaris von Olfers, De vegetativis et animatis corporibus
in corporibus animatis reperiundis commentarius, page 84.

1913. Linoinathus caviae-capensis (Pallas), Cummings, Bulletin of Entomo¬
logical Research 4:37; figures 2, 3.

1932. Prolinoinathus caviae-capensis (Cummings), Ferris, Contributions
Toward a Monograph of the Sucking Lice, Part 5:409; figures 250A.
251A, E, F, G, H, I, J. '

248

Prolinognathus caviae-capensis (Pallas'

Figure 111

249

Prolinognathus caviae-capensis (Pallas), details

Figure 112

1939. Prolinognathus caviae-capensis (Pallas), Fahrenholz, Zeitschrift fur
Parasitenkunde 11:1:2; figure la.

1949. Prolinognathus caviae-capensis (Pallas), Hopkins, Proceedings of the
Zoological Society of London 119:2:515, 516.

HOSTS AND DISTRIBUTION. Originally described as from Procavia capensis
t rom the Cape ot Good Hope. Recorded by Cummings from this host and from a
host identified under the same name in the zoological garden at London.
The species was redescribed by Ferris on the basis of Cummings' material
and a record 1 rom a host identified as Procavia coombsi from Onderstepoort,
Pretoria, South Airica, was added. Hopkins records the species from a lono1
list ot subspecies ol Procavia capens is, this including capensis, narlothi ,
klaverensls, grlquae, albaniensis, natalensis, windhuki, reuningi, and
waterbergensls, and from Procavia johnstoni johnstoni from various local¬
ities.

250

NOTES. The present writer has previously maintained that the first gen
uine validation of the name cav iae-capens is is to be ascribed to Cummings
but Fahrenholz has given cogent reasons for accepting Pallas as the author
of the specific name.

Whether or not Cummings actually had the species described by Pallas is
open to question, but there is no special point to inquiring too closely
into this. The accompanying illustrations are based upon his specimens.

Prol inoguathus ferrisi Fahrenholz

1932. Prolinognathus leptocephalus (Ehrenberg), Ferris, Contributions
Toward a Monograph of the Sucking Lice, Part 5:142; figures 25011,
251C. (Misideiitification)

1939. Prol i nognat hus ferrisi Fahrenholz, Zeitschrift fur Parasi tenkunde

11:1:12.

HOSTS AND DISTRIBUTION. The name given by Fahrenholz seems to have been
based entirely upon the illustration given by Ferris and consequently the
type of the species must be the specimen from which this illustration was
made, this having been recorded as from Procauia brucel rudolfi, Marsabit
Road, British East Africa, which, according to Hopkins, is Heterohyrax
syriacus rudolfi. This type specimen should be in the United States
National Museum. Hopkins has also recorded the species from the same host.

Prolinognathus foleyi Fahrenholz

1939. Prolinognathus foleyi Fahrenholz, Zeitschrift fur Parasi tenkunde 11:
1:9; figures 5—7, 8b.

HOSTS AND DISTRIBUTION. Recorded as from Procavla rufescens bounhloll
from Algeria. This is the only record.

Prolinognathus leptocephalus (Ehrenberg)

1828. Pediculus leptocephalus Ehrenberg, Symbolae Physicae, Decas Prima,
page f.

1874. Haenatoplnus leptocephalus (Ehrenberg), Giebel, Insecta Epizoa, page
47. (In part)

1932. Prolinognathus leptocephalus (Ehrenberg), Ferris, Contributions
Toward a Monograph of the Sucking Lice, Part 5:142. (In part; not
as to figures)

1939. Prolinognathus leptocephalus (Ehrenberg), Fahrenholz, Zeitschrift
fur Parasi tenkunde 11: 1:8.

HOSTS AND DISTRIBUTION. Described from Procauia syriacus, presumably
from Syria- Ferris recorded a single immature specimen from this host from
Syria and Fahrenholz has given additional notes on the species.

Genus SOLENOPOTES Enderlein

1904. Solenopotes Enderlein, Zoologischer Anzeiger 28: 143-

1909. Haematopinus (Solenopotes) Neumann, Archives de Parasitologie 13:530-

1915. Cervophthirius Mj&berg, Entomologisk Tijdskrift 36:282.

1916. Linognathus, Ferris, Entomological News 27:199-

1921. Solenopotes, Bishop, Journal of Agricultural Research 21:797.

1929. Cervophthirius, Ewing, A Manual of External Parasites, page 136 .
1929. Solenopotes, Ewing, A Manual of External Parasites, pages I36, 139.
1932. Solenopotes, Ferris, Contributions Toward a Monograph of the Sucking
Lice, Part 5:395-

GENERIC TYPE. Solenopotes caplllatus Enderlein. Typ e of Cervophthirius:
Cervophthirius tarandi MJoberg.

251

CHARACTERS. Linognathidae without eyes. Antennae five-segmented. Head
variously shaped hut usually slightly elongate and broadened but little
•posterior to the antennae. Thorax with a sternal plate filling most of the
space enclosed by the coxa but not marginally or apically free from the
body. Abdomen membranous throughout except for the usual ninth tergite and
the ventral genital areas; with not more than one row of setae on any seg¬
ment either dorsally or ventrally. Spiracles, in the type species, more or
less cylindrical, their apices projecting somewhat from the body on slight¬
ly sclerotized prominences. In other species they may be shorter, in some
species being almost spherical, but some indication of the tubercle is
present. Genitalia of female including a pair of quite large and prominent
gonophyses. Ninth segment terminating in a pair of ventral, flattened
lobes which may be quite long and slender. Genitalia of the inale with the
parameres well developed, elongate and enclosing the penis and pseudopenis.

NOTES. The type species of this genus was based upon a single male
specimen which was rather poorly described and led Ferris to the belief
that it was simply an immature specimen of Linognathus vltull. This was
later shown to be quite erroneous when the species was rediscovered in
North America. MjOberg described the supposed new genus Cervophthirius in
1915> but this was placed as a synonym of Solenopotes by Ferris in 1932.

Except for the one species, capillatus, all the members of this genus at
present known are from Cervidae. It is probable that several more remain
to be found .

Key to Species of SOLENOPOTES

Only a partial key can at present be given, and identifications must de¬
pend largely upon host associations until the genus has been completely re¬
viewed by someone.

1. Head with the lateral margins posterior to the antennae tending to con¬

verge and presenting neither postantennal nor posterior lateral

angles . .

Head with the lateral margins posterior to the antennae tending to be
nearly parallel, with definite postantennal and posterior lateral
angles . g

2. Abdominal spiracles strongly protuberant; female with apical lobes of

the abdomen which are short and moderately broad and then are con¬
stricted sharply into a short, slender, terminal process; male with
the parameres enclosing a broadly Y-shaped pseudopenis; as far as

known occurring only on domestic cattle . CAPILLATOS

Abdominal spiracles but very slightly protuberant; female with the api¬
cal lobes of the abdomen constricted gradually into long, tapering
processes; parameres of the male enclosing merely a very short aedea-
gus; occurring as far as known on New World deer of the genera Odo-
co ileus and Mazama . BINIPILOSUS

3. Abdomen of the female with not more than 2 long setae in the median

group, either dorsally or ventrally, on any segment . CAPREOLI

Abdomen of the female with 4-8 long setae in the median cluster, both
dorsally and ventrally, on the abdominal segments; occurring as far

as known on New World deer of the genus Odocoileus . FERRISI

(The species tarandt and burmeistert will run in this key to the last
couplet.)

Solenopotes binipilosus (Fahrenholz)

1910. LtnoQnathus an£ulatus (Piaget), Mjbberg, Arkiv f6r Zoologi 2:157.
(Misidenti fication)

252

1916. Linognathus btnlpl losus Fahrenholz, Archiv fur Naturgeschichte, A t»-
teilung A, 81:11:11; Plate, figures 11— 1^ -
1916. Linognathus coassus Fahrenholz, Jahrbuch der Hamburgischen Wissen-
schatt Lichen Anstalten 34:2.

1927. Lino&nathus panamens is Ewing, Proceedings of the Entomological Soci¬
ety of Washington 29:119.

1932. Solenopote s binipilosus (Fahrenholz), Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 5:131: figures 245, 246.

HOSTS AND DISTRIBUTION . First described by Fahrenholz from "Mazaina
liirsch" without indication of origin, but this name implies a host of the
genus Mazama from South or Central America. Recorded by Falirenholz from
"Ooossus" from the Hamburg zoological garden, this generic mune apparently
being a synonym of Mazama. Recorded by Ewing from Odocoileus chiriqulensis
from Panama and by Ferris from the same host species from the Panama Canal
Zone. Specimens are at hand from Mazama s imp l ic icorn is at Juj uy, Argentina;
from Odocoileus couesi at Tucson, Arizona; United States; and from "deer,"
at Sonora, Texas, United States.

Solenopotes burmeisteri (Fahrenholz)

1818. Pediculus crassicornis Nitzsch, Germar's Magazin der Entomologie 3*-
305. (Not Pediculus crassicornis Scopoli, and therefore preoccu¬
pied)

1916. Lino^nathus crassicornis (Nitzsch), Fahrenholz, Archiv fur Naturge-
schichte, Abteilung A, 81:11:34.

1919. Linofnathus burmeisteri Fahrenholz, Jahresbericht des Niedersach-
sischen zoologischen Vereins zu Hannover 5-10:23-
1935- Solenopotes burmeisteri (Fahrenholz), Freund, Recueil de Travail d6-
di£ au 25me Anniversaire scientifique du Professor Eugeni Pavlov¬
sky, 1909, 1934, Leningrad et Moscow, page 278; figure A-
HOSTS AND DISTRIBUTION. Described from Cervus elaphus in Europe and
known only from a few records from that continent.

NOTES." European authors have considered that the species recorded from
North Anerica by Ferris as this, under the name of Solenopotes crassicornis,
is distinct, as is also the species illustrated by Ferris as burmeisteri in
1932, and which has been named as Solenopotes capreoli. The name crass i-
cornis, being preoccupied, was replaced by the name burmeisteri in 1919-
These three species are very closely similar and it remains to be deter¬
mined just how they can be separated.

Solenopotes capillatus Enderlein

Figures 113, 114

Solenopotes capillatus Enderlein, Zoologischer Anzeiger 28:144; lig-

ures 14, 15- ^

Linotnathus vituli (Linnaeus), Ferris, Entomological News 27:199-

(Misidentification)

Solenopotes capillatus Enderlein, Freund, Zentralblatt fur Bakteri-
ologie und Parasitenkunde (1)84:142; figure.

Solenopotes capillatus Enderlein, Bishop, Journal ol Agricultural
Research 21:797; figures.

Solenopotes capillatus Enderlein, Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 5:397; figures 243, 244.

HfKTS AND DISTRIBUTION- First recorded from domestic cattle in Germany .
Now known from domestic cattle from various localities in Europe and North

America. , . ,

NOTES. The opinion is held by the writer that this species has trans

ferred to domestic cattle from deer, although it has never been taken 1 rom

1904.

1916.

1920.

1921.

1932-

253

Figure 113

female genitalia

Solenopotes capillatus Enderlein, details

255

male genital plate
Figure 114

any cervid and this opinion may be entirely erroneous. An extended discus¬
sion of the biology of the species is given in the reference by Bishop
cited above.

Solenopotes capreoli Freund

1932. Solenopotes burmeisteri (Fahrenholz) , Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 5:404; figures 247, 248. (Mis-
identification)

1935. Solenopotes capreoli Freund, Recueil de Travail dedie au 25me Anni-
versaire, Scientifique du Professor Eugene Pawlowsky, 1909, 1934,
Leningrad and Moscow, page 278; figure B.

HOSTS AND DISTRIBUTION. The specimens upon which the record and illus¬
trations given by Ferris (reference cited above) were based were from Cap-
reolus caprea from Czechoslovakia. Freund described the species as new on
the basis of specimens from the same host without indication of origin.

NOTES. This species, as Freund remarked, is "In alien Details . prak-

tisch identisch mit Solenopotes ferrisi . " differing only in the ar¬

rangement of the abdominal setae. Differences cited between this and bur¬
meisteri are so slight as to offer no definite "key characters" for the
separation of these two species.

Solenopotes ferrisi (Fahrenholz)

1916. Cervophthirius crassicornis (Nitzsch), Ferris, Entomological News
27:197; figures. (Misidentification)

1919. Linoinathus ferrisi Fahrenholz, Jahresbericht des Niedersachsischen
zoologischen Yereins zu Hannover 5-10:24.

1932. Solenopotes ferrisi (Fahrenholz), Ferris, Contributions Toward a Mon¬
ograph of the Sucking Lice, Part 5:134; figures 247, 248.

HOSTS AND DISTRIBUTION. Type from Odocoileus columbianus at Laytonvi lie,
Mendocino County, and recorded also from the same host at San Gregorio, San
Mateo County, California, United States.

NOTES. This species was identified by Ferris as the European species
which is now known as Solenopotes burmeisteri, but Fahrenholz considered it
to be distinct and named it as new on the basis of Ferris' description.
For this reason it appears that the types of the species must be regarded
as being in the material recorded by Ferris. Freund has given a redescrip¬
tion of burmeisteri, but it is still not clear wherein the two species dif¬
fer, if they do so at all.

Solenopotes muntiacus Thompson

193S. Solenopotes muntiacus Thompson, Annals and Magazine of Natural His¬
tory (Series 11) 1:634; figures.

HOSTS AND DISTRIBUTION. Known only from the original record from Munti¬
acus malabaricus at Mousakande, Gammaduwa, Ceylon. According to Hopkins
this is Muntiacus muntjak.

NOTES. Unfortunately the description and illustrations of this species
are not adequate to permit any very clear concept of it or to afford any
"key characters" for its identification. It is said to be very similar to
binipilosus.

Solenopotes tarandi (Mjbberg)

1915. Cervophthirius tarandi Mjoberg, Entomologisk Tijdskrift 36:283; fig¬
ures 1-4.

1932. Solenopotes tarandi (MjOberg), Ferris, Contributions Toward a Mono-

256

graph of the Sucking Lice, Fart 5:136.

HOSTS AM) DISTRIBUTION. Described as from Han^ifer tarandus at K;tresu-
ando, Sweden. Hopkins, in his recent host list, indicates another record,
which has not been traced in connection with the present work.

NOTES. The original description and illustrations of this species are
inadequate and offer no basis for its separation from such species as bur-
nelsterl and ferrtsi.

Family NEOLINOGNATHIDAE Fahrenholz

1929. Neol inognathidae, Ewing, Manual of External Parasites, page 133*
1936. Neolinoguathidae, Fahrenholz, Zeitschrift fur Farasi teukunde 9:1:56.

DESCRIPTION OF THE FAMILY. Auoplura in which the abdominal spiracles
are reduced to a single pair, this belonging to the eighth abdominal seg¬
ment. Abdomen membranous throughout except for the usual sclerotizations
of the genital region and the terminalia and except for the presence at
times of minute sclerotized points; almost devoid of setae except for a
pair at each lateral angle of segment eight and in the genital area. Legs
with the first pair small and slender with sleuder claw; second and third
pairs enlarged and stout, with stout claw. Antennae five- segmented, sexu¬
ally dimorphic. Thoracic sternal plate present but not apically free, di¬
vided into two longitudinal plates.

NOTES. This family was first recognized as an entity by Ewing who es¬
tablished it as the subfamily Neolinoguathinae of the family Haematopinidae .
It was elevated to family rank by Fahrenholz in 1936.

The single included genus with two species occurs on members of the fam¬
ily Macroscelididae of the Order Insectivora.

The members of this family are peculiar forms. The probabilities are
that their actual relationships are with the family Hoplopleuridae but they
are so peculiar that even if referred to this family they would have to be
maintained as a subfamily.

Genus NEOLINOGNATHUS Bedford

1920. Neol i nognat bus Bedford, Entomologist's Monthly Magazine (3)6:88.
1922. Neol inognathus, Ferris, Contributions Toward a Monograph of the Suck¬
ing Lice, Part 3:166.

GENERIC TYPE. Neol inognathus elephantuli Bedford, by monotypy. One
other species is included in the genus.

CHARACTERS. Without eyes. Antennae five-segmented, not sexually dimor¬
phic. Head fusiform. Thorax with the sternal plate not apically or mar¬
ginally free, divided longitudinally into two small plates. Anterior legs
small and with slender claw. Middle and posterior legs enlarged, somewhat
flattened, with stout claw. Abdomen membranous throughout except for the
ninth tergite and the genital sternites beset with small, sclerotized
points and entirely without setae except for a pair at each lateral margin
of segment eight. Abdominal spiracles present only on segment eight, these
noticeably enlarged.

NOTES. The two species referred to this genus may be separated by the
following key.

With a retrorse, tooth-like process on the dorsal (outer) distal angle of

the tibia of legs 2 and 3 . ELEPHANT! LI

Without such a tooth . PRAELAl ITS

257

Neolinognathus elephantuli Bedford
Figure 115

1920. Neolinognathus elephantuli Bedford, Entomologist's Monthly Magazine
(3)6:89-90; figure.

1922. Neolinognathus elephantuli Bedford, Ferris, Contributions Toward a
Monograph of the Sucking Lice, Part 3:166; figures 110-111.

HOSTS AND DISTRIBUTION. First described as from Elephantulus rupestris
( myurus) jamesoni at Onderstepoort, Transvaal, South Africa. Later re¬
corded by Ferris from Petrodromus tetradactylus and Nasilio brachyrhynchus
delameri from British Central Africa and Loita Plains, British East Africa.

Neolinognathus praelautus Ferris

1922. Neolinognathus praelautus Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 3:169; figure 111E and 112.

HOSTS AND DISTRIBUTION. Type from Elephantulus pulcher phaeus at Lime
Springs, British East Africa. Also from Elephantulus rufescens at Vor,
British East Africa.

Family PEDICULIDAE Leach

1817. Leach, The Zoological Miscellany 3:64.

1842. Denny, Monographia Anoplurorum Britanniae, page 1.

1880. Piaget, Les Pediculines, page 6 15-
1904. Enderlein, Zoologischer Anzeiger 28:136.

1929. Ewing, A Manual of External Parasites, page 141.

DESCRIPTION OF THE FAMILY. Anoplura in which the eyes are very definite¬
ly present externally as a pair of distinct lenses which are accompanied
each by a distinct spot of pigmentation that shows in uncleared specimens.
Antennae five-segmented, not sexually dimorphic. Legs variable in form,
either all practically the same or with the first pair small and slender
and the second and third pairs large and stout. Paratergal plates repre¬
sented on certain of the abdominal segments by a sclerotization which covers
the apex of lateral lobes of the abdomen and which never has any part free
from the body wall, although at times with lateral lobes. Abdomen other¬
wise membranous except for the usual terminal and genitalic plates and
small tergal plates in the male. Female with well-developed gonopods on
the eighth segment but with no definite indication of the gonopods of seg¬
ment nine present. Genitalia of the male with the parameres fused basally
with the aedeagus.

NOTES. As here understood, this family — which at one time included all
the Anoplura — is reduced to two genera that are here considered to involve
not more than four unquestioned species. Ewing has held that the genus
Phthirus should also he removed from it, leaving only the genus Pediculus
in the family. While an argument can he made for this step it is not here
accepted.

A step is taken, however, which undoubtedly will not meet with general
approval, but which is supported by the facts of morphology. That is& to re¬
move the genus Pedic inus from the Pediculidae. Those who base their concept
of the classification of the Anoplura upon the relationships of the hosts —
real or supposed— will object to this procedure, but the fact remains that,
considering the question from a morphological basis, the genus Pedic inus
appears to have very little to do with Pediculus, being apparently more
closely related to the members of the family Hoplopleuridae as here under¬
stood. The question is discussed at length in connection with Pedicinus.

The two genera remaining in the Pediculidae can be separated from each
other so readily that no key for their differentiation is here presented.

258

/ 1 i V,*

I'XAv^J’^v v

f" t 1 ♦ t

abdominal ornamentation
Neolinognathus elephantuli Bedford

female or nit alia

thoracic sternal i>late

2nd or 3rd clam

i » f

r??*1

^ i *

1 T y i i *•
f Y > f' < >

r I I » V’ *

Vj ;

/ . v\'-:r,s;

V Vv* * * »*•!'«

;> i, ~ ' » >v’

' ' V >\v 0\

'■f '#yif

I)/;;-.:;-: ;

r *• -f r, V , *» s - V4-

y/.;

>•“ . .. •

Genus PEDICULUS Linnaeus

It would be utterly impracticable here to present a complete bibliography
of this genus. Consequently, only those references are given to which a
specialized student of the group may refer for his purposes.

1758. Pedlculus Linnaeus, §ystema Naturae, Edition X, page 610.

1926. Pediculus, subgenus Parapediculus Ewing, Proceedings of the United
States National Museum 68: Article 19: 7.

1926. Pediculus, subgenus Paenipediculus Ewing, Proceedings of the Biolog¬
ical Society of Washington 45:117-

1935- Pediculus, Ferris, Contributions Toward a Monograph of the Sucking
Lice, Part 8:534.

1938- Pediculus, Ewing, Journal of Parasitology 24:13.

GENERIC TYPE. The type of Pediculus is Pediculus humanus Linnaeus. The
type of the proposed subgenus Parapediculus was stated by Ewing to be
Pediculus consobrinus Piaget. The consequences of this latter type selec¬
tion will be discussed in the notes which follow. The type of the proposed
subgenus Paenipediculus was stated by Ewing to be Pediculus simiae Ewing,
which is here considered to be a synonym of Pediculus schaffi Fahrenholz.

CHARACTERS. Anoplura in which eyes are very distinctly developed. Legs
all of essentially the same size and shape, the tibiotarsal articulation
distinct, the claws slender. Margins of the abdomen more or less strongly
lobed, the lobes covered by the sclerotized paratergal plates which are not
at all or at the most but in part and then only slightly free from the body
at any point on their margins. Thorax with a distinct notal pit. Thoracic
sternal plate sclerotized, but with its margins not free from the body.
Dorsum of the abdomen in the female membranous or at the most with slightly
developed, sclerotized plates; that of the male usually with small tergal
plates, the surrounding derm not minutely wrinkled. Spiracles present°in
normal position on the abdomen, six pairs being present, these all enclosed
within the borders of the paratergal plates. Male with the genitalia bear¬
ing very' small parameres which are united basally with the pseudopenis.

HOSTS AND DISTRIBUTION. Occurring on members of the Order Primates, es¬
pecially on man, the chimpanzee, and the New World monkeys of the family
Cebidae. There are records of its occurrence on gibbons.

It may be stated at the outset that the two supposed subgenera, Para¬
pediculus and Paenipediculus are here categorically rejected. The author
of these two subgenera, himself, indicated (1938) doubts as to the justifi¬
cation of the first of these but suggested that the second might well be
raised to the rank of a genus. That anything at all is to be gained by the
recognition of either remains to be demonstrated. In the case of Para¬
pediculus we have a question arising from a misidentified generic type.
The type of this proposed subgenus was definitely stated to be Pediculus
consobrinus Piaget, a species that has been shown/ on the basis of an exam¬
ination of the sole remaining specimen in the Piaget Collection at the
British Museum, to be Pediculus humanus. Ewing, however, had before him
when he named this subgenus specimens which were not this species and he
had never seen this type specimen. If Parapediculus, as a name for the
lice of the New World Cebidae, is ever recognized the question as to the
status of its type species will need to be settled, but since it is here
rejected no time will now be spent on the question.

Key to Species of PEDICULUS

Only those forms are included in this key which in the writer's opinion
have a reasonable claim to be recognized as species.

260

Spiracles of altdominal segments 3~5 each borne within a very small, cir

cular, sclerotized area; occurring on chimpanzees . SCHAFFI

Not so . 2

2. Paratergal plates throughout clearly without evidence of lateral lobes;

occurring normally on man although at times to be found on monkeys of

the family Cebidae and on gibbons in captivity . HUMANUS

Paratergal. plates of at least some abdominal segments showing clear ev¬
idence of lateral lobing, both dorsally and veutrally . 3

3. Paratergal plates of abdominal segments 5-6 bearing strong lateral

lobes, both dorsally and veutrally; occurring on monkeys of the fami¬
ly Cebidae . MJOfflRG]

Paratergal plates of abdominal segments 5-1 with slight, but distinct,
evidence of lateral lobes, both dorsally and ventrally; occurring on
man and Cebidae in the New World tropics and on man in the southwest¬
ern Pacific area . PSEUDOHUMANUS

In presenting the following review it has seemed desirable to consider
the species by host group rather than alphabetically.

Pediculus humanus Liunaeus
Figures 116, 117, 118, 119

The literature on this species is very extensive, but for the most part
is not pertinent in considering the taxonomy of the species. Only those
references are cited which are of importance in establishing synonymy or in
presenting evidence concerning the status of the various forms that have
been described.

17f>8. Pediculus humanus Linnaeus, Systema Naturae, Edition X, page 610.

(The original description of the species, containing no indication
that the head and body lice were considered to represent varieties. )
1761. Pediculus humanus Linnaeus, Linnaeus, Fauna Suecica, Edition 2, page
475. (The beginning of the controversy concerning the two varie¬
ties. "Qui in vestimentis victitat ab eo, qui in capite vivit,
non differt ut species, sed tantum varietas.")

1767. Pediculus humanus Linnaeus, Linnaeus, Systema Naturae, Edition XII,
page 1016. (The two supposed varieties are designated as 1 and 2,
respectively, for the head louse and the body louse and character¬
ized thus: "Varietas Capitis durior, coloratior; Vestimentorum
laxior, magis cinerea.")

1778. Pediculus humanus Linnaeus, de Geer, Memoires pour servir a l'his-
toire des insec tes 7:67; Plate 1, figures 6, 7. (Here the terms
"capitis" and "corporis" are first employed. "II y a done une
difference palpable entre ces deux sortes de poux, et qui semble
indiquer qu'ils sont d'espece differente, a moins qu'on ne veuille
plutdt, comme a fait M. de Linne, les regarder comme deux variates.
Quoiqu'il en soit, on pourroit les distinguer par les demoninations
suivantes: (1) Pediculus (humanus capitis) cinereus, thorace ab-
domineque fascia interrupta nigra marginatus ; (2) Pediculus (hu¬
manus corporis) albidus, totus intmaculatus. ")

1803. Pediculus humanus Linnaeus, Latreille, In Nouveau dictionnaire d'his-
toire naturelle 18:403- (This reference not seen. According to
Nuttall, the name "humanus" is here definitely restricted to "le
pou du corps," which would constitute the first type fixation.)
1803. Pediculus cervicalis Latreille, In Nouveau dictionnaire d'histoire
naturelle 18:403- (This reference according to Nuttall. The name
is indicated as applying to "le pou de tete.")

1805. Pediculus nigritarum Fabricius, Systema Antliatorum, page 340. (Es-

261

Pediculus humanus Linnaeus, the typical form called capitis

Figure 116

10-.C tablished for lice having their "habitat in nigritarum corpore. ")

1816. Pediculus ni£rescens von Olfers, De vegetativis et animatis corpor¬
is5 in corporibus animatis reperiundis commentarius, Part 1, pare
81 . 13

1816. Pediculus albidior von Olfers, De vegetativis et animatis corporibus
in corporibus animatis reperiundis commentarius. Part 1, page 81.
(Merely a new name lor the body louse.)

1816. Pediculus pubescens von Olfers, De vegetativis et animatis corpor¬
ibus in corporibus animatis reperiundis commentarius, Part 1, page
81. (Merely a new name for the head louse.)

1818‘ pp'ilJLulu? vesttmenti Nitzsch, German's Magazin der Entomologie 3;
306. (New name for the body louse.)

1824. Pediculus tabescentium Alt, De Phthiriasi, page 7.

1834. Pediculus capitis de Geer, liumieister, Genera^lnsectorum, Rhynchota,
Order 1, Tribe 1, Family 1, Species 1.

1880. Pediculus consobr inns Piaget, Les Pediculines, pa<re 626: Plate 61
figure 4. (Doubtfully described as new. )

262

Pediculus humanus Linnaeus, abdomen of male

Figure 117

263

Pediculus mjobergi Ferris, from Ateles dariensis .genitalia of female

Pediculus humanus Linnaeus, genitalia of female

Figure 118

264

corporis capitis

typical form typical form

Pediculus.paratergal plates

rnrfiN'v

pseudohumanus
from man,
Marquesas Islands

mjobergi

from

Ateles dariensis

Figure 119

1911. Pedtculus capitis de Geer and Pediculus capitis vestimenti Nitzsch,

Neumann, Archives de Parasitologie 14:410-413- (Neumann concludes
that the body louse is but a variety of the head louse and assigns
to it, contrary to all rules of nomenclature, the name combination
given above. He concludes also that Pediculus consobrinus Piaget
is identical with Pediculus capitis.)

1912. Pediculus capitis de Geer and Pediculus corporis de Geer, Fahrenholz,

Jahresbericht des Niedersachsischen zoologischen Vereins zu Han¬
nover 2-4:2-12; text figures 1-7; Plate 2, figures 16-19; Plate 3,
figures, 1-4. (Maintains the distinctness of head and body lice.)

1915. Pediculus corporis niiritarum Fabricius, Fahrenholz, Zeitschrift fUr
Morphologie und Anthropologie 17 : 596-597 ; text figure 1. (Assumes
to recognize this form on the basis of a single specimen.)

1915. Pediculus capitis aniustus Fahrenholz, Zeitschrift fur Morphologie
und Anthropologie 17:597; text figure 2; Plate 21, figure 1. (From
Japanese. )

1915. Pediculus capitis maculatus Fahrenholz, Zeitschrift fur Morphologie
und Anthropologie 17:598; text figures 3, 4; Plate 21, figures&2,
3. (For the head louse of African negroes.)

1915. Pediculus corporis marginatus Fahrenholz, Zeitschrift fur Morphologie

und Anthropologie 17:599. (The body louse of Japanese.)

1916. Pediculus humanus Linnaeus, Fahrenholz, Zoologischer Anzeiger 47: 269-

271. (Points out the proper application of the name "humanus, "
and gives data on literature.)

1916. Pediculus friedenthali Fahrenholz, Archiv fur Naturgeschichte, Ab-
teilung A, 81:11:2; text figures 1, 2; Plate, figure 1. (For a
louse from Hylobates mulleri. Date of issue indicated as July.)
1916. Pediculus oblongus Fahrenholz, Archiv fur Naturgeschichte, Abteilung
A, 81:11:15; text figure 14. (Fora louse from Hylobates syndacty-
lus.) (Not Pediculus oblongus Geoffroy)

1916. Pediculus humanus marginatus Fahrenholz, Zoologischer Anzeiger 48:

87 . (This apparently was intended as a preliminary diagnosis to
appear before the description in the reference cited above which,
however, has priority. Date of issue indicated as October.)

1916. Pediculus corporis aniustus Fahrenholz, Zoologischer Anzeiger 48:88.
(The same note applies.)

1916. Pediculus capitis maculatus Fahrenholz, Zoologischer Anzei rer 48'88
(The same note applies.)

1916. Pediculus friedenthali Fahrenholz, Zoologischer Anzeiger 48 :88 . (The
same note applies. The intended later description ^f this species
apparently has priority of three months.)

1916. Pediculus oblonius Fahrenholz, Zoologischer Anzeiger 48:88. (The
same note as for the next preceding species applies.)

1916. Pediculus humanus chinensis Fahrenholz, Zoologischer Anzeiger 48:87.

(Preliminary description of the body louse of Chinese.)

19 1 / . Pediculus humanus chinensis Fahrenholz, Fahrenholz, Mitteilungen aus
dem zoologischen Museum zu Hamburg (Beiheft zum .Jahrbuch der Ham-
burgischen Wissenschaftlichen Anstalten [2] ) 34:2, 6; text figure
1. (Definitive description.)

1917. Pediculus capitis maculatus Fahrenholz, Fahrenholz, Mitteilungen aus

dem zoologischen Museum zu Hamburg (Beiheft zum Jahrbuch der Ham-
burgischen Wissenschaftlichen Anstalten [2] ) 34:2. (Records this
form from Negroes in Dutch Guiana.)

1917. Pediculus humanus Linnaeus, Nuttall, Parasitology 10:1-79. (An ex¬
tensive bibliography is presented.)

^•919. Pediculus assimilis Fahrenholz, Jahresbericht des Niedersachsischen
zoo.1 ogischen Vereins zu Hannover 5-10:27. (New name for Pediculus
oblonius, which was preoccupied.)

266

1919. Pedlculus humanus Linnaeus, Nuttall, Paras! tology 11 : 3^9- (Sy s
tematic position, syuouymy , and iconography. All the species of
Pedlculus thus far described are regarded as being probably syno¬
nyms of humanus.)

1919. Pedlculus humanus Linnaeus, Nuttall, Parasitolojp’ 11: 279-328 ; 27 text

figures; Plates 12-17. (Records of abnormalities, together with
consideration of hybridism between "capitis" and " corporis " and
other evidence tliat they constitute but races of a single species.)

1920. Pedlculus humanus Linnaeus, Nuttall, Parasitology 12:136-153- (On

Fahrenholz1 purported new species, subspecies, and varieties of
Pediculus . A scathing and well-justified criticism in which it
is, however, erroneously concluded that all the named forms of
Pedlculus belong to the same species.)

1924. Pedlculus capitis de Geer and Pediculus vestimenll Nitzsch, Freund,
Tiex'arztliches Archiv, Prag 4(A) :42; text figures 1-4. (Revives
the idea of the specific distinctness of head and body lice and
presents evidence intended to support this view.)

1926. Pedlculus ( Pedlculus ) humanus nlgritarum Fabricius, Ewing, Proceed¬
ings of the United States National Museum 68:19:16; text figures
1C° 2, 3C, 5. 6; Plate 2, figures 6, 7- (Revived for lice from
Negroes. )

1926. Pediculus ( Pediculus ) humanus angustus Fahrenholz, Ewing, Proceed-
ingsofthe United States National Museum 68:19:19- (While assuming
to identify this form on the basis of two specimens, the author
united with it Pedlculus capitis mariinatus Fahrenholz and Pedlrul us
humanus chinensls Fahrenholz.)

1926. Pediculus (Pediculus) humanus americanus Ewing, Proceedings of the
United States National Museum 68:19:20; text figures IB, 2, 3B ;
Plate 3, figures 9, 10, 11. (Established for lice from the heads
of Peruvian mummies.)

1933. Pediculus humanus americanus Ewing, Bequaert, Carnegie Institution
of Washington Publication Number 431, page 573- (From Maya Indi¬
ans, Yucatan and Guatemala.)

1936. Pediculus humanus americanus Ewing, Proceedings of the Helmintholog¬
ical Society of Washington 3:36.

1938. Pediculus pseudohumanus Ewing, The Journal of Parasitology 24:23;
figures 5a, 6a.

Review of the PEDICULI Ascribed to Man

The nomenclatorial history of Pediculus humanus begins with the tenth
edition of Linnaeus' "Systema Naturae" in 1758. Here he named the species
but trave no indication of recognizing more than one form. In his Fauna
Suecica" of 1761 he indicated the head and body infesting forms as varieties
and in the twelfth edition of the "Systema Naturae they were numbered as

varieties 1 and 2 respectively. ,

In 1778, de Geer named these varieties as Pedlculus humanus capitis and

Pediculus humanus corporis. We need not go into the nomenclatorial problem
as to which of these varieties should be called Pediculus humanus humanus,
in accordance with the present rules of nomenclature, other than to indi¬
cate that following the literature it appears that this name should be used
for the head louseT while Pediculus humanus corporis should be used for the
body louse, if the distinction between them is to he recognized in nomen¬
clature. , „ .. , . *>

In 1805, Fabricius named a Pediculus nitritarum trom Negroes and it the

form which ’seems actually to occur on Negroes is to be recognized nomencla-
torialiy it would appear that this name is available tor it.

In 1816, von Olfers named a Pedlculus nigrescens from Negroes and re-

named the head and body lice as pubescens and albidior respectively.

In 1818, Nitzsch used the name capitis for the head louse and employed
the name vestimenti for the body louse.

JD Son’ ProPosed the name Pediculus tabescentium for the body louse.

In 1880, Piaget named Pediculus consobrinus from a New World monkey of
ghe tenus Ateles, this bein^, according to the evidence of the one specimen
remaining in the Piaget collection, nothing more than Pediculus humanus.

Now the idea came to be generally accepted that the head louse and the
body louse constitute two distinct species, these being referred to under
various combinations of the names previously mentioned.

In 1911, Neumann concluded that these two forms can at the most be re¬
garded as only subspecies and employed the names Pediculus capitis and
Pediculus capitis vestimenti for them in complete disregard of the accepted
rules of nomenclature.

Fahrenholz, beginning in 1912, maintained the distinctness of the head
louse and the body louse as species and began the process of supplying each
presumed race of man with a subspecies or variety of each of them. He as¬
sumed to recognize Pediculus corporis nigritarum— on the basis of a single
specimen and named the varieties Pediculus capitis angustus and Pediculus
corporis marginatus from Japanese. In 1916, he named Pediculus humanus
chinensis from Chinese and added, also, the supposed species Pediculus
friedenthali and Pediculus oblongus, both from gibbons, and the species
Pediculus lobatus from a New World monkey of the genus Ateles. The name
oblongus being preoccupied, he later altered it to assimilis.

During the first World War the recognition of the importance of the lice
as the carriers of disease, most importantly typhus, led to a very' large a-
mount of study of them. Professor 6. H. F. Nuttall gathered a large amount
of material from various parts of the world and various races of man and
came to the conclusion that the lice of man constitute at the most two un-
stable races of a single species. Work done by Bacot seemed to show that
the head louse can be converted into the body louse experimentally, although
he did not demonstrate the converse of this. His experiments are open to
question in regard to the material that he employed, but he at least demon-
strated that the two forms will interbreed successfully. Nuttall concluded
that all records up to that time of supposedly distinct species of Pedicul us
on New World monkeys referred also to humanus, although this conclusion can
not now be supported. He vigorously— and from the point of view here held
quite justifiably— criticized the work of Fahrenholz, although in some re¬
spects his conclusions were erroneous.

Freund, ina series of papers (1924, 1925, 1927) maintained the distinct¬
ness of head and body lice as species.

In 1926, Ewing accepted the opinion that head and body lice of Europeans
constitute a single species, but clung to the opinion that they represent
subspecies. Although he was unable to accept Fahrenholz' assignment of
three forms to Japanese and Chinese, he assumed to recognize one^of these
forms and also accepted the Pediculus nigritarum of Fabricius for lice from
Negroes. He then, for his own part, added a supposed subspecies, Pediculus
humanus americanus, for lice from American Indians. In addition, he named
two supposedly new species for lice from New World monkeys and accepted two
already established names. ^

The result of alJ this naming stood at this time at about twelve names
tor the lice of man himself, two names for lice from gibbons, two names for
Lice of the chimpanzee— which are discussed elsewhere— and seven names for
theT in*?* new W°rid monkeys> whic6 "ill 6e discussed elsewhere.

In 1935, Perris reported upon his studies upon the large collection of
lice assembled by Professor Nuttall and material accumulated from other
sources. He was able to examine types or other authentic specimens of the
forms named by Fahrenholz and the type of Pediculus humanus americanus,

named by Ewing, as well as the solo remaining specimen from the type lot of
Piaget's Pedlcu l us consobr intis.

It muy as well be frankly stated that his conclusions were to a consid¬
erable degree influenced by disgust at what he had seen in the course of
this work and revolt against the methods that had been employed in the sys-
tematics of this group. It was clear that some forms had been named solely
upon differences in the method of preparation of specimens, upon supposed
differences of the utmost triviality which had not been checked against a
series oi specimens to determine normal variation, and upon a philosophy of
taxonomy which apparently adopted the concept that a species "is a specimen
which looks different from other specimens." It is entirely possible that
iu this revolt against such methods he may himself have gone too fax and in
his turn made errors that arose from too great conservatism, but he still
holds that the revolt was sound in principle.

In 1936, Ewing, in a paper dealing specifically with the lice of New
World monkeys, expounded the view that Piaget must have had two species in
his material of Pedlculus consobrinus, one from monkeys and one from man.
He recognized, however, that in selecting the sole remaining specimen from
the Piaget collection as type Perris had relegated this species to synonymy
with Pediculus humanus. In addition he named a new species, Pedlculus
pseudohumanus , which, while based upon lice from monkeys, also included
specimens from American Indians. It will be discussed especially in con¬
nection with the lice of the Cebidae.

This, then, iu brief is how the matter of the lice of the genus Pedlculus
occurring on man stands at the present moment.

The Problem of the PED1CULI of Man

This problem has been recognized and argued about for nearly a hundred
and fifty years, with still no satisfactory solution. Ferris (1935) has
presented the story in detail up to that date and it will here merely be
abstracted.

As early as 1761, Linnaeus, in his "Fauna Suecica," recognized the exis¬
tence of two forms of Pediculus on man, these being the head louse and the
body louse. In 1878, de Geer applied names to these forms, calling them
respectively Pediculus hrnanus capitis and Pediculus humanus corporis. Un¬
der our now accepted rules of nomenclature one of these forms should have
been called Pediculus humanus humanus. Tracing the matter out it appears
that the first restriction, at least by implication, was by Latreille about
1805, in such a manner that the name Pediculus humanus humanus should be
applied to the body louse and humanus capitis he used for the head louse,
if the two forms are considered sufficiently distinct to be worthy of sci¬
entific names.

Since our present rules of nomenclature were not well developed and not
widely followed until almost 1900, other names came to be applied to these
lice/ Thus, the names vestimenti, cervlcalis, and tabescentium , as well as
both capitis and corporis were variously employed and the name vestimenti
especially was much used for the body louse. Also, the idea developed that
the head louse and the body louse constitute two distinct species and it
was not until 1911 that this idea was challenged by Neumann. Since that
time it has been both supported and attacked. Fahrenholz and Freund espec¬
ially have supported the idea that two species are involved, while Nuttall
and Ferris have maintained the opposing point of view. It should here be
emphasized that at the bottom of this difference of opinion lies the funda¬
mental question of "what do we mean by species?" So important is that
question that a brief discussion of it and of the writer's point ol view
will be presented somewhat farther along.

Concurrently with the idea that two species are involved has gone also

269

the idea that different forms occur on the different races of man. As
early as 1805, Fabricius named a Pediculus ni£ritarum from Negroes and in
1816 von Olfers named a Pediculus nigrescens from the same source. In
1912, Fahrenholz not only maintained the distinctness of head and body lice
as species, but apparently set out to provide every race of man with its
own "variety" of each of these species and this was added to by Ewing, who
as late as 1936 named Pediculus humanus americanus from American Indians.

Now, in the light of the evidence afforded by some other quite clear ex¬
amples of the occurrence of two or more species of lice of the same genus
on hosts of the same species, the possibility must be admitted that we may
have to do with a similar situation in connection with the Pediculi of man.
Thus, leaving out of consideration the extraordinary' situation connected
with the biting lice oi the Order Hyracoidea, it appears that something of
this sort occurs in the sucking lice of the Hyracoidea. Three clearly dis¬
tinct species of the same genus of Anoplura occur on domestic sheep. Two
clearly distinct species of the same genus occur on rodents of the genus
Thryonomys . Apparently' two quite distinct species of Haematopinus occur on
zebras. So in the light of these considerations there is no a priori reason
to assume that a parallel situation could not occur in the case of the lice
of man. The question is merely as to whether or not it does.

It is a risky business for an entomologist to become involved in anthro¬
pology without any personal basis of knowledge and in the face of the very
considerable differences of opinion which exist among anthropologists.
However, there are certain ideas which seem rather widely to be accepted.
Thus it appears rather generally agreed that present-day Homo sapiens falls
into three broad groups which seem to be subspecies as that term is general¬
ly understood by mammalogists. Furthermore, it is to be recalled that up to
a relatively few thousand years ago there existed what is generally consid¬
ered to have been a distinct species of man, Homo neanderthalensis — if one
can untangle the weird nomenclatorial practices of the anthropologists— who
must certainly' have had contact with Homo sapiens and whose females may
very well have furnished sport and variety for the males of a conquering
race. Somewhat parallel situations are not unknown today. It must also be
recalled that there is evidence of other ancient forms of man regarded by
some anthropologists as distinct species, which doubtless were to°some de¬
gree contemporaneous with the ancestors of the living subspecies.

In fact, it appears that these three existing groups or subspecies,
which display' an apparently unlimited capacity for exchanging genes with
unimpaired fertility to unnumbered generations, offer one of the best ex¬
amples of the biological meaning of the word "species." Relatively pure
representatives ot each of these groups exist and if considered bv systema—
tists working from a few preserved specimens would almost inevitably be re¬
garded as belonging to distinct species. But there exist in the total pop¬
ulation of man every conceivable degree of variation and every conceivable
combination of the characters that mark these races as they presumably were
constituted in their original state of nature.

Thus man, as he exists today, is a species within the formula which
seems best to express the biological concept of that term as used in zool-
ogyr . That formula, as the writer has attempted to express it after a care¬
ful weighing of every word is this:

A species is a population, the members of which are parts of' a continu¬
ously interlinked genetic complex, which is separated from other such com¬
plexes by barriers or incompatibilities of genetic origin and which under
natural conditions, that is, the conditions which have been concerned with
evolution, maintains itself from its own genetic resources.

An enormous experiment lias been going on for many thousands of years, in
which man has been the experimental animal. In the course of that experi¬
ment we may assume that certain mutants appeared from an original common

270

stock. These mutants became geographically localized and distributed as
subspecies and minor groupings within the population of man.

Certain of these minor genetic groups, being perhaps more aggressive
than others, spread from their original centers, and since no genetic bar¬
riers existed betweeu them and the groups which they overcame or with which
they mingled, hybridization resulted and finally we have a world population
in which all the mutations are mingled in every combination and to such a
degree that the population as a whole is continuously genetically inter¬
linked. Nor do the hybrid offspring have to be renewed by continued hy¬
bridization in order to maintain themselves. In contrast to this we may
cite the case of the domestic mule, which must continually be re-established
by renewed crossings and which otherwise cannot maintain itself.

This would seem to be a reasonably objective statement of the conditions
which exist in the human population of the earth and of the basis for con¬
sidering man to constitute a single species.

Now it would seem probable that something of this sort may very well
have happened in connection with the Pediculi which have been the constant
companions of man and which have accompanied him in his original process of
diverging into subspecies and his later reunification by hybridization.

These Pediculi may very well have begun to develop into genetically dif¬
ferentiated forms upon the various species of man, for the past existence
of which there is evidence, and the various subspecies that still exist.
But as their hosts have intermingled with each other the opportunity for
the intermingling of the parasites also has occurred. Thus we would arrive
at a condition among the parasites which may rather closely parallel the
condition found among their hosts. A population has resulted which con¬
forms to the same formula as does the host population. Here and there rel¬
atively pure populations of the parasites may exist. It is conceivable
that certain forms may have inherited physiological as well as morphological
differences which would lead to some degree of segregation in the hybrids
in accord with the degree of inheritance of these physiological character¬
istics. But the population as a whole presents a picture which closely
parallels that presented by man himself.

It is the writer's belief, based upon an examination of many specimens
of lice from different races of man and from various parts of the world,
that the situation above described is that which actually exists in the
population of Pediculus on man.

Material at hand from Negroes in Africa and South America would offer to
a systematist working merely from a few specimens a real basis for the be¬
lief that a distinct species of Pediculus occurs on Negroes. Ten specimens
of this form compared with ten specimens of the characteristic "body louse"
of Europeans would almost inevitably lead to such a conclusion. These lice
from Negroes are very darkly pigmented, their bodies are very compact,
their length is scarcely more than one-half the length of the European body
lice. However, other specimens present in a lot from Negroes in Africa
show every degree of gradation into normal head lice of Europeans.

And so with a comparison between normal "head lice" and normal "body
lice" of Europeans. The body louse, in its most characteristic lorm, is
much larger than the typical head louse, is paler in color and differs mor¬
phologically in the fact that the parater^al plates of the abdomen do not
extend around the apex ot their abdominal lobes into the intersegmental
notch as is the case with the paratergal plates of typical head lice. But
every degree of variation among these forms exists.

There have been available a wide range of specimens from Europeans,
Eskimos, Hindus, Arabs, Negroes, .American Indians, Chinese, and so on.
From the point of view here adopted these constitute a single species,
within the definition oi the term explained above.

There remains the questions involved with the nomenclatorial procedure

271

which should be followed in dealing with this material.

Within this material certain rather well-marked forms can be recognized
and certainly some basis exists for the recognition of two or more groups
that might be called subspecies, within the meaning of that term as em¬
ployed by those who accept it on a biological basis. Actually, however,
onl;y a certain portion ot this material will fall within these subspecies.
The remainder consists of variants from the typical forms in such combina¬
tions that nothing more can be said of them than that they are Pediculus
humanus.

Under these conditions and until the whole problem can be submitted to
examination by experimentation the opinion is here held that nothin* what¬
soever is to be gained by naming a series of "varieties" or subspecies. In
fact, if this process is once started and carried to its logical conclusion
the number of named forms could be extended indefinitely. “ For example, a
lot is at hand from natives of Rennell Island, one of the Solomon Islands
group, that could certainly be named as a new species if we were to employ
criteria of the order of those used by Fahrenholz and Ewing. And when we
have carried this process of naming to its ultimate limits, what have we
gained? Nothing more than a series of names of forms which can be recog¬
nized only if a perfectly typical example is at hand, which will be rela¬
tively seldom. When the needed experimental work has been done it may
prove that some definite nomenclature can be supported, but there seems to
be no justification for complicating the nomenclatorial situation in ad¬
vance of such work.

The employment of the name Pediculus humanus Linnaeus to cover this pop¬
ulation as a whole, with the addition of the vernacular names head louse
and body louse for those forms when the occasion demands, would seem ade¬
quately to take care of the situation as it exists at present.

Review of the Purported Species of PEDICULUS from Gibbons

In the bibliography of Pediculus humanus there are included two names
which have been given to lice reputed to have been taken from gibbons,
which are Primates of the genus Hylobates. The two purported species are
Pediculus assimilis Fahrenholz and Pediculus friedenthali Fahrenholz.

The writer has earlier seen specimens of the first-named of these, "deter¬
mined by Fahrenholz, which came from Hylobates syndactylus in the Zoological
Garden at Berlin, but has not seen specimens of the second, which was de¬
scribed as from Hylobates miilleri without indication of locality.

The examination of specimens of assimilis and the description of frieden¬
thali indicate no reason whatsoever for the recognition of these species as
distinct from Pediculus humanus.

Review of the Purported Species of PEDICULUS from New World Monkeys

As will appear from the following discussion the question of the name to
be used for the characteristic louse of the New World monkeys, which belong
to the family Cebidae, is much confused. In the opinion here held there
is but one species, apart from the at present very dubious question of

Pediculus humanus Ewing, but if this be true what shall this species be
called/

Actually, not until the matter has been subjected to an extended inves¬
tigation involving an examination of a large mass of material, including
the types— such as still exist — and an experimental genetic study of all
the forms involved, will it be possible to arrive at absolution that may be
generally satisfactory. J

Leaving aside the name quadrumanus, for which no evidence of any sort
is— or is likely to become— available, the first possible name is Pediculus

mjobergi Ferris. But this is clouded by the deficiency of the original de¬
scription and b} the tact that the type — the whereabouts of which is un¬
known has not been re-examined. The ntuue lobatus Fahrenholz is entirely
unclouded, although the type is probably not now in existence.

Pending the tinul stud} which must be made, the writer is inclined to re¬
tain the use of the name Pedtculus mjobergi Ferris pending a final settlement.

Realizing fully that the questions concerning the species of lice on the
New World monkeys involve many differences of opinion and that future wor¬
kers may decide that certain of the purported species are valid, the fol¬
lowing review lists these names and their bibliographies separately in or¬
der to minimize future confusion as far as may be.

Pediculus atelophilus Ewing

1926. Pediculus ( Parapediculus) atelophilus Ewing, Proceedings of the
United States National Museum 68:19:9; figures 4A, 6-
1931- Pediculus ( Parapediculus ) atelophilus Ewin % Hinman, Parasitology 23:
488.

1935- Pediculus mjdbergi Ferris, Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 8:688; figures 318E, 319E, 322F, 3231,
328-332.

1938. Pediculus atelophilus Ewing, Ewing, The Journal of Parasitology 24:
26; figures 2, 6b, 6b.

HOSTS AND DISTRIBUTION. The type of this species was recorded as from
Ateles geoffroyi with unspecified type locality. Recorded at the same time
from skins of the same host taken in Costa Rica. Recorded by Ewing in 1938
from Ateles pan from Guatemala; from Ateles dar iensis from Panama; from
Ateles hybridus from the National Zoological Park at Washington, D. C.

There is at hand a considerable amount of material undoubtedly referable
to this name, including specimens from the material recorded by Hinman out
of a lot identified by Ewing from Ateles geoffroyi in captivity and others
from this host in Panama; numerous specimens from Ateles dariensis, Cebus
capuchinus, and Alouatta palliata, all from Panama from monkeys in captiv¬
ity; "ring-tailed monkey" from the Bronx Zoological Park in New York.

NOTES. No basis appears in all this material for the recognition of
more than one species. It is here held that all these specimens are cov¬
ered by Pediculus lobatus Fahrenholz.

Pediculus chapin'i Ewing

1926. Pediculus ( Parapediculus ) chapini Ewing, Proceedings of the United
States National Museum 68:19:13; figs. 2, 4b, 6; Plate 1, figs. 3 -4.
1935* Pediculus mjobergi Ferris, Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 8:588.

1938. Pediculus chapini Ewing, Ewing, The Journal of Parasitology 24:23;
figures 4a, 5c, 6c.

HOSTS AND DISTRIBUTION. The types were recorded as from Ateles ater
from the National Zoological Park at Washington. Specimens considered by
Ewing to be stragglers were later recorded by him (1938) from Ateles geof¬
froyi from the same place and from Cebus capuchinus from Panama.

NOTES. Unfortunately the only specimens at hand from the type host of
this supposed species are immature. In the absence of authentic specimens
it is not possible to assert that this species is identical with any other,
although in the opinion here held it is the same as atelophilus.

Pediculus lobatus Fahrenholz

1916. Pediculus lobatus Fahrenholz, Archiv fur Naturgeschichte , Abteilung

273

A, 81:11:16; Plate, figures 6, 7. (August)

1926. Pedi cuius ( Parapediculus) lobatus Fahrenholz, Ewing, Proceedings of
the United States National Museum 68:19:8; Plate 2, figure 5.1"
1935- Pediculus mjbbergi Ferris, Ferris, Contributions Toward °a Monograph
of the Sucking Lice, Part 8:588.

1938. Pediculus lobatus Fahrenholz, Ewing, The Journal of Parasitology 24:
29; Plate 2, figure 5-

HOSTS AND DISTRIBUTION. Recorded by Fahrenholz from Ateles rellerosus
from the Berlin Zoological Gardens. Ewing (1938) identified with this
specimens from Ateles paniscus from the National Zoological Park at Washing¬
ton and recorded specimens which he considered to be stragglers from Leon-
tocebus nigricollis at the same institution.

NOTES. The original description of this species is composed chiefly of
useless detail which aids not at all in recognizing the species and the ac¬
companying photographic illustrations do nothing more than indicate from
the form that the species is probably of the type of those known from New
World monkeys. Actually, comparing the illustration of the male given by
Fahrenholz with that given by Mjoberg for his Pediculus af finis {-mjobergi)
there is no more basis for the identification of a species on the basis of
one of these illustrations than of assuming it to represent some other
species.

Pediculus mjobergi Ferris

1910. Pediculus af finis Mjoberg, Arkiv for Zoologi 6:169, 258; text fig¬
ures 85, 151 ; Plate 5, figure 8. (Preoccupied)

1916. Pediculus mjobergi Terris, Proceedings of the California Academy of
Sciences (Series 4) 6:136. (May)

1935* Pediculus mjobergi Ferris, Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 8:588.

HOSTS AND DISTRIBUTION. The types were recorded from Ateles sp. in a
traveling menagerie. This constitutes the only positive record of the
species.

NOTES. The name mjobergi vtas proposed by Ferris to replace af finis of
Mjoberg which was preoccupied by an earlier Pediculus 'af finis. A question
might arise as to the priority of mjobergi and lobatus, both of which were
established in 1916. In 1913> Fahrenholz used the name lobatus without any
accompanying description and the name was not nomenclatorially established
until August, 1916. Unfortunately, the name mjobergi was established by
publication in May, 1916, to replace the preoccupied name af finis of Mjoberg
and therefore has priority in case any question arises in the future con¬
cerning these names.

If it be concluded that the lice of the New World monkeys represent but
a single species it apparently should be called by the name mjobergi.

Ewing (1936) has put forward the thesis that MjSberg's description indi¬
cates his specimens to have been Pediculus humanus . Actually, Mjobercr's
* pp0ns<T description and the very poor accompanying illustrations
offer little or nothing to demonstrate anything, one way or another. The
photograph of a male contains nothing more than a faint suggestion, which
can be magnified by a slight application of the imagination, into the char-
actenstic louse of the New World monkeys. Ewing has maintained that
Mjoberg s illustration of the egg indicates lumanus, but in fact the draw¬
ing is erroneous even for that.

Hie writer will concede that until and unless Mjoberif's types axe redis¬
covered the status of this name will have to be te Id in’ abeyance, although
on the basis of a theory of probabilities it is used in this work to cover
the entire list of names employed for the lice of Cebidae except as is
later indicated.

274

Pediculus pseudohuuutuus Ewing
Pi gu re 120

1938. Pediculus {Parupediculus) pseudohumanus Ewing, The .Journal of Para-
si tology 24:23; figures 3. 5a., 6a.

H0ST8 AND DISTRIBUTION. Type from a monkey, Pitchecla monachus. Also
recorded by Ew ing from another monkey, Cacajao rubicundus, from the National
Zoological Park at Washington. Also recorded by Ewing from an Indian at
Cohan, Guatemala.

NOTES. We have here a most extraordinary situation. The lorm which
Ewing described exists, without question, but its distribution is extremely
peculiar. Ferris (1935) mentioned the presence in his material of speci¬
mens from Central American Indians and from natives in the Marquesas Islands
in the south Pacific which show a sli dit lateral iobin of certain ol the

CD O

paratergal plates. This is the form that Ewing ascribes to his pseudo-
humanus and the illustration here given, based upon a specimen from the
Marquesas Islands, almost duplicates that given by him. It may be noted
that the specimens from the south Pacific all liave a noticeably larger num¬
ber of setae on the dorsum of the abdomen than do those from the New World.

The material at hand which appears to be covered by the name pseudo-
humcinus is as follows: from natives at the village of Kakahitau, Uapou,
Marquesas Islands; from natives at Hitiaa, Tahiti; from Indian hut at Santa
tinilia, Guatemala; from "dried head from Ecuador." Specimens from head of
Maya Indian, Xichel, Yucatan (from a lot one time identified by Ewing as
americanus) , and others from "natives," at Guayabilete, Panama, have the
lobing of the paratergites of segments 5~6 even more strongly developed and
approximating that to be seen in typical specimens of atelophilus.

Unfortunately, no specimens of this form are at hand from monkeys, but
since Ewing himself has recorded this supposed species from man, there
should be no objections to the records given above.

The name pseudohimanus is here recognized to cover a form which may pos¬
sibly be worthy' of recognition.

The Problem of the Lice of the Cebidae

This problem has been reviewed by Ferris (1935) and by Ewing (1926,
1938) , these authors coming to quite different conclusions. The general
story is briefly this:

In 1877, Murray ("Economic Entomology, Aptera, " page 3) described a
Pediculus quadrumanus which was said to have been taken from a captive mon¬
key of the genus Ateles. The species is entirely unrecognizable from the
description and inquiry' has led to the conclusion that if there ever were
any' types these are not now in existence. Whatever the possibilities may¬
be there is certainly no justification for dispossessing any later name in
favor of quadrumanus.

In 1880, Piaget ( "Les Pediculines, " page 626; Plate 51, figure 4) de¬
scribed a Pediculus consobrinus taken from Ateles pentadactylus. Museum of
Leyden, whether from a living or preserved animal not being indicated. No
indication was given as to the number of specimens examined, except for the
statement that no male was observed. Ferris (1935) reported upon the single
specimen from the Piaget Collection, which still exists at the British
Museum, and illustrated this specimen. It is in his opinion definitely not
separable from Pediculus humanus. There is no reason to suppose that this
specimen is not one of those which Piaget had before him but Ewing, entire¬
ly on the basis of a comparison of the figures given by Piaget and by
Ferris, has offered the entirely gratuitous assumption that Piaget had two
species in his material. Only some exercise of the imagination and a lirm
will to believe in the results of that imagination could lead to such a

275

Pediculus mjobergi Ferris, from Ateles dariensis

Figure 120

conclusion. The opinion is here maintained that we must accept the evi¬
dence from this one remaining specimen and place Pediculus consobrinus as a
synonym of Pediculus humanus. This is supported by the fact that othpr
specimens of what seems undoubtedly to be hLnus, Lcribed to monkeys of
this group, have been seen by the writer. y 1

to have19belnM'!»l;!rgfdeS0,'ibed “ Pedicul“s “Minis a louse which was said
to have been taken from some species of Ateles in a traveling menagerie

pr4eo,ccupied u was later changed to \jbberii by
Dn Unfortunately, the description given by Miobere is useless

and the illustration of the male given by him falls short of being actually

decisive. Not until the types have been re-examined — if they exist _ can

the question of what this species actually is be definitely settled. ' Perris
UJJ.)) was willing to accept the species, but Ewing (1938) considered the

^ °" the °f “ i^ticm of [he £

The next name available is Pediculus lobatus Fahrenholz (1916). The

276

mune mjdberfil has precedence of about three mouths over lobatus which, al
though tirst proposed in 1913> remained a nomen nudum until 1916. The
present writer has examined the types of lobatus and illustrated them
(19*1*1). This species is here held to be distinct from humanus on good mor¬
phological grounds. The illustration given by Ferris was based upon the
uncleared type specimen and represents that particular specimen as well as
possi ble.

In 1926, Freund recorded specimens from A tele s ater from the Leipzig
Zoological Garden and employed for them the name Pedlculus capitis forma
atelis. This is unrecognizable from the description.

In 1926, Ewing presented an extensive paper on the New World lice of the
genus Pedlculus (Proceedings of the United States National Museum 68, Ar¬
ticle 19) in which he presumed to recognize Pedlculus consobr tnus Piaget
and Pediculus lobatus Fahrenholz and named two new species, Pedlculus
atelophilus and Pediculus chapini from New World monkeys.

In 1935, Ferris (Contributions Toward a Monograph of the Sucking Lice,
Part 8) reviewed the whole question of the species of Pediculus and came to
the conclusion that there is but one valid species on the family Cebidae —
exclusive of occasional occurrences of what seems definitely to be Pediculus
humanus. For this species he employed the name Pediculus mjdberfi Ferris.

Iu 1938, Ewing (The Journal of Parasitology 24:13- 33; figures 1-6) pre¬
sented another paper on the lice of the New World monkeys, maintained the
validity of the two species previously described by him and named another,
Pedlculus pseudo humanus.

A considerable amount of material has been available in connection with
the present work and on the basis of this material the writer still main¬
tains the opinion that — apart from Pediculus humanus, which apparently can
transfer to and survive upon members of the Cebidae, and Pediculus pseudo-
humanus — there is but one actual species which seems to be normal to the
New World monkeys of the family Cebidae. That species seems to be distinct
from Pediculus humanus.

Pediculus sch&ffi Fahrenholz
Figure 121

1910. Pediculus schdffi Fahrenholz, Jahresbericht des Niedersachsischen
zoologischen Yereins zu Hannover 1:57; Plate 1, figures 1-3; Plate
3, figures 1, 2, 4. 5; Plate 4, figures 2, 6.

1915* Pediculus schdffi Fahrenholz, Fahrenholz, Archiv fur Naturgeschichte,
Abteilung A, 81:11:1.

1919. Pediculus humanus race schdffi Fahrenholz, Nuttall, Parasitology 11:
336.

1932. Pediculus (Paenipediculus) s imiae Ewing, Proceedings of the Biologi¬
cal Society of Washington 45:117.

1933- Pediculus (Paenipediculus) simiae Ewing, Ewing, Proceedings of the
Biological Society of Washington 46:168; figure 2c.

1935- Pediculus schdffi Fahrenholz, Ferris, Contributions Toward a Mono¬
graph of the Sucking Lice, Part 8:599; figures 322H, 333> 334 -
H0ST5 AND DISTRIBUTION. Recorded by Fahrenholz from chimpanzee, Pan
(= Simla) troilodptes from the Zoological Gardens in Hamburg, and by Ewing
from the same host from the London Zoological Gardens.

NOTES. The accompanying illustrations, which are those given by Ferris
in 1935, were made from specimens which are apparently a part of the same
lot on which Ewing based his name simiae.

The type of schdffi was not seen by Ferris, but the description is suf¬
ficient to indicate clearly that only one species is involved.

Ewing has based the subgenus Paenipediculus upon this species. This is
not here accepted.

277

Pediculus schaffi Fahrenholz

Figure 121

278

Suggestions for Future Research

In tiie species of Perlicul us occurring on man and on the New World monkeys
ot tiie tamily Cebidae there appear a number of problems which should be ap¬
proached t rum a biological point ot view and submitted to examination by
the methods ot experimental biolo^. It is evident that no solution can be
achieved merely by the contemplation of preserved specimens.

First of all there is the problem of the lice of man himself. While
this has been submitted to a certain amount ot experimental investigation
those experiments are not entirely free from criticism and they are not
sufficiently extensive to permit definite biological conclusions. Thus the
experiments of Bacot seemed to show that it is possible to convert the head
lice of man into forms having the appearance of body lice, but the converse
of this experiment was not described. The experiments of Bacot showed a
complete hybridization of what he considered to be head lice with what he
considered to be body lice. But in the offspring of these hybridizations
there appeared a large number of abnormalities that have been considered by
Keilin and Nuttall (Parasitology 11:279-328; text figures 1-26; Plates 12-
17. 1919). Some occurrence of abnormalities seems to appear in "wild" pop¬
ulations, but that shown in this material is relatively large. Certain of
the abnormalities are perhaps genetic. The suspicion occurs that these ab¬
normalities may have in part resulted from the hybridization of genetically
disharmonious individuals. Also, as the work was done in Egypt there is a
suggestion that perhaps the supposed head louse employed represented a
strain that is possibly farther removed from typical humanus than are typi¬
cal head and body lice from each other. Attention has been called to the
existence on African Negroes of a form which is so different from the typi¬
cal head lice of Europeans that if specimens of both extremes were placed
side by side the conclusion would almost inevitably be reached that we are
dealing with different species. It is conceivable that Bacot had this ex¬
treme form actually in his possession and that the experiments on hybridi¬
zation were in part based upon it. If so, these abnormalities might be ex¬
plained on this basis.

Nuttall records experiments (Parasitology 11:345. 1919) which led him to
the conclusion that the two supposed subspecies, capitis and corporis, rep¬
resent merely two unstable races of a single species. He indicates that
the head louse can be converted into the body louse by environmental condi¬
tions. But it remains to be shown that the changes of color and size re¬
corded by him also extend to the apparent differences in structure which
are revealed by typical specimens of either race.

So it appears that this experimental work should be done again, with
proper precautions as to the status of all the material employed and proper
attention to all changes that might be ascribed to changes in environment
and proper consideration of all genetic factors that might be involved.

But this problem of the lice occurring normally on man is by no means
all of the problem. There remains that of the lice of the Cebidae.

The Cebidae are supposed to be rather far removed from the remainder of
the Primates. Simpson (The Principles of Classification and a Classifica¬
tion of the Mammals, "Bulletin of the .American Museum of Natural History,"
Volume 85, 1945) divides the Primates into three superfamilies — Ceboidea,
Cercopithecoidea, and Hominoidea. There occurs on the Cercopithecoidea a
group of lice, the genus Pedicinus, which is far removed from the lice of
the Hominoidea. Why is it that the lice of the Ceboidea seem to be so
closely related to those of the Hominoidea that they belong in the same
'■enus? The Ceboidea are considered to be connected with the remainder of
the Primates only at the base of the stem of the Order. Is it to be sug¬
gested that the members of the Ceboidea have retained the genus Pediculus
from the time when the various stocks diverged, or is it to be supposed

279

that they have received them from man in the time that man has been in the
New World? How close are they genetically to the lice of man?

And yet it is supposed that the lice of the Cehidae are sufficiently
plastic in an evolutionary sense, to have formed at least four species, al¬
though at the best the characters assigned to those species are extremely
unconvincing. What are the genetic relationships of these supposed species
to each other? It certainly appears on morphological grounds that the New
World monkeys have upon them a species that can he easily distinguished
from the lice of man, but even this is somewhat clouded.

And, to add the last hit to this tantalizing problem, what is the situa¬
tion in regard to the supposed Pediculus subhumanusl Here is a form that
is supposed to occur both on New World monkeys and upon man. More than
that, it occurs not only upon man in the region where these monkeys occur
naturally but what is apparently the same form occurs on man in the far
distant South Sea Islands? What are the genetic relationships of the lice
oi this form in the New World to those on man in the New World and to those
of the South Pacific?

These are problems for which the solution is to be obtained — if at all—
by experimental methods. Speculation on the basis of the present known
specimens might easily lead the systematist to the endorsement of some idea
that this form developed on man and monkeys after man appeared in the New
World and then passed to the South Pacific following the migration of man
from the New World to that area, supporting the views of the 'author of Ron-
Tiki. It is futile to speculate upon such matters when all that is known
of the lice is actually but a relatively few specimens.

The author holds that this is a problem which is eminently suited to an
approach by experimental methods. Should some one gather together at one
place representatives of typical head lice and bod} lice of Europeans, typ¬
ical lice from African Negroes, lice of the type of Pediculus pseudohumanus
from both man and monkeys from Central America and from the South Seas, and
typical lice from a wide assortment of New World monkeys, subjecting them
to careful genetic experiments, we might at last put an end to the other-
& n concerning thei r relationships to each other. More
than that, we might possibly have an answer that would apply to other cases
involving parasitic insects and thus to a general biological problem.

Until that has been done the problems associated with the genus Pediculus
will remain for mere opinion and for speculation.

Genus PTHIRUS Leach

1815- Pthirus Leach, Edinburgh Encyclopaedia 9:77.

1817. Pthirus, Leach, Zoological Miscellany 3:365.

1935- Phthirus, Ferris, Contributions Toward a Monograph of the Sucking
Lice, Part 8:602.

GENERIC TYPE. Pediculus pubis Linnaeus.

CHARACTERS. Pediculidae with distinctly five-segmented antennae which
are not sexually dimorphic. Anterior legs very slender, with slender claw;
middle and posterior legs very large and stout, with stout claw; the coxae
of all the legs set at the extreme margins of the thorax and thus the mem¬
bers of corresponding pairs set far apart. Thorax very wide, forming the
greater part ol the body, without a sternal plate and without a notal pit,
the sclerotization of the apparent notum confined to the lateral areas.*
Abdomen relatively small., as broad basally as the posterior part of the
thorax and tapering somewhat posteriorly, membranous except for the projec¬
ting lateral, segmentaJly arranged tubercles. These lateral tubercles
sclerotized and prominent, there being one at the margin on each side of
segments 6-8. Spiracles of these sejjments borne slightly removed from the
base of tubercles, those of segments 3-4 being crowded close to those of

280

sequent, tour, somewhat displaced toward the median line and being the only
cleax- evidence of these segments. The spiracles are of a peculiar fonn,
the atrium be in^r much enlarged, conical, with the base of the coue at the
inner end, the walls marked by parallel, longitudinal ridges. Female with
well-developed gonopods on segment eight and with a very large spennatheca.
Male with small, pointed paramei*es which articulate near their apices with
a pair ot small sclerotized points; no endomeral plate present and no scle-
rotized peuis, the gonopore being surrounded merely by a partially sclero-
tized ring.

NOTES. This genus was employed by Ewing (1929) as the type of the family
Phthiridae, it beiug the only genus of this family. Other woi’kers have rec¬
ognized it as the type of a subfamily, Phthirinae, of the Pediculidae. It
is a peculiar form, indeed, yet its relationship seems to be with Pedieulus
and to place it in a separate family is to obscure this relationship. Its
peculiar position is here recognized to the extent of accepting the subfam¬
ily Pthirinae of the Pediculidae.

Pthirus pubis (Linnaeus)

Figures 122, 123, 124

1758. Pedtculus pubis Linnaeus, Systema Naturae, Edition X, page 611.

1815* Pthirus inguinal is Leach, Edinburgh Encyclopaedia 9:77-
1816. Pediculus ferus von Oilers, De vegetativis et animatis corporibus in
corporibus animatis reperiundis commentarius, page 83- (Definite¬
ly a synonym of pthirus pubis (Linnaeus)

1904. Pthirus pubis (Linnaeus), Enderlein, Zoologischer Anzeiger 28:136;
figures 10, 11.

1918. Phthirus pubis (Linnaeus), Nuttall, Parasitology 10:383; figures 1,

3-5, 7-9.

1935- Phthirus pubis (Linnaeus), Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 8:603; figures 335> 336, 337.

1935* Phthirus chavesi Escomel and Velando, Cronicas de Medicina (Lima,
Peru) 52:335-

1936. Phthirus pubis (Linnaeus), Bedford, Onderstepoort Journal of Veteri¬
nary Science and Animal Medicine 7: 105-
19 39 - Phthirus pubis (Linnaeus), Buxton, The Louse, page 93; figure 25.

HOSTS AND DISTRIBUTION. Occurring on man in many parts of the world.
Bedford (1935) has recorded it from a chimpanzee from the French Congo.
There have been occasional records of its occurrence on dogs, but no infor¬
mation exists that these are anything more than incidental.

NOTES. The original description of Phthirus chavesi Escomel and Velando
has not been seen in connection with this work, but the species was recorded
as occurring in the eyebrows of man and since this is well known for pubis
there is no reason to suppose that any other species was involved.

The account by Buxton (1939) sums up what is known concerning the biology
of the species.

Pthirus gorillae Ewing

1927. Phthirus gorillae Ewing, Proceedings of the Entomological Society of
Washington 29 : 120 .

1933. Phthirus gorillae Ewing, Ewing, Proceedings of the Biological Soci¬
ety of Washington 46:170; figures lc, 2b.

1935. Phthirus gorillae Ewing, Ferris, Contributions Toward a Monograph of
the Sucking Lice, Part 8:608.

HOSTS .AND DISTRIBUTION. Described from eggs and first-stage nymphs taken
from skins of Gorilla berengeri in the Belgian Congo.

NOTES. In the absence of any information other than that given by its

281

Pthirus pubis (Linnaeus)

Figure 122

describer, nothing can be said concerning this species other than that it
presumably demonstrates the occurrence of a species of Phthirus onagorilla.

Unplaced Names

Th® following species names have been proposed, but the original descrip¬
tions are unavailable or are so inadequate that the genera to which they be¬
long cannot be determined. Perhaps future workers may be able to clarify

them either through the rediscovery of their tj^pes or through circumstan¬
tial evidence.

Pediculus aquaticus Pontoppidan

1763. Pediculus aquaticus Pontoppidan, Danske Atlas 1:69Q.

The original description of this species has not been available and I
know nothing of it except a bibliographic reference by Fahrenholz.

Pediculus clavicornis Nitzsch

1864. Pediculus clavicornis Nitzsch, Zeitschrift lur die gesamten Natur-
wissenschaften 23:32.

1874. Haematopinus clavicornis (Nitzsch), Giebel, Insecta Epizoa, pa^e 37
Perhaps a species of Hoplopleura, if we may judge by Giebel' s redescrip-
tion, which was based upon a single female from Meriones sp., from Africa

282

(UltflllUl

male abdomen

male yenitalia

Pthirus pubis (Linnaeus) .details Figure 123

Pediculus spiculifer Gervais

1844. Pediculus spicultfer Gervaris, In Walckenaer's Histoire naturelle des
insectes apteres 3:302.

This is apparently a species either of Polyplax or Hoplopleura. It was
recorded as from "Hus barbarus" from Algiers. This host is now called
Lemniscomy barbarus.

Polyplax miacantha Speiser

1905. Polyplax miacantha Speiser, Centralblatt fur Bakteriologie, Originate
38:318- (The figure does not belong with this.)

193?. Polyplax miacantha Speiser, Ferris, Contributions Toward a Monograph
of the Sucking Lice, Part 4:237.

Recorded as from a small rat with very thick, spine-like hair, from
Salomona in Abyssinia, in the collections of the Museum at Konigsberg in
Prussia. The original description is entirely inadequate for placing the
species even generically, although it is probably either a Polyplax or a
Hoplopleura.

283

1st clam

Pthirus pubis (Linnaeus), details

2nd or 3rd claw

Figure 124

284

ERRATA

The following references have been omitted from the preceding text:

Haematopinus asini (Linnaeus)

1865. Haematopinus equi Simmonds, Journal of the Royal Agricultural Soci¬
ety of England (2)1:60.

1880. Haematopinus astni, variety colorata Piaget, Les Pediculines, page
654.

Schizophthirus pleurophaeus (Burmeister)

1874. Haematopinus leucophaeus Giebel, Insecta Epizoa, page 37.

Haemodipsus lyriocephalus (Burmeister)

1864. Pediculus lyriceps Nitzsch, Zeitschrift fur den gesamten Naturwis-
senschaften 23:24.

285

CHAPTER VII
Host List

The preparation of a dependable list of the mammal species from which
Anoplura are known is not entirely a simple matter. In the first place,
some of the hosts have been recorded merely by their vernacular names and
these are not always sufficient to fix the record to a particular animal
species. Then there have been a certain number of definite errors by the
entomological recorders and there have been errors in the transcription of
names from mammal specimens and also in regard to the identifications of
mammals from which Anoplura have been taken in zoological gardens where the
mammals may have been misid entitled. But the greatest source of difficulty
arises Irom the tact that in many instances mammalogists themselves are not
or have not been in agreement either as to the nomenclature or even the
zoological status of the mammals.

Furthermore, there is no one master list of the mammals of the world
which we may utilize as a source of reference. The catalogue of the mam¬
mals of the world, which was published many years ago by Trouessart, has
long been obsolete and nothing has been published that takes its place. We
are therefore compelled to rely upon a series of lists, some regional, some
applying only to the mammals of a limited group, and where these lists
overlap they are not always in agreement either as to nomenclature or as to
i status °f some particular species. Under such circumstances an
additional source of error is added, since the compiler of a host list, in
pursuing names through these various lists of mammals, may himself go wrong
and commit errors o f his own. All this should be understood by future
students of the parasites.

In preparing the following list the following sources have especially
been utilized.

First of all appreciation needs to be expressed of the work of Mr. G. H.
E. Hopkins, who has published a host list of the Mallophaga and Anoplura of
mammals. This was very carefully done and has aided greatly in checking
the records of the names which had already independently been arranged in
such a list for this work. Hopkins' paper is replete with notes, many of
which are beyond the scope of the list here presented. The reference to
his work is as follows.

HOPKINS, G. H. E. The host association of the lice of mammals "Pro—

^d!nfS -i0£nthe Zool°gical Society of London, " Volume 119, Part 2, pages
387-604, 1949. ^

Fortunately, a large percentage of the known species of the Anoplura
occur on rodents and for this group we have available the great work of
Ellerman which relatively simplifies the preparation of this portion of the
host list.

ELLERMAN, J. R. "The Families and Genera of Living Rodents." Tliree vol¬
umes. Published by the British Museum (Natural History) . Volume 1, issued
June, 1940; Volume 2, issued March, 1941; Volume 3, issued March, 1949.

for the so-called "Ungulates," the only work that the writer has been
abfe to find that is at all comprehensive is that of Lydekker, in which
although it is scarcely recent, one can at least find the names that have
been employed for most of the mammals of this series, very' few havin '- been
described since its publication.

/.I. 1 1,1 f V II ,1- 13 1 ^ Mamina Ls in the British Museum

(NaturaJ History). five volumes. Published by the British Museum (Natur¬
al History). Volume 1, issued 1913; Volumes 2 and 3, issued 1914; Volumes

286

4 and 5> issued 1915*

For the Primates the large work of Elliot appears to be the only avail¬
able general reference although the entire group seems still to be in a
cout'used condition nomenclatorially . Thus, Simpson (1945, page 181) re¬
marks that "most Primates have alternative names and hardly any two stu¬
dents use the same nomenclature for them." Simpson's work (cited below)
has been employed for the names of genera and larger groups and Elliot's
for the names of species.

ELLIOT, D. G. A review of the Primates. "Monographs of the American
Museum of Natural History," Volumes 1-3, 1913.

Of regional lists the most useful are the following. The first named of
these is especially important because of the large number of mammal species
involved.

ALUU , G. M. A checklist of African mammals. "Bulletin of the Museum of
Comparative Zoology at Harvard College," Volume 83, 1939.

MILLEK. G. S. "Catalogue of the Mammals of Western Europe (Europe exclu¬
sive of Russia) in the Collections of the British Museum." Published by the
British Museum. 1912.

MILLER, G. S. List of North American laud mammals in the United States
National Museum. "Bulletin of the United States National Museum" 79, 1912.

For the higher categories of the mammals the recent work by Simpson is
here relied upon.

SIMPSON, G. G. The principles of classification and a classification of
the mammals. "Bulletin of the American Museum of Natural History'" 85, (350
pages) 1945.

For a few forms such as the Pinnipedia it has seemed necessary to accept
all names as given by the recorders of the lice.

Since the hosts of all the comparatively few sucking lice known from
South America are, with few exceptions, rodents, they are cared for in the
work of Ellerman, as are also the rodents of Asia. The Carnivora, except
for the Canidae and the Pinnipedia, have no sucking lice and the few names
here needed cause no special difficulty. The Insectivora are cared for
chiefly in the list of African mammals given by Allen ahd in various other
regional lists.

Names of hosts are usually given in the review of species of lice in the
text as given by the original recorders of the lice, but the correct names
— as far as they can be determined — will be found in the host list.

One other source of possible error in this list should be noted. This
begins with the collecting of the mammals. The writer has himself partici¬
pated in such field collecting and is familiar with the problem involved.
The collector of mammals is likely to obtain a considerable number of spec¬
imens and to pile them up on his work table before preparing the skins.
As the animal becomes cold the parasites tend to come out to the ends of
the hairs and from that position may cross over to other animals that may¬
be in contact, thus leading to their ascription to the wrong host. Then
the dried skins may be packed together in shipment, leading to the possi¬
bility that the dead lice may be shaken from one host to another and this
may be followed by other possibilities of accident.

Many of the records of lice, especially from the rodents, have been
based upon specimens secured by- examination of museum skins. It is there¬
fore probable that at least some of these records are erroneous. Not until
repeated collections, or the finding of the parasites on freshly collected
animals, have been made can this possibility be eliminated. On the other
hand, merely to dismiss any peculiar record immediately as the result of
such straggling or contamination is probably also a mistake and is likely
to produce a too great assurance that normal transfers never occur.

A small proportion of the records of lice from the larger mammals are
based upon specimens taken in zoological gardens. Some of these are espe-

287

cially questionable, but they must be accepted until more positive informa¬
tion is available.

Considerable thought has been given to the question as to whether or not
the host list should include subspecific mammalian names. The decision has
been to ignore subspecific names and to present all mammal names under the
species names to which the subspecies are to be referred. The reasons for
this action are as follows. In the first place, the writer knows of no in¬
stance where the subspecies make any difference in the distribution of the
lice, except perhaps in the Hyracoidea. The whole subject of the lice of
the Hyracoidea must be subjected to later review on the basis of much more
extensive material of the lice than is available to anyone at present. In
the second place, a listing of the mammalian subspecies would greatly ex¬
tend the host list, to no special purpose. Mammalogists have been much
enamored of the subspecies concept and have employed it to such a degree
that in the rodents, for example, most "full species" have at least five
supposed subspecies, if not more. Peromyscus maniculatus is credited with
forty- five subspecies. Since the Anoplura from Peromyscus seem not even to
discriminate among species, it becomes, from the point of view here held,
merely pretentious nonsense solemnly to list the parasites according to the
subspecies of the host.

In the following list the arrangement of the higher categories of the
mammals is in accord with that presented by Simpson, but the families, sub¬
families, tribes, and genera are arranged alphabetically under the next
higher category.

From the writer's point of view nothing is to be gained in such a list
by attaching the names of the authors of mammalian names. They are there¬
fore omitted.

Class MAMMALIA
Subclass IHERIA
Infraclass EUTHERIA
Order INSECTIVORA
Superfamily MACROSCELI DI DAE
Family Macroscelididae

Elephantulus pulcher and rufescens
Neolinognathus praelautus Ferris
Elephantulus rupestris

Neolinognathus elephantuli Bedford
Nasilio brachyrhynchus

Neolinognathus elephantuli Bedford
Petrodromus tetradactylus
Neolinognathus elephantuli Bedford

Super family SORI COIDEA
Family Soricidae
Subfamily Soricinae

Sorex araneus

Polyplax reclinata (Nitzsch)

Subfamily Crocidurinae

Crocidura aranea (see Sorex)
Crocidura coerulea

Polyplax asiatica Ferris
Crocidura horsfieldi
Ancistroplax crocidurae Waterston
Crocidura leucodon

Polyplax reclinata (Nitzsch)
Pachyura luzonensis

Polyplax reclinata (Nitzsch)
Scutisorex congicus
Polyplax reclinata (Nitzsch)

Suncus coeruleus

Polyplax reclinata (Nitzsch)

Family Talpidae
Subfamily Scalopinae

Parascalops brewer i
Haematopinoides squamosus (Osborn)
Scalopus aquaticus
Haematopinoides squamosus (Osborn)

Order DERMOPTERA
Family Qynocepalidae

Qynocephalus sp. (-variegatus, ac¬
cording to Hopkins)
Hamophthirius galeopitheci Mjoberg
Galeo pith ecus sp .=Qynocephalus

288

Order PRIMATES
Suborder PKDSIMII
1 Hi niorder LEMUR I FORMES
Super family LEMUKOLDEA
Family ludridae

Prop i the cus diadema

Phthirpediculus propitheci Ewing
Propithecus edwardsi. diadema

Super family TUP AIO IDEA
Family Tupaiidae
Subfamily Tupaiinae

Anathana ellioti

Docophthirus acinetus Waterstou

Infraorder L0R1S1F0RMES
Family Lorisidae
Subfamily Galaginae

Cheirogaleus sp.

Lemurphtbirus verruculosus Ward
Galago demidovii

Lemurphtbirus galagus Bedford
Galago moholi

Lemurphthirus galagus Bedford
Galagus-misspelling of Galago
Mouse lemur (-Cheirogaleus)

Suborder ANTHROPOIDEA
Superfamily CEBOIDEA
Family Cebidae
Subfamily Pithecinae

Cacajo rubicundus

Pediculus subhumanus Ewing
Pi thee is monacha

Pediculus pseudohumanus Ewing

Subfamily Alaouttinae

Alouatta palliata

Pediculus mjobergi Ferris

Subfamily Cebinae

Cebus capuchinus

Pediculus atelophilus Ewing
Pediculus chapini Ewing

Subfamily Atelinae
Ateles ater

Pediculus chapini Ewing
Ateles geoffroyi

Pediculus atelophilus Ewing
Pediculus chapini Ewing

Ateles dariensis

Pediculus atelophilus Ewing
Ateles hybridus

Pediculus atelophilus Ewing
Ateles pan

Pediculus atelophilus Ewing
Ateles paniscus

Pediculus lobatus Fahrenholz
Ateles vellerosus (recorded as rel-
lerosus) . This name apparently
has been applied to Ateles
beelzebul and Ateles pan.
Pediculus lobatus Fahrenholz
Lagothrix sp.

Pediculus pseudohumanus Ewing

Subfamily Callithricidae

Leontocebus nigricollis

Pediculus lobatus Fahrenholz

Superfamily CERC0PITHEC01DEA

In speaking of this group Simpson
(1945, page 185 and following) refers
to the "confusion bequeathed to us by
swarms of students, of all degrees of
competence and shades of judgment,"
and further remarks "the macaques have
been placed in at least twenty-five
different genera or subgenera, yet it
is the present consensus that all be¬
long to one genus with perhaps three
subgenera, requiring a total of three
names, only one of generic (and hence
also subgeneric) rank." It is hope¬
less for an entomologist to try to
untangle all of this. Consequently,
records are cited as they were given,
with such accompanying notes as are
possible. There is always the possi¬
bility that in trying to do even this
many new errors have been added. Al¬
so probably many of the host names
are misidentifications.

Cercopithecus aethiops (synonyms

griseoviridis and pygerythraeus)
Pedicinus obtusus
Cercopithecus albogularis=mitis
Cercopithecus cynomolgus-Macaca
cynomolgus
Cercopithecus diana
Pedicinus obtusus

Cercopithecus griseoviridis-aethiops
Cercopithecus kolbi-mitis
Cercopithecus martini-nictitans

289

Cercopithecus mona

Pedicinus obtusus (Rudow)

Pedicinus eurygaster (Burmeister)
Cercopithecus mitis
Pedicinus patas (Fahrenholz)
Cercopithecus patas- Erythrocebus
patas

Cercopithecus pruinosus, apparently
-Pygathrix cristata
Cere o p i th ec us py ge ry thr aeus-ae th i ops
Comop ithecus hamadryas

Pedicinus hamadryas MjOberg
Qynocephalus-Papio
Cynomolgus-Macaca
Erythrocebus patas

Pedicinus patas (Fahrenholz)
Erythrocebus whitei
Pedicinus patas (Fahrenholz)

Guenon. Perhaps an Erythrocebus
Hamadryas, as a generic name this ap¬
pears to =Comopithecus
Innuus nemestrinus=Macaca nemestrina
Innuus sinicus=Macaca sinica
Lasiopyga*=Cercopithecus, under which
genus all names are listed.
Macaca. Frequently spelled Macacus.

Includes Silenus and Rhesus.

Macaca adusta

Pedicinus eurygaster (Burmeister)
Pedicinus obtusus (Rudow)

Macaca albibarbata

Pedicinus eurygaster (Burmeister)
Macaca andamensis

Pedicinus obtusus (Rudow)

Macaca arctoides-speciosa
Macaca cynomolga. Probably -Macaca
fascicularis, although according
to Elliot the name was long mis¬
applied to Macaca ira.

Macaca ery thraea-rhesa
Macaca fascicularis, possibly', for
records under the name Macacus
cynomolgus.

Pedicinus eurygaster (Burmeister)
Macaca innuus, most probably Macaca
sylvana.

Macaca ira, possibly for records
under Macacus cynomolgus.

Pedicinus eurygaster (Burmeister)
Macaca mindanensi-philippinensis
Macaca mindora

Pedicinus eurygaster (Burmeister)
Pedicinus obtusus (Rudow)

Macaca nemestrina, recorded as Innuus
nemestrinus

Pedicinus obtusus (Rudow)

Macaca philippinensis

Pedicinus obtusus (Rudow)

Macaca rhesa

Pedicinus eurygaster (Burmeister)
Pedicinus obtusus (Rudow)

Macaca silena. Apparently -Macaca
albibarbata.

Macaca sinica

Pedicinus obtusus (Rudow)

Macaca speciosa

Pedicinus obtusus (Rudow)

Macaca sylvana

Pedicinus albidus (Rudow)

Papio. Includes Qynocephalus.

Papio sp.

Pedicinus obtusus (Rudow)

Papio griseipes

Pedicinus hamadryas Mjoberg
Pithecus-Macaca ; all names attributed
to it will be found under the lat¬
ter, except as listed.

Pithecus patas. Is probably Erythro¬
cebus patas.

Simia sylvanus-Macaca sylvana

Subfamily Colobinae

Colobus cauda tus- Colo bus polykomos
Colobus polykomos

Pedicinus pictus Ferris
Pedicinus obtusus (Rudow)

Nasalis larvalis, error for larvatus.
Nasalis larvatus

Pedicinus obtusus (Rudow)

Presbytis. Includes all records un¬
der Semnop ithecus
Presbytis cristata

Pedicinus ancoratus Ferris
Presbytis entellus

Pedicinus obtusus (Rudow)

Presbytis germaini

Pedicinus ancoratus Ferris
Presbytis pullata. Regarded by

Elliot as a subspecies of cristata
Presbytis rubicunda

Pedicinus ancoratus Ferris
Presbytis sanctorum

Pedicinus obtusus (Rudow)

Presbytis schistaceus
Pedicinus ancoratus Ferris
Pygathrix aurata

Pedicinus obtusus (Rudow)

Pygathrix priamus

Pedicinus ancoratus Ferris
Rhinopithecus concolor. Presumably
is Simias concolor.

Simias concolor

Pedicinus eurygaster (Burmeister)
Semnopithecus entellus. Here placed
as =Presbytis entellus.

290

Semuopithecus inaurus. Apparently Ochotona danurica

-Presby tis aumta. Hoplopleura ochotonae Ferris

Semuopithecus pruinosus. Here placed Ochotona roylei

as -Presby tis cristatus. Hoplopleura ochotonae Ferris

Ochotona thibetana

Super tamily H0M1N0IDEA Hoplopleura ochotonae Ferris

Family Pongidae

Subfamily Hylobatinae Order R0DENT1A

Hylobates syndactylus

Pediculus humanus Linnaeus
Symphalanges syndacty lus=Hy lotates
syndactylus

Subfamily Ponginae

Gorilla berengeri

Phthirus gorillae Ewing
Pan sp.

Pediculus sch&ffi Fahrenholz
Phthirus pubis (Linnaeus)

Family Hominidae

Homo sapiens

Pediculus humanus Linnaeus
Pediculus subhumanus Ewing
Phthirus pubis (Linnaeus)

Cohort GL1RES
Order LAGOMORPHA
Family Leporidae

Lepus arcticus

Haemodipsus lyriocephalus (Bur-
meister)

Haemodipsus setoni Ewing
Lepus cal i torn icus

Haemodipsus setoni Ewing
Lepus europaeus

Haemodipsus lyriocephalus (Bur-
meister)

Lepus glacialis-arcticus
Lepus saxatilis

Haemodipsus africanus Bedford
Lepus townsendi

Haemodipsus setoni Ewing
Lepus zuluensis=saxatilis
Oryctolagus cuniculus

Haemodipsus ventricosus (Denny)
Rabbit, domestic

Haemodipsus ventricosus (Denny)
Sylvilagus sp.

Haemodipsus setoni Ewing

Family Ochotonidae

Ochotona cansus- thibetana

The arrangement here followed is
strictly according to El ierman, except
that his ending oldae for the super-
family is altered to ouiea in accord
with more generally accepted pi'actice
and families are arranged alphabeti¬
cally under super lami 1 i es, subfam¬
ilies alphabetically under families,
and genera alphabetically under suit-
families.

Superfamily BATHYERG01DEA
Family Bathyergidae

Bathyergus maritimus

Eulinognathus lawrensis (Bedford)
Geo rhyc hus hotten Lotus- Cry p tomy s
hottentotus
Cryptomys hottentotus

Eulinognathus hiili (Bedford)

Super family HYSTK1C0IDEA
Family Chinchillidae

Lagidium inca-peruanum
Lagidium peruanum

Lagidiophthirus parvus (Kellogg and
Ferris)

Family Echimyidae
Subfamily Abrocominae

Abrocoma cinearea

Neohaematopinus longus Wemeck

Subfamily Echimyinae

Cercomys cunicularius

Ctenophthirus cercomydis Ferris
Cercomys sosteri-cunicularius
Proechimys cayennensis

Pterophthirus audax (Ferris)
Proechimys oris=cayennensis
Proechimys semispinosus-cayennensis

Subfamily Octodontinae

Ctenomys b nasi liens is

Eulinognathus americanus Ewing

291

Ctenomys sericeus

Eulinognathus americanus Ewing
Octodontomys gliroides
Hoplopieura disgrega Ferris
Octodontomys simonsi-gliroides

Subfamily Petromyinae

Petromus typicus

Scipio tripedata Ferris
Petromys-Petromus

Subfamily Thryonomyinae

Aulacodus-Thryonomys
Tbryonomys aulacodus-swinderianus
Thryonomys gregor=gregorianus
Thryonomys gregorianus
Scipio longiceps Ewing
Proend erleinellus calva (Waterston)
Thryononys swinderianus
Scipio aulacodi (Neumann)

Scipio breviceps Ferris

Superfamily CAVIODEA
Family Caviidae

Cavia apearea

Pterophthirus imitans Werneck
Caviella australis
Pterophthirus alatus (Ferris)

Galea leucoblephara=musteloides
Galea musteloides

Galeophthirus caviae (Werneck)
Kerodon australis apparently-
Caviella australis

Superfamily SCIUROIDEA
Family Sciuridae

Ellerman recognizes no subfamilies
in this family^, but employs the terms
"group" and "section. " These are
here arranged alphabetically.

Pteromys group

Glaucomys sabrinus

Hoplopieura trispinosa Kellogg and
Ferris

Microphthirus uncinatus (Ferris)
Neohaematopinus sciuropteri
(Osborn)

Glaucomys volans

Hoplopieura trispinosa Kello^ and
Ferris

Neohaematopinus sciuropteri
(Osborn)

Petaurista batuana-petaurista
Petaurista inornata
Neohaematopinus petauristae Ferris
Petaurista petaurista

Neohaematopinus batuanae Ferris
Pteromys volans

Enderleinellus replicatus
Redikorzey

Sc iur op terns -Pteromys in part and
Glaucomys in part.

Sciurus group

Lariscus section

Lariscus diversus=insignis
Lariscus insignis

Enderleinellus larisci Ferris
Neohaematopinus sciurinus (Mjdberg)
Lariscus obscurus- insignis
Menetes berdmorei

Enderleinellus menetensis Ferris
Neohaematopinus sciurinus (Mjdberg)

Marmota section

Ammo sp ermo ph i lus sp .

Neohaematopinus patiki Rubin
Callospermophilus-Citellus
Cite llus adocetus
Neohaematopinus traubi Rubin
Citellus barrowensis=parryi
Citellus beecheyi

Enderleinellus osborni (Kellogg and
Ferris)

Neohaematopinus laeviusculus
(Grube)

Citellus beldingi

Enderleinellus suturalis (Osborn)
Neohaematopinus laeviusculus
( Grube)

Citellus buckleyi-variegata
Citellus buxtoni-eversmanni
Citellus castanurus- lateral is
Citellus chrysodeirus-lateralis
Citellus columbianus
Neohaematopinus laeviusculi
(Grube)

Citellus dauricus

Enderleinellus suturalis
(Osborn)

Citellus douglasii-beecheyi
Citellus elegans-richardsoni
Citellus eversmanni
Enderleinellus suturalis (Osborn)
Neohaematopinus laeviusculus
(Grube)

292

Cite 11 us trank 1 ini i

Enderleinellus sutumlis (Osboni)
Neohaematop inus laeviusculus
(Grube)

Citellus liarrisi

Neohaematopinus citeilinus Ferris
Citellus grummurus*variegatus
Citellus lateralis

Enderleinellus suturalis (Osborn)
Neohaematop inus laeviusculus
(Grube)

Citellus leptodactylus, belongs to
Spermophilopsis
Citellus madrensis

Enderleinellus occideutalis Rellogg
and Ferris
Citellus mexicanus

Enderleinellus suturalis (Osborn)
Citellus mol lis= towns end i
Citellus mongolicus-dauricus

Enderleinellus suturalis (Osborn)
Citellus nelsoni

Enderleinellus suturalis (Osborn)
Citellus oregonus^beldingi
Citellus osgoodi

Enderleinellus suturalis (Osborn)
Neohaematop inus laeviusculus
(Grube)

Citellus parry i

Neohaematopinus laeviusculus
(Grube)

Citellus plesius=parryi
Citellus richardsoni

Enderleinellus suturalis (Osborn)
Neohaematopinus laeviusculus
(Grube)

Citellus rufescens

Neohaematopinus laeviusculus
(Grube)

Citellus spilosoma

Enderleinellus suturalis (Osborn)
Neohaematopinus citeilinus Ferris
Neohaematopinus laeviusculus
(Grube)

Neohaematopinus marmotae Ferris
Citellus tereticaudus

Enderleinellus osborni Rellogg
and Ferris

Neohaematopinus citeilinus Ferris
Citellus towns end i

Enderleinellus suturalis (Osborn)
Citellus tridecemlineatus

Enderleinellus suturalis (Osborn)
Neohaematopinus laeviusculus
(Grube)

Citellus variegatus

Enderleinellus osborni Rellogg and
Ferris

Neohaeina top inus m&thesoni Kubiu
Qynomys guunisoni

Enderleinellus suturalis (Osborn)
Qyuoiqys leucurus

Enderleinellus sutumlis (Osboni)
Neohaematopinus laeviusculus
(Grube)

Qynomys lucovicianus

Neohaematopinus marmotae Perris
Marmota aurea

7 Neohaematop in us marmotae Ferris
Marmota flaviventer=flaviventris
Maniiota llaviventris

Neohaematopinus marmotae Ferris
Manuota monax

Enderleinellus marmotae Ferris

Paraxerus section

Indicated by Ellerman merely as
section D, without name, but for our
purposes it may be listed as the
Paraxerus section.

Ueliosciurus daucinus=gambianus
Heliosciurus gambianu^

Enderleinellus heliosciuri Ferris
Neohaematopinus keniae Ferris
Heliosciurus keniae=gambianus
Heliosciurus multicolor=gambianus
Heliosciurus palliatus-Paraxerus
palliatus

Heliosciurus rufobrachiatus=gambianus
Heliosciurus ruwenzorii

Enderleinellus heliosciuri (Ferris)
Neohaematopinus keniae Ferris
Ueliosciurus undulatus=gambianus
Parasciurus=an error for Paraxerus
Parasciurus animosus^Paraxerus
ochraceus

Paraxerus jacksoni=capitis
Paraxerus ochraceus

Werneckia minuta (Werneck)
Neohaematopinus heliosciuri
Cummings

Neohaematopinus suahelicus Ferris
Paraxerus palliatus

Neohaematopinus heliosciuri
Cummings

Neohaematopinus suahelicus Ferris
Werneckia paraxeri (Werneck)
Protoxerus stangeri

Enderleinellus heliosciuri Ferris

Sciurus section

Callosciurus benticanus=caniceps
Callosciurus borneoensis=prevosti

293

Callosciurus caniceps
Elide rleinellus malaysianus Ferris
Hoplopieura erismata Ferris
Neohaematopinus sciurinus (Mjdberg)
Callosciurus davisoni=caniceps
Callosciurus domelanus, misspelling
ior domelicus—caniceps
Callosciurus ferrugineus
Hoplopieura erismata Ferris
Neohaematopinus sciurinus (MjoLerg)
Callosciurus fin lay son i
Neohaematopinus sciurinus (Mjoberg)
Callosciurus juvencus
Neohaematopinus sciurinus (Mjoberg)
Callosciurus lancavensis—caniceps
Callosciurus lucas=caniceps
Callosciurus maclellandi
hoplopieura erismata Ferris
Callosciurus prevosti
Enderleinellus malaysianus Ferris
Neohaematopinus sciurinus (Mjoberg)
Callosciurus procerus
Neohaematopinus sciurinus (Mjoberg)
Callosciurus vestitutus
Hoplopieura distorta Ferris
Funambulus species (recorded as

palmarum)

Hoplopieura maniculata (Neumann)
Neohaematopinus echinatus (Neumann)
Funambulus palmarum
Enderleinellus platyspicatus Ferris
Neohaematopinus ceylonicus Ferris
Funambulus tristriatus (presumably=
palmarum)

Hoplopieura maniculata (Neumann)
Microsciurus mimilus

Enderleinellus microsciuri Wemeck
Sciurus aberti

Enderleinellus longiceps Kellogg and
Ferris

Neohaematopinus sciurinus (Mjoberg)
Sciurus aestuans

Neohaematopinus sciurinus (Mjdberg)
Sciurus alleni

Enderleinellus longiceps Kellogg and
Ferris

Neohaematopinus sciurinus (Mjdberg)
Sciurus anomalus

Enderleinellus nitzschi (Burmeister)
Neohaematopinus syriacus Ferris
Sciurus apache

Neohaematopinus sciurinus (Mjdberg)
Sciurus arizonensis
Hoplopieura sciuricola Ferris
Sciurus aureogaster

Enderleinellus extremus Ferris
Neohaematolinus sciurinus (Mjdberg)
Sciurus bentincanus-Callosciurus

Sciurus borneoensis=Chllosciurus
Sciurus carolinensis
Enderleinellus longiceps Kellogg and
Ferris

Hoplopieura sciuricola Ferris
Neohaematopinus sciurinus (Mjoberg)
Sciurus colliae

Enderleinellus mexicanus Weraeck
Neohaematopinus sciurinus (Mjdberg)
Sciurus deppei

Enderleinellus extremus Ferris
Sciurus domele ns is= Callosciurus
Sciurus douglasi=Taroiasciurus
Sciurus f remonti=Tamiasciurus
Sciurus lerrugineus=Callosciurus
Sciurus gerrardi

Enderleinellus venezuelae Werneck
Hoplopieura sciuricola Ferris
Neohaematopinus sciurinus (Mjdberg)
Sciurus goldmani=variegatoides
Sciurus griseoflavus
Enderleinellus extremus Ferris
Sciurus griseogena
Enderleinellus venezuelae Werneck
Hoplopieura sciuricola Ferris
Sciurus griseus

Enderleinellus kelloggi Ferris
Hoplopieura sciuricola Ferris
Neohaematopinus griseicolus Ferris
Sciurus ignitus
Hoplopieura sciuricola Ferris
Sciurus kaibabensis

Enderleinellus longiceps Ferris
Sciurus lancavensis= Callosciurus
Sciurus lucas=Callosc iurus
Sciurus melania-variegatoides
Sciurus meridensis=griseogena
Sciurus nayaritensis
Enderleinellus longiceps Ferris
Sciurus negligens
Enderleinellus extremus Ferris
Sciurus nelsoni

Enderleinellus mexicanus Wemeck
Sciurus nesaeus

Enderleinellus insularis Werneck
Hoplopieura sciuricola Ferris
Sciurus niger

Enderleinellus longiceps Kellogg
and Ferris

Neohaematopinus sciurinus (Mjdberg)
Sciurus oculatus

Enderleinellus longiceps Kellogg
and Ferris
Sciurus poliopus
Enderleinellus extremus Ferris
Sciurus syriacus«-anomalus
Sciurus truei

Enderleinellus mexicanus Werneck

294

Sciurus variabi 1 is-gerrardi
Sciurus vulgaris

Enderleinellus nitzschi
(Burmeister)

Neohaematopinus sciuri Jan eke
Sciurus vuLp inus-niger
Uro sciurus- Sciurus

Tam Las section

Kutamias is retained by Ellerman
soLeiy for certain Asiatic species,
whi Le the North American forms common¬
ly referred to it are placed by him
under the genus Tam i as.

Sciurotamias davidianus

Enderleinellus sciurotamiasis
Ferris

Hoplopleura emarginata Ferris
Tamias alleni

Hoplopleura arboricola Kellogg and
Ferris

Neohaematopinus paci ficus Kellogg
and Ferris
Tamias alpinus

Neohaematopinus paci ficus Kellogg
and Ferris
Tamias amoenus

Hoplopleura erratica (Osborn)

Tamias hindsi-townsendi
Tamias merriami

Hoplopleura arboricola Kellogg and
Ferris

Tamias quadrivittatus

Hoplopleura arboricola Kellogg and
Ferris

Neohaematopinus paci ficus Kellogg
and Ferris

Tamias speciosus-quadrivittatus
Tamias striatus

Enderleinellus tamiasis Fahrenholz
Hoplopleura erratica (Osborn)

Tamias townsendi

Hoplopleura arboricola Kellogg and
Ferris

Neohaematopinus paci ficus Kellogg
and Ferris

Tamiasciurus douglasi

Enderleinellus nitzschi Fahrenholz
Hoplopleura sciuricola Ferris
Neohaematopinus sciurinus (Mjoberg)
Tamiasciurus fremonti

Enderleinellus nitzschi Fahrenholz
Tamiasciurus hudsonicus

Enderleinellus nitzschi Fahrenholz
Hoplopleura sciuricola Ferris

Neohaematopinus sciurinus (Mjoberg)

Xerus section

Atlautoxerus getulus

Neohaematopinus pectinifer
(Neumann)

Euxerus microdon-Xerus erythropus
Geo sc i uru s-Xe ru s
Spemophilopsis leptodact} lus

Neohaematopinus cite lli (Cummings)
Xerus capensi s-inauris
Xerus erythropus

Enderleinellus euxeri Ferris
Xerus inauris

Neohaematopinus faurei (Bedford)

Superfamily GE0MY01DEA
Family Heteronyidae

Dipodomys califoruicus-heermanni
Dipodomys deserti

Fahrenholzia piunata Kellogg and
Ferris

Dipodomys heermanni

Fahrenholzia pinnata Kellogg and
Ferris

Dipodomys merriami

Fahrenholzia pinnata Kellogg and
Ferris

Dipodomys ordi

Fahrenholzia pinnata Kellogg and
Ferris

Dipodomys ornatus

Fahrenholzia pinnata Kellogg and
Ferris

Dipodomys phillipsi

Fahrenholzia pinnata Kellogg and
Ferris

Heteronys goldmanni

Fahrenholzia microcephala Ferris
Heteronys pictus-Liomvs
Liomys irroratus

Fahrenholzia microcephala Ferris
Liomys pictus

Fahrenholzia microcephala Ferris
Liomys texensis- irroratus
Microcipodops polionotus

Fahrenholzia pinnata Kellogg and
Ferris

Pero dip us- Dipodomys
Perognathus californicus

Fahrenholzia tribulosa Ferris
Perognathus formosus

Fahrenholzia reducta Ferris
Perognathus hispidus

Fahrenholzia zacatecae Ferris

29^

Perognathus parvus

Fahrenholzia pinnata Kellogg and
Ferris

Perognathus sp.

Fahrenholzia pinnata Ferris

Superfamily PEDETOIDEA
Family Pedetidae

Pedetes cafer

Eulinognathus denticulatus Cummings
Pedetes larvalis=surdaster
Pedetes surdaster

Eulinognathus denticulatus Cummings

Super family DIPODOIDEA
Family Dipodoidea
Subfamily Dipodinae

Allactaga mongolica-sibirica
Allactaga sibirica
Eulinognathus aculeatus (Neumann)

Dip od ipus= Di pus

Dipus sp. Probably a species of
Jaculus

Eulinognathus aculeatus (Neumann)
Dipus sagitta

Eulinognathus biuncatus Ferris

Superfamily MUROIDEA
Family Lophiomyidae

Lophiomys ibeanus=imhausi
Lophiomys imhausi

Eulinognathus lophiomydis Ferris
Lophiomys thomasi* imhausi

Family Muridae
Subfamily Cricetinae

Ellerman remarks concerning this
subfamily that the South American mem¬
bers are "an appalling chaos." For¬
tunately, from a nomenclatorial point
of view, very few Anoplura are known
from the South American species.

Akodon arenicola
Hoplopleura affinis (Burmeister)
Akodon arviculoides
Hoplopleura affinis (Burmeister)
Akodon aerosus

Hoplopleura affinis (Burmeister)
Akodon aurosus=misspeiling of aerosus
Akodon cursorvarviculoides
Akodon mollis

Hoplopleura affinis (Burmeister)

Akodon pulcherrimus
Hoplopleura affinis (Burmeister)
Cricetulus andersoni=longicaudatus
Cricetulus incanus=triton
Cricetulus longicaudatus
Polyplax dentaticornis Ewing
Cricetulus migratorius
Neohaematopinus citelli (Cummings)
(Probably an error)

Eligmodontia collisae, misspelling of

specific najne=Hesperomys callosus
Euneomys pictus=Phyllotis pictus
Graomys griseoflavus
Hoplopleura affinis (Burmeister)
Hesperomys. In part=Peromyscus
Hesperomys callosus
Hoplopleura hesperomydis (Osborn)
Hesperomys venustus

Hoplopleura nesoryzomydis Ferris
Hodomys alleni

Neohaematopinus neotomae Ferris
Holochilus balnearum
Hoplopleura nesoryzomydis Ferris
Holochilus sciureus
Hoplopleura nesoryzomydis Ferris
Holochilus squamipes=Nectomys
squamipes

Holochilus vulpinus
Hoplopleura nesoryzomydis Ferris
Nectomys palmipes=Nectomys squamipes
Nec tone's squamipes

Hoplopleura quadridentata (Neumann)
Neotoma albigula
Neohaematopinus neotomae Ferris
Neotoma cinerea

Neohaematopinus inornatus Kellogg
and Ferris
Neotoma fuscipes
Neohaematopinus neotomae Ferris
Neotoma micropus
Neohaematopinus neotomae Ferris
Neotoma streatori- fuscipes
Neotomodon allstoni

Polyplax auricularis Fello^r and
Ferris

Nesoryzomys-Oryzonrys
Nesoryzomys defessus=Oryzomys
indefessus

Onychonys leucogaster
Hoplopleura hesperomydis (Osborn)
Polyplax auricularis Kellogg and
Ferris

Onychomys torridus
Hoplopleura hesperomydis (Osborn)
Polyplax auricularis Kellogg and
Ferris

Oryzomys angouya

Hoplopleura nesoryzomydis Ferris

296

Oryzomys chap ureas is

Hop lop leant hesperomydis (Osbora)
Oryzomys eliurus

Hoplopleara affinis (Burmeister)
Oryzomys flavescens

Hoplopleura travassosi Werueck
Oryzomys fulvescens

Hoplopleura hesperomydis (Osbora)
Oryzomys indefessus

Hop lop Leara aesoryzomydis Ferris
Oryzomys narboroughi

HoplopLeura aesoryzomydis Ferris
Oryzomys rat ti ceps

Hoplopleura quadrideotata (Neumann)
Oryzomys rostratus

HoplopLeura quadridentata (Neumann)
Oryzomys xantheolus

Hoplopleura aesoryzomydis Ferris
Peromyscus boylei

Hoplopleura hesperonydis (Osbora)
Peromyscus californicus

Polyplax auricularis Kellogg and
Ferris

Peromyscus leucopus

HoplopLeura hesperomydis (Osborn)
Peromyscus maniculatus

Hoplopleura hesperomydis (Osborn)
Polyplax auricularis Kellogg and
Ferris

Peromyscus sitkensis

Polyplax auricularis Kellogg and
Ferris

Phyllotis arenarius

Hoplopleura affinis (Burmeister)

Phy llotis boliviensis

Hoplopleura affinis (Burmeister)
Phyllotis domorum-Graorays griseo-
flavus

Phyllotis micropus

Hoplopleura affinis (Burmeister)
Hoplopleura reducta Ferris
Phyllotis pictus

Hoplopleura af finis (Burmeister)
Phyllotis ricardulus

Hoplopleura af finis (Burmeister)
Reithrodon cuniculoides

Hoplopleura affinis (Burmeister)
Reithrodon hatcheri-cuuiculoides
Reithrodontomys raexicanus

Polyplax auricularis Kellogg and
Ferris

Rhipidomys leucodactylus
Hoplopleura angulata Ferris
Rhipidomys venezuelae

Hoplopleura angulata Ferris
Rhipidomys venustus

Hoplopleura angulata Ferris
Hoplopleura hirsuta Ferris

Sigmodou hispidus

Hoplopleura hirsuta Ferris
Sigmodou ochrognathus

Hoplopleura hirsuta Ferris
Si.Tnodon peruanus

Hoplopleura hirsuta Ferris
Thoinasomys cinereus

Hoplopleura angulata Ferris
Xeuomys nelsoni

Hoplopleura hirsuta Ferris

Subfamily Dendromyinae

Dendromus mesomelas

Hoplopleura intermedia Kellogg and
Ferris

Subfamily Gerbilliuae

Gerbilius indica-Tatera indica
Gerbillus pyramidum

Polyplax gerbilli Ferris
Meriones auceps=meridianus
Meriones ineridianus

Polyplax chinensis Ferris
Hoplopleura merionidis Ferris
Meriones psammophilus=meridi anus
Pachyuromys duprasi

Polyplax werneri ( Glinkiewi csz)
Tatera angolae

Hoplopleura biseriata Ferris
Tatera boehmi

Hoplopleura veprecula Ferris
Tatera brantsi

Hoplopleura biseriata Ferris
Polyplax biseriata Ferris
Tatera indica

Polyplax stephensi (Christophers
and Newstead)

Tatera joanae

Hoplopleura biseriata Ferris
Tatera liodon

Polyplax taterae Bedford
Tatera lobengulae

Hoplopleura biseriata Ferris
Polyplax biseriata Ferris
Tatera nigricauda

Polyplax neumanni Fahrenholz
Tatera vicina

Polyplax taterae Ferris
Taterillus emini

Polyplax taterae Ferris

Subfamily Hydromyinae

Chrotomys whiteheadi

Hoplopleura chrotomydis Ferris
Hydromys chry sogaster

Hoplopleura bidentata (Neumann)

297

Subfamily Microtinae

Arvicola-Microtus in large part
Arvicola anphibius
Polyplax spiniger (Burmeister)
Clethrionomys gapperi
Hoplopleura acanthopus (Burmeister)
Clethrionomys nivarius
Hoplopleura acanthopus (Burmeister)
Clethrionomys rufocanus
Polyplax alaskensis Ewing
Clethrionomys rutilus
Hoplopleura edentulus Fahrenholz
Dicrostonyx torquatus
Hoplopleura acanthopus (Burmeister;
Lagurus intermedius
Polyplax abscisa Fahrenholz
Lemmus alascensis

Hoplopleura acanthopus (Burmeister)
Lemmus obensis
Hoplopleura hispida (Grube)

Microtus agrestis

Hoplopleura acanthopus (Burmeister)
Microtus arvalis

Hoplopleura acanthopus (Burmeister)
Microtus californicus
Polyplax abscisa Fahrenholz
Microtus constrictus-californicus
Microtus mordax

Polyplax abscisa Fahrenholz
Microtus nivalis

Hoplopleura acanthopus (Burmeister)
Microtus pennsylvanicus
Polyplax abscisa Fahrenholz
Microtus sp.

Polyplax alaskensis Ewing
Phaiomys blythei=leucurus
Phaiomys leucurus
Hoplopleura phaiomydis Ferris
Phenacomys longicaudus
Polyplax spinulosa (Burmeister)
Pitymys pinetorum

Hoplopleura acanthopus (Burmeister)
Pitymys savii

Hoplopleura acanthopus (Burmeister)
Pitymys subterraneus
Hoplopleura aequidentis Fahrenholz

Subfamily Murinae

Acomys cahirinus*

Symoca brachyrrhyncha (Cummings)
Polyplax oxyrrhynchus Cummings
Acomys hystrella

Polyplax oxyrrhyncha Cummings
Symoca brachyrrhyncha (Cummings)
Acomys percivali
Polyplax oxyrrhyncha Cummings

Symoca brachyrrhynchus (Cummings)
Acomys sp. (Probably)

Polyplax miacantha Speiser
Aethomys chrysophilus
Polyplax cummingsi Ferris
Apodemus agrarius

Polyplax affinis (Burmeister)
Apodemus insignis
Hoplopleura apomydis Ferris
Apodemus speciosus
Polyplax serrate (Burmeister)
Apodemus sylvaticus
Polyplax spinulosa (Burmeister)

( questionable)

Polyplax serrate (Burmeister)
Hoplopleura affinis (Burmeister)
Apodemus bardus-insignis
Arvicanthis abyssinicus
Polyplax abyssinica Ferris
Arvicanthis niloticus
Polyplax abyssinica Ferris
Arvicanthis pumilio=Rhabdomys pumilio
Arvicanthis univittatus-Hybomys uni-
vittatus

Bandicota bengalensis
Polyplax spinulosa (Burmeister)
Bandicota malabarica
Polyplax asiatica Ferris
Cricetomys cosensi
Proenderleinellus calvus
(Waterston)

Criceton^'s emini

Proenderleinellus calvus
(Waterston)

Cricetomys gambianus
Proenderleinellus calvus
(Waterston)

Dasymys incomtus
Hoplopleura oenomydis Ferris
Hoplopleura somereni Waterston
Polyplax cummingsi Ferris
Dasymys helukus-dncomtus
Epimys-rattus
Eropeplus canus
Polyplax eropepli Ewing
Grammomys i be an us
Polyplax phthisica Ferris
Grammomys surdaster
Hoplopleura oenomydis Ferris
Gunomys= Bandicota
Hybomys univittatus
Hoplopleura laticeps Ferris
Lemni scones barbarus

Hoplopleura enormis Kellow and
Ferris

Lemni sconrys griselda
Hoplopleura enormis Kellog.r and
Ferris

298

Lemni scomys quiche L l us~striatus
Lemiiiscomys striatus
Hoplopleura pelomy dis Kerris
Limnomy s mearnsi
HopiopLeuru oeuomydis Ferris
Lop hu romy s sikapusi

Poiypiax phthisica Kerris
Lophuromys*squi Lus

Poiypiax phthisica Ferris
Lophuromys zena=aquiius
Mastomy s«Rattus
Micromys minutus

HopiopLeura longula Fahrenhoiz
Poiypiax ^raciLis Fahrenhoiz
Mus coucha=Rattus coucha
Mus chrysoph i lus-Aethomy s
chrysophilus

Mus decumauus= Rattus decuinanus
Mus gansus, error for gansuensis=
musculus

Mus ininutus-Micromys minutus
Mus musculus

Hoplopleura acanthopus (Burmeister)
HopiopLeura hesperomy dis (Osborn)
Poiypiax serrate (Burmeister)

Mus spicilegus=musculus
Mus triton

Hoplopleura rukenyae Ferris
Mus wagneri=musculus
Mylomys cuninghamei

Hoplopleura mylomydis Ferris
Mylomys roosevelti=cuninghamei
Nesokia indica

Poiypiax asiatica Ferris
Nesokia hardwickei- indica
Oenomys hypoxanthus

Hoplopleura oenonydis Ferris
Pelomy s fall ax

Hoplopleura pelomydis Ferris
Praomys=Rattus

Praomys namaquensis= Thallomys
Rattus calcis

Hoplopleura oenomvdis Ferris
Poiypiax spinulosa (Burmeister)
Rattus coucha

Hoplopleura intermedia Kellogg and
Ferris

Poiypiax praecisa (Neumann)
Eulinognathus denticulatus Cummings
Rattus culmorum

Poiypiax spinulosa (Burmeister)
Rattus exulans

Hoplopleura oenomvdis Ferris
Rattus hawai iensis

Hoplopleura oenomvdis Ferris
Rattus migricaudus- Thallomys
Rattus norvegicus

Poiypiax spinulosa (Burmeister)

Hoplopleura oenomvdis Kerris
Eulinognathus denticulatus
(Cummings)

Rattus rattus
Rattus sabanus

Hoplopleura mainysiana Ferris
Poiypiax insulsa
Rattus stridens

Poiypiax spinulosa (Burmeister)
Rattus surifer

Hoplopleura pectinata (Cummings)
Rattus tullbergi

Hoplopleura intermedia Kellogg and
Kerris

Poiypiax waterstoni Bedford
Rattus vor.iferans
lloplopleum malaysiana Ferris
Rhabdomys pumilio

Poiypiax arvi can this Bedford
Saccostomus campestris

Poiypiax jonesi Kellogg and Kerris
Thallomys moggi

Hoplopleura affinis (Burmeister)
Poiypiax spinulosa (Burmeister)
Thallomys namaquensis

Poiypiax praomydis Bedford
Thallomy's nigricauda

Hoplopleura affinis (Burmeister)
Thamnomy s=Grammomys
Zelotomys hildegardae

Hoplopleura intermedia Kellogg and
Ferris

Subfamily Otonyinae

Otomys angoniensis

Poiypiax otomydis Cummings
Otomys brantsii=Parotorays brants ii
Otomys irroratus

PolyplaLX otomydis Cummings
Otomys irroratus tropical is=
tropicalis
Otomys tropicalis

Polyp lax otomydis Cunmings
Parotomys brants ii

Poiypiax otomydis Cummings

Family Muscardinidae
Subfamily Graph iur in ae

Graphiurus alticola

Schizophthirus graphiuri Ferris
Graphiurus murinus

Schizophthirus graphiuri Ferris
Graphiurus nanus

Schizophthirus graphiuri Ferris
Graphiurus raptor-murinus

299

Subfamily Muscardininae

Dryomys nitedula

Schizophthirus pleurophaeus
(Burmeister)

Dryomys pallidus=nitedula
Eliomys pallidus=Dryomys nitedula
Eliomys quercinus
Schizophthirus pleurophaeus
(Burmeister)

Muscardinus avellanarius
Schizophthirus pleurophaeus
(Burmeister)

Cohort FERUNGULATA
Superorder FERAE
Order CARNIVORA
Suborder FISSIPEDA
Superfamily CANOIDEA
Family Canidae
Subfamily Caninae

Alopex lagopus

Linognathus setosus (von Olfers)
(blue fox and arctic fox-pre-
sumably Alopex lagopus)

Canis aureus

Linognathus setosus (von Olfers)
Canis brasiliensi^Dusicyon
Linognathus taeniotrichus Werneck
Canis cupus, misprint for lupus
Canis lupus

Linognathus setosus (von Olfers)
Canis mesomelas

Linognathus setosus (von Olfers)
Domestic dog

Linognathus setosus (von Olfers)
Du si cy on fulvipes
Linognathus taeniotrichus Werneck
Dusicyon thous

Linognathus taeniotrichus Werneck
Vulpes fulva

Linognathus setosus (von Olfers)
Vulpes vulpes

Linognathus setosus (von Olfers)

Suborder PINNIPEDIA
Family Odobenidae

Odobenus obesus

Antarctophthirus trichechi
(Bohemann)

Odobenus rosmarus

Antarctophthirus trichechi
( Bohemann)

Family Otariidae
Callorhinus alascanus

Antarctophthirus callorhini
(Osborn)

Proechinophthirius fluctus
(Ferris)

Eumetopias jubata

Antarctophthirus microchir
(Trouessart and Neumann)

Otari a hookeri

Antarctophthirus microchir
(Trouessart and Neumann)

Pho care to s«0 1 ari a
Zalophus californianus
Antarctophthirus microchir
(Trouessart and Neumann)

Family Phocidae
Subfamily Phocinae

Greenland seal

Echinophthirius horridus (von Olfers)
Halichoerus grypus

Echinophthirius horridus (von Olfers)
Phoca foetida*=hispida
Phoca groenlandica

Echinophthirius horridus (von Olfers)
Phoca hispida

Echinophthirius horridus (von Olfers)
Phoca richardii

Echinophthirius horridus (von Olfers)
Phoca sibirica

Echinophthirius horridus (von Olfers)
Phoca variegata

Echinophthirius horridus (von Olfers)
Phoca vitulina

Echinophthirius horridus (von Olfers)

Subfamily Lobodontinae

Hydrurga leptonyx

Antarctophthirus ogmorhini
Enderlein

Leptonychotes weddelli
Antarctophthirus ogmorhini
Enderlein

Lobodon carcinophagus
Antarctophthirus lobodontis
Enderlein

Ogm o rh i n u s-Hy d r u rga
Ommatophoca rossi
Antarctophthirus mawsoni Harrison

Subfamily- Oystophorinae

Qystophorus cristata
Echinophthirius horridus (von Olfers)
Macro rhinus-Mirounga
Mirounga leonina

Lepidophthirus macrorhini Enderlein

300

Superorder PROTON GULATA
Order TOBULLDENTATA
Family Oi'ycteropodidae

Orvcteropus a t er

ily bophUi i rus notophallus (Neumann)
Orvcteropus capeusis

Hybophthirus uotophalLus (Neiuitann)

Order HYRACOIDEA
Family Procaviidae

There seems to be considerable dis¬
agreement concerning the species of
this group. They are listed here ac¬
cording to Lyddeker, 1916, merely as
a standard point of reference.

Procavia bouahioli=ruficeps
Procavia Lrucei=ruficeps
Procavia capensis

Prolinognathus aethiopicus
Fahrenholz

Prolinognathus caviae-capensis
(Pallas)

Procavia coombsi=capensis
Procavia natalensis=capensis
Procavia rufescens=misspelling for
ruficeps

.Procavia ruficeps

Prolinognathus foleyi Fahrenholz
Procavia shoana (properly sciona)-
habessinica
Procavia habessinica

Prolinognathus aethiopicus
Fahrenholz

Prolinognathus arcuatus
Fahrenholz

Heterohyrax syriacus

Prolinognathus leptocephalus
(Ehrenberg)

Superorder MESAXONIA
Order PERISSODACTYLA
Suborder HIPPOMORPHA
Family Equidae

Equus asinus

Haematopinus asini (Linnaeus)
Ratemia squamulata (Neumann)

Equus burchelli

Raenatopinus acuticeps Ferris
Haematopinus asini (Linnaeus)
Ratemia squamulata (Neumann)

Equus caballus

Haematopinus asini (Linnaeus

Superorder PAKAGONIA
Order ARTIODACTYLA
Suborder SU1F0RMES
Intraorder SU1NA
Family Suidae
Subfamily Suinae

Koiropotamus choeropotamus apparent-
ly-Potomoclioerus choeropotamus
Phacochoerus aeJ iani-aethiopicus
Phacochoerus aethiopicus

Haematopinus phacochoeri Euderleiu
Phacochoerus sundevali i-aethiopicus
Potamoclioerus affinis (unable to
trace)

Haematopinus latus Neumann
Po tamocho e rus a f r i can us- larva tus
Potamochoerus choeropotamus
Haematopinus latus Neumann
Sus cristatus

Haematopinus suis (Linnaeus)

Sus scrota

Haematopinus apri Goureau
Sus vittatus

Haematopinus suis (Linnaeus)
Domestic swine

Haematopinus suis (Linnaeus)

Family Tayassuidae
Subfamily Tayassuinae

Dicoty-les-Tayassu

Pecari-Tayassu

Pecari javalii-Tayassu angulatus
Tayassu javalii

Pecaroecus javalii Babcock and
Ewing

Infraorder 0RE0D0NTA
Suborder TYLOPODA
Family Camelidae

Auche n i a-- Lama
Auchenia lama=Lama glama
Camelus dromedarius

Haematopinus tuberculatus
(Burmeister)

Microthoracius cameli (Linnaeus)
Lama glama

Microthoracius mazzai kerneck
Microthoracius minor Werneck
Microthoracius praelongiceps
(Neumann)

Lama huanaca-glama
Lama pacos=glama

301

Suborder RUMINANTIA
Infraorder PEOORA
Super family CERVOIDEA
Family Cervidae
Subfamily Cervinae

Cervus elaphus

Solenopotes bunneisteri Fahrenholz
Cervus nippon

unnamed species of Solenopotes
Cervus unicolor
Haematopinus longus Neumann

Subfamily Muntiacinae

Muntiacus muntjak

Solenopotes muntiacus Thompson

Subfamily Odocoileinae

Capreolus capreolus
Solenopotes capreoli Freund
Coass us— Mazama
Mazama rondoni

Solenopotes binipilosus
(Fahrenholz)

Mazama simplicicornis
Solenopotes binipilosus
(Fahrenholz)

Odocoileus chiriquiensis=rothschildi
Odocoileus columbianus

Solenopotes ferrisi (Fahrenholz)
Odocoileus couesi

Solenopotes binipilosus
(Fahrenholz)

Odocoileus hemionis

Solenopotes ferrisi (Fahrenholz)
Rangifer tarandus
Solenopotes tarandi (Mjoberg)

Super family GIRAFPOIDEA
Family Giraffidae

Giraffa Camelopardalis
Linognathus brevicornis (Giebel)

Superfamily BOVOIDEA
Family Bovidae
Subiamily Antilopinae
Tribe Antilopini

Aepyceros melampus

Linognathus aepyceros Bedford
Antidorcas marsupialis
Linognathus bedfordi Ferris
Linognathus tibialis (Piaget)

Anti lope cervicapra
Linognathus cervicaprae (Lucas)
Linognathus pithodes Cummin rs

Antilope euchore-Antidorcas marsu¬
pialis

Antilope maori (as recorded by

Piaget. The name does not ap¬
pear in available lists).
Linognathus tibialis (Piaget)
Antilope subgutturosa
Linognathus tibialis (Piaget)
"Antilope"

Linognathus tibialis (Piaget)
"Antilope, North African"

Linognathus petasata Ferris
Gaze 11a (possibly either a misidenti-
fication or owing to straggling)
Linognathus gazella Mjoberg
Gaze 11a thorns on i
Linognathus lewisi Bedford
Oreotragus saltator-oreotragus
Oreotragus oreotragus

Linognathus africanus Kellogg and
Paine

Raphicerus campestris
Linognathus tibilais (Piaget)

Tribe Bovini

Bison americanus=bison
Bison bison

Haematopinus tuberculatus
(Burmeister)

Bos grunniens
haematopinus tuberculatus
(Burmeister)

Bos indicus

Haematopinus eurysternus (Nitzsch)
Bos taurus

Haematopinus eurysternus (Nitzsch)
Linognathus vituli (Linnaeus)
Solenopotes capillatus Enderlein
Bubalus bubalis
Haematopinus tuberculatus
(Burmeister)

Syncerus cafer

Haematopinus bufali (de Geer)

Tribe Strepsicerotini

According to Simpson the generic
name Strepsiceros antedates Tra^ela-
phus and Limnotra$us should be regar¬
ded as a subgenus of the former.
There seems to be some question here,
however, and the latter names are re¬
tained for this list.

Also, there seems to be confusion
concerning Rose laphus. The name has
been used for Taurotraius, but seems
to apply to an entirely different
animal.

302

Boselaphus oreas»Tauro tragus oryx
Limno tragus gratus

Linognathus limuotragi Cumrn i ngs
Tiiuro tragus oryx

Haematopiuus tauro tragi Cummings
Linognathus taurotragus Bedford
Tragelaphus scriptus

Linognathus Iractus Ferris
Linognathus limno tragi Cummings
Tragelaphus syivaticus-scriptus

Subfamily Caprinae
Tribe Capriui

Capra hi reus

Liuognathus africanus KelLogg and
Paine

Linognathus stenopsis (Burmeister)
Capra ibex

Linognathus stenopsis (Burmeister)
Capra manifricus (no such name in
any list)

Linognathus oviformis (Kudow)
Caprella rupicapra=Rupicapra
Domestic goat=Capra hi reus
Ovis aries

Linognathus africanus Kellogg and
Paine

Linognathus ovillus (Neumann)
Linoguathus pedalis (Osborn)

Ovis longipes

Linognathus ovillus (Neumann)
Rupicapra rupicapra

Linognathus stenopsis (Burmeister)

Subfamily Cephalophinae
Tribe Cephalophini

Cephalophus grimmi=§ylviacapra grimmi
Cephalophus inaxwelli

Linognathus breviceps (Piaget)
Cephalophus natalensis

Linognathus breviceps (Piaget)
Cephalophus nigri Irons

Linognathus angulata (Piaget)
Philautomba. At times used lor some
species generally referred to
Cephalophus.

Sylviacapra grimmi

Linognathus breviceps (Piaget)

Subfamily Hippotraginae
Tribe Alcelaphini

Connochaetes gnu

Linognathus gnu Bed lord
Damaliscus alii Irons

Linognathus damaliscus Bedford
Damaliscus dorcas*. Damaliscus pygargus
Damaliscus korrigum

Linoguathus damaliscus Bed lord
Damaliscus pygargus

Linognathus damaliscus Bed lord
Gorgon taurinus

Linognathus gnu Bedford
Linoguathus hologastrus toerneck
Linognathus spicatus Ferris

Tribe Hippotragiui

tlippotragus niger

Linognathus hippotragi Ferris

Tribe Reduncini

Cervicapra-Redunca
Pelea capreolus

Linognathus peleus Bedford
Redunca arundinum

Linognathus fahrenholzi Paine
Redunca fulvorufula

Linognathus fahrenholzi Paine
Redunca redunca
Linognathus fahrenholzi Paine

303

CHAPTER VIII

The Distribution of the Anoplura

The problems associated with the spatial distribution of plants and ani-
JTfipH 7 a,t,tract1ed a ™st amount of attention and have come to represent
™nhvid tk the^?e11Je? whlch 1S gathered under the general title of biogeo r-
k-k' + k 11)13 tle,ld lmpin®es upon other fields of inquiry in relation to
w ich there may be a high degree of reciprocal agreement and support, or
conversely, an equal degree of entertaining disagreement. Thus, the geo¬
logical concept of continental drift may on the one hand be supported by
conclusions drawn from biogeography or on the other hand he exposed to most
vigorous criticism from the same source. The strictly practical field of
economic entomology may find a utilitarian aid since it may employ the Geo¬
graphical distribution of insect pests for which the discovery of their na-
tural enemies is desirable. The purely philosophical aspects of biolo.ry
and geography may be concerned with the problems of the spread of naturS
populations and the light that present-day distribution of plants and ani-
malsnmy throw upon the distribution of ancient land masses.

The distribution of parasitic forms is a specialized aspect of bio'?eo *-
raphy and is one of special philosophical interest. It is a matter of* com¬
mon biological knowledge that in the evolution of their parasitic habit
many or^msms— whether piant or animal— are very closely adjusted, both
p >sicaliy and physiologically, to the environment of a single host or host

7? Aaoplura there is a general adjustment°to the environ-
ment afforded by the hair of the hosts. This involves the loss of the
ngs, the reduction or loss of the eyes, the modification of the le rs for
lasping the hairs, and the alteration of the ovipositor for the lavin r of
eggs attached to the hairs of the host. Obviously, this physical ‘adjust¬
ment has been carried still farther into an adjustment to a particular
iTr.’ Anopluran which can grasp the delicate hairs of a mouse would

be completely helpless among the huge bristles of a pig. But beyond such
obvious adjustments as these, there seems also to bean adjustment to ^e
ot the host. There are indications that a sucking louse trans-
} j ™ lts n?rmal {iosb to one of a very different kind may find the
h«vp hpL 6 DZ physiologically unsuited. One may surmise that we
ha e here something analagous to the allergies which at least make life
miserable tor many people. These factors seem to apply rather generally
among many kinds of parasites and they appear very strongly to influence
the distribution of the parasites. uce

Nor is this all. The adjustment of the parasite may be so delicate that
it cannot pass from one host to another of the same species except under a

t", ?°ndiiio;s I*™' in the Aa°p^a thePmodificaUons of [he

for clasping the hairs of the host have proceeded so far that the
louse is unable to move about effectively when detached from its host

Shpne?hpDhlyt SUCh aniJ)ial can pass readi1}' from one host to another only
when the hosts are in contact and a bridge of hair is available.

The outcome ol all these factors is that one might expect to find a par¬
ticular louse species confined to a particular species of host. 4t times
this is true, but usually the distribution of the louse species is broader

IpK1pT[ SpcciesT;,a>- and very frequently does, occur on various closed
related hosts. It seems to be a quite tenable assumption that the louse
bpecies has been inherited from the common ancestor of the various host
KST ^ that hG 1106 haVe evoived somewhat more slowly than have the

304

It this hypothesis be pursued lurther iu its implications, it may very
well be utilized to account for the occurrence of related species of lice
upon related hosts, such as the occurrence ol species of the louse ,'enus
Knderlatnel lua upon rodents of the family Sciuridae throughout the world
and ot other instances that can be cited.

Now, the question arises as to how far this hypothesis may be pursued.
Can it be that the phylogeny of the lice has paralleled the phylogeny of
the hosts back to the time when the ancestors of the lice first became es¬
tablished on the ancestors ot the mammals? It is most tempting to surest
that the c lassi t ication ot the lice, which are a small and compact group,
mit^ht be utilized to throw light upon the phytogeny of the much larger and
mote diverse group ot ttie mammals. These are most intriguing speculations
tor the expression ot hypotheses. But betore we convert speculations into
hypotheses and hypotheses into expressions of opinion we need to inquire
into all the available tacts and to take into consideration all the various
pitfalls that may be encountered. Before the student of lice presumes to
offer advice to the mammalogist concerning moot points in mammal classifi¬
cation he should first be sure that the classification of the lice is
placed upon a sound basis and that he genuinely knows whereol he speaks.

In considering such matters, it should be emphasized at the outset that
at the present time we know probably only about hall' the species of Ano-
plura that may be expected to occur in the world and that until we know

substantially the whole fauna we are scarcely justified in devising a clas¬
sification of the Anoplura that may be regarded as approaching finality.
At the moment, as is emphasized elsewhere, we do not have a satisfactory
classification of the Anoplura as a whole. Most of the genera are relati¬
vely clear, but the way in which these genera may be assembled into genu¬
inely significant higher categories is not yet so. Tribes, subfamilies,
families, superfamilies, these elements in any classificatory system that
will be biologically sound, remain still to be elucidated.

A lew things, however, appear to be reasonably clear.

First, it appears that the Anoplura are confined to the mammals.

Next, it appears that they are confined to the Eutheria, although it is
still possible that they may be found upon the Monotremata which have not
yet been adequately examined. No evidence exists that they occur on the
Metatheria (the Marsupials) although biting lice are known from this group.

Then it appears that they do not occur on certain large groups of the
Eutheria. They seem definitely not to be included among the weird assort¬
ment of parasites that occur on the bats. Among the Ferungulata, which in¬
cludes the living Orders Carnivora and Pinnipedia, the Carnivora seem to
have no Anoplura — with the exception of two species — although they are well
supplied with Mallophaga. The Pinnipedia, on the other hand, seem to pos¬
sess only Anoplura.

Further it appears that there are groups of Anoplura which are definite¬
ly restricted to groups of related hosts, so much so that if any Anopluran
specimen of one of these groups came to hand with no host data it would be
possible to predict with a high degree of probability the family of mammals
from which it came. But not always I There are enough discrepancies to in¬
troduce the possibility of error into such a prediction. Some of these
discrepancies are very strange and at the moment admit of no logical solu¬
tion on the basis of present evidence.

Thus, we have the Anopluran genus Linoinathus, with about 30 included
and apparently evidently related species, which is restricted absolutely to
hosts of the Order Artiodactyla, with two strange exceptions. One species
occurs on the domestic dog and the arctic fox and another species occurs on
a wild dog of South America. These are all the Anoplura that are known
from the terrestrial Carnivora.

There is also the genus Polyplax, with about 30 included species, which

occurs on rodents of the family Muridae, except that two species — clearly
belonging to this genus— occur on shrews of the family Soricidae.

But apart from such glaring exceptions, the species of the various gen¬
era of Anoplura are as tar as known rather generally restricted to host
groups among which some degree of relationship is commonly recognized.

Thus the genus Enderleinellus with about 28 species is, as at present
known, restricted to rodents ot the tamily Sciuridae. The genus Neohaema-
topinus with 25 known species is restricted to the same family, except for
two species which occur on members of the family Muridae. The" ^enus fficro-
thoracius, with four known species, is restricted to the family0 Camelidae,
occurring in both South America and the Old World. The peculiar genus Heo-
l inognathus , with two known species, is confined to the elephant shrews of
the family Macroscel ididae . The genus Prol inognathus , with at least five
apparently distinct species, is confined to the Hyracoidea. The renera
Antarctophthirius and Echinophthirius are confined to the Pinnipedia! The
oenus Schtzophthirus, with two included species, is confined to the rodent
family Muscardinidae. The genus Fahrenholzia , with about five known spe¬
cies, is contined to the rodent iamily Heteromy idae . The genus Pedicinus
with seven included species is confined to the monkey group Ce°rcop i the co idea.

These instances are clear enough. It matters not at all if there be
diife rences ot opinion concerning whether these Anopluran groups be called
genera or are broken each into more than one genus. The group — submenus
genus, supergenus, tribe, or whatever it may be called — as a whole is asso¬
ciated with a particular group of mammals.

Up to this point everything works out nicely. The hypothesis that the
phytogeny of the lice and the phylogeny of the Anoplura are correlated is
entirely tenable. But beginning with these instances we must proceed in
two directions. On the one hand, how much farther does the correlation <ro
in associating these groups of lice with larger groups? On the other hand,
how much farther does it go in associating closely related species of Ano¬
plura with closely related hosts?

Here again we have some beautiful instances of correlation and equally
we have some discrepancies. J

Enderleine l l us nitzscht occurs in North America on squirrels that ac¬
cording to the most recent classification by mammalogists are placed in the
genus Sciurotamias. But what is apparently the same species occurs also on
the European Sciurus vulgaris, while other species of Enderletnellus occur
on other North American species of Sciurus. We have here a partial agree¬
ment and a larger area of disagreement. It would certainly be presumptious
for a student of the Anoplura to advise the mammalogists that Tamiasciurus
should be made to include a species which they utilize as the type of the
genus Sciurus and that other species should be removed from Scturusl

However, within one section of the genus Enderleinellus we have a very
interesting condition. There is a certain group of squirrels that has been
recognized by mammalogists as a category, although the status of this cate-
gory seems not generally to be agreed upon. By some the group has been
called the subfamily Marmotiuae, but Ellerman places it simply as a "Sec-
tion of the Sciuridae. The basic genera are Harmota, Citellus, and Cyno-
nys. I he indications at present concerning the members of the ^enus Ender¬
letnellus occurring upon them are as follows.

The section of the genus Enderleinellus involved has been elevated to a
genus, Cyclophthirus, by Ewing and whatever opinion be held concerning this
genus it exists as a group. Much more collecting needs to be done before
any finaf conclusions are possible, but the evidence at present indicates
that the members of this group of lice are restricted to what we may call
the Marmotinae. The genus Harmota seems to have a distinct species. The
:^enus Cynomys has upon it a species that seems possibly to be distinct
But otherwise the problem of the possible species of Enderleinellus is much

ntused, and recent studies concerning the relationships of the squirrels
H**avfS seem to be equally so. We caunot here go into the matter in

nt.taiL, hut the situation in the classification of the squirrels seems very
much to parallel that found among the lice. When a stud.) of the lice has
been carried to completion it may' show a very interestin '"parallel with the
conclusions o t mainnialojists.

Other illustrations are even more confusing. Thus, the species of the
genus Hop lop leura range over a wide variety of rodents of the various sub-
amiiies oi what Ellerman calls the family Muridae and the Sciuridae. A
louse ot very similar type occurs ou the Lagomorph genus Oc hotona. Within
the genus Hoplopleura various minor groups can be recognized, which seem to
be associated with minor groups of rodents, there being a fairly well de¬
fined group on the Sciuridae. But one species, or at least a series of
very' closely similar forms, ranges from Europe and Asia into South America
on various genera of both Murinae and Cricetinae.

A rather striking instance of an anomalous distribution is that ol the
species Hop lopleura oenomydls , which was described from a rodent of the
Murine genus Oenomys in Africa and recorded from other genera of this group
in Africa and Asia, but which has apparently tiansferred to domestic rats
and is now common in southern United States. The identification is on
available evidence correct.

At the other end of the scale are such genera as Ancistroplax and Haema-
topi no ides which on the totality of their structure seem merely to be some¬
what modified relatives ot Hoplopleura, but occur on Insectivora.

A very peculiar genus is Lemurphthlrus, with one species, which occurs
on a lemuroid. In the characters of head and thorax it seems evidently re¬
lated to Heohaenatoplnus , which is a genus that occurs on squirrels, but in
the characters of its abdomen it is very much a thing by itself although it
is here placed in company with Heohaematopinus in the Polyplacinae . Other
species from lemuroids show no general agreement among themselves and are
placed in this subfamily, which mostly occurs on rodents.

A very illuminating instance is furnished by the one known species of
Phthirpedlculus. In describing this genus Ewing remarked that it stands
"intermediate between Pediculus Linnaeus and Phthirus Leach." This seems
reasonable enough, since Phthirpedlculus is from a lemur and the lemurs are
supposed to stand somewhere near the bottom of the phylogeny of the Pri¬
mates. But the genus Phthirpedlculus, considered on morphological grounds,
has nothing to do with either of these other two genera. In fact, none of
the few lice known from lemuroids seems to have anything especially to do
with the lice of the Primates.

We may here once more call attention to the genus Pedicinus. In all
previous classifications this has been placed in the Pediculidae as a rela¬
tive of Pediculus. This has seemed eminently reasonable, since the hosts
of Pedicinus and Pediculus are Primates. But it is here questioned that
these two genera of lice have anything to do with each other and Pedicinus
is here removed to the Hoplopleuridae, where it is placed as constituting
a subfamily by itself. The morphological evidence does not indicate any
close relationship with Pediculus .

Webb has been much impressed by considerations of host relationship and
the phylogeny of the Anoplura and has used host relationships to influence
his opinions concerning louse classification to a degree that seems to the
writer not to be supported by the evidence of morpholo^' other than his
somewhat questionable evidence derived from the spiracles.

Hopkins (1949) has considered this matter at considerable length in con¬
nection with his list of host-associations of the lice of mammals and point¬
ed out factors concerned with the presence — or, of equal interest, the ab¬
sence — of certain groups of lice on various host groups. He suggests that
where Anoplura and Mallophaga occur on the same host there is perhaps a

307

competition between the members of the two groups of lice which may result
unfavorably for the sucking forms and lead to their disap pearance . It may
be pointed out that the rodent family Heteromyidae supports only sucking
lice oi the peculiar genus Fahrenholzia , while the Geomyidae support only
biting lice of the family Trichodectidae . let while in most lists of mam¬
mals these two groups of rodents are placed as separate families there are
mammalogists who have maintained that they are nothing more than subfami¬
lies of the same family. Again, the rodents of the family Erethizontidae,
the porcupines, seem to possess only biting lice, although in general suck¬
ing lice are much developed on the rodents. However, there are groups of
mammals, such as many of the Ungulates, on which both biting and sucking
lice are well represented. The idea that the absence of the sucking lice
may be due to competition with biting lice must be considered, but there
are other causes that can be envisioned and the present writer is not con¬
vinced that this can have been a tremendously important factor in the pres¬
ent host distribution of the Anoplura.

With such matters as these in mind we may approach the problem of the
correlation of a classification of the higher groups of the Anoplura with
their distribution.

From the writer's point of view the matter of classification should be
developed first and then the problem of distribution should be considered
after the morphological evidence has been developed.

From the point of view here adopted the distribution of the lice in gen¬
eral is on the whole rather consistent with the idea of a correlation of^the
phytogeny of the lice with the phytogeny of their hosts. But above that
point it becomes increasingly inconsistent, and finally fades away into the
realm of pure speculation. If it were pushed to its extreme limits one
would assume that the original ancestor of the sucking lice boarded the
original ancestor of the mammals and the descendants of the one have divid¬
ed and followed the various successors of the original ancestor of the mam¬
mals with the interposition of various accidents — ever since. But it
seems probable that the lice could not have joined the mammals until the
latter had already reached some measure of differentiation into groups.
Under this speculation it may be assumed that some of the ancestral mammals
never achieved the company of any sucking lice. Also, it may be speculated
that the originals of the lice were not specifically adapted to a limited
host group and therefore may have passed rather freely from host to host,
even long after the hosts were quite widely separated phylogenetically ,
thus placing the members of a single phylogenetic line of the parasites
upon many phylogenetic lines of hosts. Even after the lice had differenti¬
ated to a considerable degree this possibility of transfer may have re¬
mained. Then there may have been a certain number of what may be consi¬
dered to be accidental transfers having nothing to do with phytogeny.

Out of all this the correlation of the phytogeny of the parasites with
the phytogeny of the hosts may have begun a process of obscuration early in
the period of differentiation of both groups. This being possible, it
therefore becomes improbable that the phytogeny of the two groups would
follow parallel paths back to the points of origin of both.

The problem then centers merely upon the question of how far these par¬
allel paths extend before they become entangled in the criss-crossing of
roads that may have occurred before a system of parallel paths had devel¬
oped .

From the writer's point of view it is upon this question that future in¬
terest in the general problem should be concentrated.

The conclusions here adopted are as follows:

1. The distribution of the smaller groups — such as genera — is in Gen¬
eral reasonably consistent with the idea that they have followed the phyto¬
geny of the hosts in their development. J

308

2. It. must be recoguized that even here there have been occasional
anomalies ot distribution which must be accepted as they are.

.{• lo a more limited degree the phytogeny of the groups above the gen¬
et a have followed a similar pattern. Liut here the number of apparent dis¬
crepancies increases.

4. When we come to the larger’ groups of genera, which we may call fami¬
lies, this parallelism of phytogenies is but faintly shown.

5. Speculation in regard to the problem should remain firmly within the
bounds of ascertained fact.

6. Discrepancies which cannot be accounted for on a lias is of known fact
should be accepted as they exist and uot forced into some framework of
ideas as to how they ought to be accounted for b}' the system of classifica¬
tion.

7. Future work should be directed in large part toward .the completion
of our knowledge of the species of Anoplura that still remain to be found
and the fitting of these into a system of classification that is based on
the Anoplura themselves.

S. Then and then only will we be in a position to theorize concerning
the relation ol the phytogeny of the lice to the phylogeu} of their hosts.

309

INDEXES

Index to Anoplura

Acanthopinus, in synonymy, 185
antennatus , 195
sclurinus, 195
Achimella, in synonymy, 72
callorhint , 72

Ahaematopinus, in synonymy, 185
inornatus, 192
tnsulsus, 208
ANCISTROPLAX, 120
crocidurae, 120
ANOPLURA, 58, 61
ANTARCTOPHTHIRUS, 71, 72
Antarctophthiriinae, 63
c al i for ni anus, 73
callorhini, 72
lobodontis, 73
mawsoni, 72, 73
microchir, 72, 73
monachus, 72
ogmorhini, 72, 73
trichechi, 72, 75
Arctophtirius, 72
Bathyergicola, in synonym}', 169
hilli, 172
lawrensis, 173
lophiomydis, 173

Bed i'ordia, in synonymy , lf»l
tripedata, 156
Cervophthirius, 251
crassicornts, 256
tarandi, 256

CTENOPHTHIRUS, 163
cercomydis, 164
Cte no pleura, in synonymy, 125
Ctenura, in synonymy, 125
crypt tea, 135
pectinata, 142

Qyclophthirus, in synonymy, 102
suturalis, 113
D000PHTHI RUS, 163, 164

acinetus, 168

ECHLNOPilTHI RII DAE , 63, 68, 71, 72, 77
ECHLNOPHTHIRIUSIINAE, 63, 71
ECHINOPHTHIRUS, 75
baikalensis, 76
fluctu#, 81
groenlandicus, 76, 78
horridus, 75
microchir, 73
sericans, 76
setosus, 73

Ellipoptera, 59
ENDERLEINELL1NAE, 64, 68, 101
ENDERLEI.NELLUS, 101, 102, 103
arizonensis, 105, 106, 108
brasiliensis, 105, 108
dreuiomydis, 104, 108
euxeri, 104, 108
extremus, 106, 108
heliosciuri, 103, 108
houdurensis, 105, 106, 108
insularis, 105, 106, 109

kelLoggi, 105, 106, 109
larisci, 103, 109
longiceps, 104, 105, 106, 109
longiceps group, 104
malaysianus, 104, 105, HO
marmotae, 103, HO
menetensis, 104, 110
mexicanus, 105, 106, 110
microsciuri, 106, 110

minutus, 115, 118
nannosciuri, 104, 110
nitzschi, 104, 110
occidentalis, 113
osborni, 103, 113
paraxert, 115
platyspicatus, 103, 113
replicatus, 113
sciurotamiasis, 104, 113
sphaerocephalus, 110
suturalis, 103, 113
tamiasis, 104, 113
uncinatus, 115
urosciuri, 105, 114
venezuelae, 106, 114
zonatus, 103, H4
Eremophthirius, in s}-non}Tny, 199
biseriatus, 207
ierbtllt, 208
praecisa, 209
stephensi, 214
subtaterae, 214
tarsomydis, 214
taterae, 214
werneri, 215

Euenderleinellus, in synonymy, 102
larisci, 109
Euhaematopininae, 63
Euhaematopinus, in synonymy, 120
abnormis, 120
Euhoplopleura, 125

311

Euhoplopleura —
trispinosa, 144
EULINOGNATHUS, 169

aculeatus, 169, 170
americanus, 170
biuncatus, 170
caviae, 177
denticulatus, 369, 170
hilli, 170, 172
lawrensis, 170, 173
lophiomydis, 170, 173
parvus, 184
surdasteri, 170

FAHREN1I0LZIA, 163, 173
microcephala, 174
p inn at a, 174
reducta, 174
tribulosa, 174
tribulosa, 175
zacatecae, 174, 175
Ferrisella, in synonymy, 125
ochotonae, 125
GALEOPHTHIRUS, 163, 177

caviae, 177
Haeinatomyzus, 59, 60
HAJEMATOPINIDAE, 63, 64, 68, 81, 199
Haematopininae, 63, 64
HAEMATOPINOIDES, 64, 119, 120

squaraosus, 120
Haematopinoidinae, 64
HAEMATOPINUS, 83, 84
acanthopus, 130
aculeatus, 170
acuticeps, 83, 84
adventicius, 91
affinis, 130
albtdus, 158
annulatus, 75
antennatus, 195
aperis, 84
appendiculatus, 240
apri, 84
asini, 84, 85
aulacodt, 153
bicolor, 236
bidentatus, 133
breviceps, 228
brevicornis, 229
breviculus, 87
bufali, 84, 88
bufalt-capensis, 88
bufall-europaet, 95
burchellls, 85
camp.lt, 244
cerutcaprae, 229
chinensis, 91, 94
clavlcornls, 282
coloratus, 285

HAEMATOPINUS—
columbianus, 193
crassicornis, 253
echinatus, 191
elevens, 85
equii, 285
erraticus, 136
eurystemus, 84, 88
forficulus, 238
iermanicus, 94
germanus, 94
hesperomydis, 136
incisus, 90
irritans , 91
laeviusculus, 193
latissimus, 90
latus, 84, 90
leptocephalus, 251
loniulus, 137
longus, 83, 91
lyriocephalus, 179
macrocephalus, 85
mniculatus, 138
microcephalus, 231
minor, 85
montanus, 193
notophallus, 151
obtusus, 159
oviformts, 231
outllus, 231
pectinifer, 194
pedalis, 231
peristictus, 90, 91
phachochoeri, 84, 91
pi lifer us, 236
pleurophaeus, 285
praecisus, 139, 209
praecitus, 138, 209
praelonitceps, 247
praviprocursus, 88
punctatus, 95
quadridentatus, 142
quadripertusus, 88
reclinatus, 210
rupicaprae, 238
saccatus, 235
sardiniensis, 94
sciuropteri, 1%
serratus, 210
setosus, 194
sphaerocephalus, 110
spiculifer, 283
spiniger, 211
spinulosus, 211
squamulatus, 217
stephenst, 214
suis, 84, 91
suturalis, 113

312

UAEMATOPINUS—
taurotrajdj 83, 95
ttbtal Is, 238
trlchecbl, 75
tuberculatus, 83, 95
tuberculatum, 244
ungulatus, 228
ux’ius, 84
ventrlcosus, 179
vituli, 241

Haematopinus, in synonymy, 199, 251
HAEMODIPSUS, 163, 177

at'ricanus, 179
lyri ocephalus, 179
parvus, 183
setoni, 179
ventricosus, 179
Hamophthiri inae, 64
HAMOPTdIRUS, 163, 183

^aleopitheci, 183
RaseLlus, in synonymy, 147
pleurophaeus, 14?
daustelleta, 59
Hemiptera, 59

HopLophthirus, in synonymy, 102
euxeri, 108
HOPLOPLEURA, 119, 125

acanthopus, 128, 130
acanthopus, 135
affinis, 127, 130
affinis, 133. 134
alata, 144
angulata, 129, 133
apomydis, 127, 133
arboricola, 129, 133
argentina, 129, 133
audax, 147
bidentata, 126, 133
biseriata, 128, 134
brasiliensis, 126, 134
chilensis, 125, 134
chrotonydis, 129, 134
cricetuli, 127, 134
cryptica, 126, 135
dis^rega, 125, 135

dlsgrega, 134
distorta, 128, 135
edentula, 135

emar^inata, 125, 135
enormis, 127, 135

erismata, 128, 136
erratica, 128, 136
erratica, 133
fonsecai, 128, 136
hesperomydis, 127, 136
hirsuta, 129, 137
hispida, 137
intermedia, 126, 137

dOPLOPLEUKA—

laticeps, 126, 137
lineata, 138
lon^ula, 128, 138
malaysiana, 129, 138
maniculata, 129, 138
inerionidis, 129, 138
mylomydis, 128, 138
nesoryzomydis, 127, 138
neumanni, 127, 138
ochotouue, 126, 139

oenomydis, 129, 139
oryzomydis, 127, 139
oxynycteri, 128, 139
pacifica, 139
pectinata, 126 , 142
pelomydis, 126, 142
phaiomydis, 128, 142
quadridentata, 127, 142
reducta, 129, 142
reithrodontomydis, 128, 143
rukenyae, 128, 143
sciuricola, 129, 143
soraereni, 127, 143
sukenyae, 143
travassosi, 127, 143
trispinosa, 126, 143
veprecula, 126, 144
HOPLOPLEURIDAE, 68, 98
doplopleurinae, 68, 101, 119
HYBOPHTBIRIINAE, 64, 68, 101, 148
HYBOPUTHIRIUS, 148, 151
notophallus, 151
Inrostrata, 60
LAGI DI0PHTH 1RUS, I63, 183
parvus, 183

LEMURPHTHIRUS, 163, 185

^alaeTis, 185
verruculosus, 185
LEPID0PHTH1RIINAE, 64
LEPIDOPHTdIRUS, 78
macrorhini, 78
LINOGNATHIDAE, 69, 220
Lino^nathinae, 63, 64, 220
Lino^nathoides, in synonymy, 185
citelli, 191
faurei, 191
inornatus, 192
pectinifer, 194
spermophtli, 191
LINOGNATOUS, 222, 223
aepycerus, 225
africanus, 225
angulatus, 225, 223
angulatus, 252
antennatus, 238
appendiculntus, 240
bedfordi, 22 3, 228

313

LINOGNATHUS —

btnipilosus, 253
breviceps, 225, 228
brevicornis, 225, 229
burmeisteri, 253
cauiae-capensis, 248
cervicaprae, 225, 229
coassus, 253
crassicornis, 253
damaliscus, 229
euchore, 240
fahrenholzi, 224, 229
ferrisi, 230
ferrisi, 256
forficulus, 229
fractus, 225, 230
iazella, 228
iiluus, 228
gorgonus, 230
gnu, 230

hippotragi, 224, 230
hologastrus, 224, 230
lewisi, 223, 231
limnotragi, 225, 231
oviform is, 231
ovillus, 224, 231
panamensis, 253
pedalis, 224, 231
peleus, 225, 231
petasmatus, 224, 233
pithodes, 223, 234
praelongiceps, 244
saccatus, 235
setosus, 224, 235
squamulatus, 217
spicatus, 223, 237
stenopsis, 224, 233
stenopsts, 225, 227
taeniotrichus, 224, 238
taurotragus, 224, 238
tibialis, 225, 238
vituli, 223, 241
vitult , 253
ungulatus, 228

Linognathus, in synonymy, 251
Lutegus, in synonym}-, 185
pectinifer, 194
Mallophaga, 59
Mandibulata, 56
MICROPHTHIRUS, 102, 115
uncinatus, 115
MICR0TH0RACIUS, 222, 243
cameli, 244
mazzai , 244
minor, 244, 247
praelongiceps, 244, 247
praelongtceps, 244
N BOH AEMATOPINUS, 164, 185

N B0 HAEMA70PIN US —
batuanae, 190
ceylonicus, 189, 190
cite lli, 188, 191
citellinus, 189, 191
echinatus, 189, 191
echinatus, 190
faurei, 190, 191
griseicolus, 189, 192
heliosciuri, 190, 192
inornatus, 189, 192
kenyae, 190, 192
laeviusculus, 190, 193
longus, 189, 193
macrospinosus, 195
marmotae, 190, 193
mathesoni, 190, 194
neotomae, 189, 194
pacificus, 189, 194
patiki, 194
pectinifer, 189, 194
petauristae, 190, 195
sciuri, 189, 195
sciurinus, 189, 195
sciurinus, 192
sciuropteri, 189, 196
semlfasciatus, 195
suahelicus, 190, 196
syriacus, 189, 196
traubi, 190, 196
NE0LINOGNATH1DAE, 257
NED LINO® ATHINAE, 64
NE0LIN0GNATHUS, 257
elephantuli, 257, 258
praelautus, 257, 258
Neopedicinus, in synonymy, 157
patas, 162

Neumannellus, in synonymy, 151
aulacodi, 151

Paenipediculus, in synonym}-, 260
schaffi, 277
slmiae, 277

Parapediculus, 260, 273
atelophilus, 260
chapini, 273
lobatus, 274
pseudohumanus , 275
Parasita, 59
Parasiti, 61
PECAROEaiS, 83, 98
javalii, 98

PEDIQNINAE, 63, .64, 69, 101, 156, 258
PEDICINUS, 157

albidus, l’57, 158
ancoratus, 157, 158
breviceps, 158, 159
col ob l, 159
eurygaster, 157, 158

314

PEDICINUb —
euryiaster , 159
graclllcepx, 159
hamadryas, 157, 159
longiceps, 158, 159
n icropi losus , 158
mlcrops, 158
obtusus, 157, 159
parallellceps, 159

patas, 157, 162
piageti, 158
pictus, 157, 162
rhesl, 159
vulgaris, 159
PEDICULIDAE, 59, 63, 258
Pediculina, 59
Pediculinae, 63, 64, 156
PEDICULUS, 260
acanthopus, 130
affinis. 130, 274, 276
albidlor, 262
americanus, 267
angustus, 266
annulatus, 75

aquaticus, 282
as ini , 84
assimilis, 266
atelis, 277
atelophilus, 273
bufali, 88
bufali-capensis, 88
bufali-europaei , 95
camel i, 244
capitis, 262
cauiae-capensls, 248
ccrv tcalis , 26 1
c bap ini, 273
chinensis, 2 66
clavicornis, 282
col laris, 248
consobrinus, 262, 275
corporis, 2 66
crassicornis, 253
denticulatus, 211
eurygaster, 158
eurysternus , 88
ferus, 281
flavidus, 236
friedenthali, 266

hispidus, 137
horridus, 75
hunianus, 261
iso pus, 236
laeviusculus, 193
leptocephalus, 251
lobatus, 273. 276
lyriocephalus, 179
macrocephalus , 85

PKUICULUS—
nac'ulatus, 266
marginatus, 266
mjober^i, 261, 274
mi crops, 158
nigrescens, 262
nigritarum, 261, 267
oblongus, 2 66
oxyrrhynchus, 241
papillosus, 88
phocae, 75
pi It ferus, 2 36
phthirlopsis, 88
pleurophaeus, 148
pseudohumanus, 26 1 , 275
pseudohumanus , 267
pubescens, 262
pubis, 281
quadnunanus, 275
reclinatus, 210
schal'fi, 261, 277
schistopygus, 238
serratus, 210
setosus, 75
simtae,

spermophili, 193
sphaerocephalus, 102, 110
spicu lifer, 283
spiniger, 211
spinulosus, 193, 211
stenopsis, 2^
suis, 91

tabescentiim, 262
tenuirostris, 241
tuberculatus, 95
urtus, 91
vestimenti, 262
vituli, 241

Petauristophthirus, in synonymy, 185
petaurtstae, 195
Phthiraptera, 60
Phthiriidae, 64
Phthirius, 144, 280
Phthirpedicinus, in synonymy, 157
micropilosus, 158
microps, 158

PHTHIRPEDICULUS, 163, 199
propitheci, 199
Phthirus, in synonymy, 280
gorillae, 281
shavest, 281

POLYPLACINAE, 69, 101, 162
POLYPLAX, 163, 199, 209
abscisa, 205
abyssinica, 204, 205
acanthopus, 130
affinis, 130
alaskensis, 205

315

POLYPLAX—

arvicanthis, 204, 206
asiatica, 203, 206
auricularis, 203, 206
bidentata, 133
biseriata, 203, 207
borealis, 205
brachyrrhyncha, 220
californiae, 206
calva, 21 6
campylopteri, 211
chinensis, 203, 207
cummingsi, 208
deltoides, 204, 207
dentaticornis, 202, 207
echinata, 191
eminata, 214
eriopepli, 203, 207
gerbilli, 203, 208
gracilis, 208
gracilis, 208
insulsa, 203, 208
j one si, 204, 208
laeuiuscula, 193
longula, 137
maniculata, 138
miacantha, 283
otomydis, 203, 208
oxyrrhynchus, 209
paine i , 206
pectinifer, 194
phthisica, 204, 209
praecisa, 203, 209
praomydis, 211
quadridentata, 142
reclinata, 204, 210
reclinata, 207
serrata, 204, 210
spinigera, 202, 211
spinulosa, 205, 211
spinulosa, 206
stephensi, 203, 214
s ubtaterae, 214
tarsomydis, 204, 214
taterae, 203, 214
waterstoni, 204, 214
werneri, 203, 215
PROECHINOPHTHIRIUS, 81

fluctus, 81

PROENDERLEINELLUS, 163, 215
afrlcanus, 21 6
calva, 216
hllli, 172
lawrensts, 173

PROLINOGNATHUS, 222, 247
aethiopicus, 248

PROLINOGNATHUS —
arcuatus, 248
caviae-capensis, 248
ferrisi, 248, 251
foleyi, 248, 251
leptocephalus, 248, 251
Pseudorhynchota, 59
PTHIRUS, 280
gorillae, 281
inguinalis, 281
pubis, 281

PTEROPHTHIRUS, 119, 144
alata, 144
audax, 144, 147
imitans, 144, 147
wemecki, 144, 147
RATEMIA, 163, 216
squamulata, 217
Rhinaptera, 59
Rhinophthirus, 102
heliosciuri, 196
Rhynchota, 59
Rhynchophthirina, 59, 60
SCHIZOPHTlilRUS, 119, 147, 148
graphiuri, 148
leucophaeus, 285
pleurophaeus, 1-6
SCIPI0, 148, 151

aulacodi, 152, 153
breviceps, 152, 154
longiceps, 154
tripedatus, 152, 156
Siphunculata, 59, 60
S0LEN0P0TES, 222, 251
binipilosus, 252
burmeisteri, 253
capillatus, 252, 253
capreoli, 252, 256
ferrisi, 25*2, 256
muntiacus, 256
panamensis, 253
tarandi, 256

Stobbella, in synonymy, 223
pithodes, 235
SYMOCA, 164, 220
brachyrrhyncha, 220
Symysadus, in synonymy, 215
calva, 216

Trichaulinae, in synonymy, 220
Trichaulus, in synonymy, 223
Waterstonia, in synonym}', 215
calva, 216
zanzibariensis, 216
WERNECKIA, 102, 115
minuta, 115, US
paraxeri, 115, 118

316

Index to Hosts

Alphabetical list of genera and higher groups only. Species are listed
alphabetical Ly under their genera in the host list and can then be found by
ret erring to the index of Auopluran species.

Abrocoma, 291

Abrocominae, Subfamily, 291
Acoinys, 298
Aepyceros, 302
Aethomys, 298
Akodon, 29b

Alcelaphini, Tribe, 303
AlLactaga, 296
Alopex, 300
ALouatta, 289

Alouattinae, Subfamily, 289
Ammospermoph i lus, 292
Anathana, 289

Anthropoidea, Suborder, 289
Antidorcas, 302
Anti lope, 302

Anti lop inae. Subfamily, 302
Anti lop ini, Tribe, 302
Apodemus, 298
Artiodactyla, Order, 301
Arvicanthis, 298
Arvicola, 298
Ateles, 289

Atelinae, Subfamily, 289
Atlantoxerus, 295
Auchenia, 301
Aulacodus, 292

Bandicota, 298

Bathyergidae, Family, 291

Bathy ergoidea, Superfamily, 291

Bathy ergus, 291

Bison, 302

Bos, 302

Boselaphus, 303

Bovidae, Family, 302

Bovini, Tribe, 302

Bovoidea, Superfamily, 302

Cacaj 0 , 289

Callithricidae, Subfamily, 289
Callorhinus, 300
Callosciurus, 293
Callospemiophilus, 292
Camelidae, Family, 301
Camelus, 301
Canidae, Family, 300
Caninae, Sublamily, 300
Canis, 300

Cano idea, Super family, 300
Capra, 303
Caprella, 303

Capreolus, 302
Caprinae, Sublamily, 303
Caprini, Tribe, 303
Carnivora, Order, 300
Cavia, 292
Caviella, 292
Cavi idae, Fam i ly , 29 2
Cavoidea, Superlainily, 292
Cebidae, Family, 289
Cebinae, Subfamily, 289
Ceboidea, Superfamily, 289
Cebus, 289

Cephalophinae, Subfamily, 303
Cephalophini, Tribe, 303
Cephalophus, 303
Cercomys, 291

Cercopithecoidea, Superfamily, 289

Cercopi thee us, 289

Cervicapra, 303

Cervidae, Family, 302

Cervinae, Subfamily, 302

Cervoidea, Superfamily, 302

Cervus, 302

Cheirogaleus, 289

Chinchillidae, Family, 291

Chrotomys, 297

Citellus, 292

Clethrionomys, 298

Coassus, 302

Colobinae, Subfamily, 290
Colobus, 290
Comopithecus, 290
Connochaetes, 303
Cricetinae, Subfamily, 296
Cr ice tomys, 298
Cricetulus, 296
Crocidura, 288

Crocidurinae, Subfamily, 288
Cryp tomys, 291
Ctenomys, 291

Qynocephalidae , Family, 288
Qynocephalus, 290
CJynomolgus, 290
Qynomys, 292

Cystophorinae, Subfamily, 300
Qvstophorus, 300

Damaliscus, 303
Dasymys, 298
Dendromus, 297

Dendromyinae, Subfamily, 297
Dermoptera, Order, 288

317

Dicotyles, 301
Dicrostonyx, 298
Dipod idae. Family, $$6
Dipodinae, Subfajiiily, 296
Dipodipus, 296

Dipodoidea, Superfamily , 296

Dipodomys, 295

Dipus, 296

Dryoroys, 300

Dusicyon, 300

Echimyidae, Family, 291
Echinryinae, Subfamily, 291
Elephantulus, 288
Eligmodontia, 296
Eliomys, 300
Epimys, 298
Equidae, Family, 301
Equus, 301
Eropeplus, 298
Erythrocebus, 290
Eumetopias, 300
Euneomys, 296
Eutheria, Infraclass, 288
Euxerus, 295

Ferae, Superorder, 300
Ferungulata, Cohort, 300
Fissipeda, Suborder, 300
Funambulus, 294

Galaginae, Subfamily, 289
Galago , 289
Galea, 292
Galeopithecus, 288
Gaaella, 302

Geomyoidea, Superfamily, 295
Georhychus, 291
Geosciurus, 295
Gerbillinae, Subfamily, 297
Gerbillus, 297
Giraffa, 302
Giraffidae, Family, 302
Giraffoidea, Superfamily, 302
Glaucomys, 292
Glires, Cohort, 291
Gorgon, 303
Gorilla, 291
Grammomys, 298
Graomys, 296

Graphiurinae, Subfamily, 299
Graph iurus, 299
Guenon, 290
Gunomys, 298

Halichoerus, 300
Ham;idryas, 290
Heliosciurus, 293

Hesperomys, 296
deterohyrax, 301
Heteromyidae, Family, 295
Heteromys, 295
Hippomorpha, Suborder, 301
llippotraginae. Subfamily, 303
Hippotragini, Tribe, 303
Eippotragus, 303
Hodonys, 296
Holochilus, 296
Hominidae, Family, 291
Hominoidea, Superfamily, 291
Homo, 291
Hybomys, 298

Hydroinyinae, Subfamily, 297

Hydromys , 297

Hydrurga, 300

Hylo bates, 291

Hylobatinae, Subfamily, 291

Hyracoidea, Superfamily, 301

Hystricoidea, Superfamily, 291

Indridae, Family, 289
Innuus, 290

Insectivora, Order, 288

Kerodon, 292
Koiropotamus, 301

Lagidium, 291

Lagomorpha, Order, 291

Lagothrix, 289

La gurus, 289

Lama, 301

Lariscus, 292

Lariscus section, 292

Lasiopyga, 290

Lemmus, 298

Lemniscomys, 298

Lemuriformes, Infraorder, 289

Lemuroidea, Superfamily, 289

Leontocebus, 289

Leporidae, Family, 291

Leptonychotes, 300

Lepus, 291

Limnomys, 299

Limnotragus, 303

Liomys, 295

Lobodon, 300

Lobodontinae, Subfamily, 300
Lophiom^ idae, Family, 296
Lopbiomys, 296
Lophuromys, 299
Lorisidae, Family, 289
Lorisi formes, Infraorder, 289

Macaca, 290
Macacus, 290

318

Macrorhinus, 3U0

Macroscel ididae, Family, 2H8

Macrosce 1 i Jo idea. Superfamily, 288

Mannota, 293

Mannota section, 292

Mas tony s, 292

Mazuma, 302

Menetes, 292

Meriones, 297

Mesaxonia, Superorder, 301

Microdipodops, 295

Micronys, 299

Microsciurus, 294

Microtinae, Subfamily, 298

Microtus, 298

Mirounga, 300

Muntiacinae, Subfamily, 302
Muridae, Family, 296
Murinae, Subfamily, 298
Muroidea, Superfamily, 296
Mus, 299

Muscardinidae, Family, 299
Muscardininae, Subfamily, 300
Muscardinus, 300
Mylomys, 299

Nasalis, 290
Nasilio, 288
Nectomys, 296
Nesokia, 299
Nesorvzomys, 296
Neotoma, 296
Neotomodon, 296

Ochotona, 291
Ochotonidae, Family, 291
Octodontinae, Subfamily, 291
Octodontomys, 292
Odobenidae, Family, 300
Odobenus, 300

Odocoileinae, Subfamily, 302
Odocoileus, 302
Oenorays, 299
Ogmorhinus, 300
Ommatophoca, 300
Onychomys, 296
Oreodonta, Infraorder, 301
0ocier°PO(lidae, Family, 301

0ryc!eroPus> 301
Oryclol01^115* 291
Oryzomys> 296

Otaria, 300

Otariidae, Family, 300
Otonyinae, Subfamily, 299
Otomys, 299
Ovis, 303

Pachyura, 288

Pachyuromvs, 297
Pan, 291
Papio, 290

Paragonia, Superorder, 301
Parascalops, 288
Parasciurus, 293
Paraxerus, 293
Paraxerus section, 293
Parotomys, 299
Pecari, 301

Pecora, Infraorder, 302
Pedetes, 296
Pedetidae, Family, 296
Pedetoidea, Superfamily, 296
Pelea, 303
Pelonys, 299

Perissodactyla, Order, 301
Perodipus, 295
Perognathus, 295
Peromyscus, 295
Petaurista, 292
Petrodromus, 288
Petromus, 292

Petromyinae, Subfamily, 292
Petronys, 292
Phachochoerus, 301
Phaionys, 298
Phenacomys, 298
Philantomba, 303
Phoca, 300
Phocarctos, 300
Phocidae, Family, 300
Phocinae, Subfamily, 300
Phyllotis, 297
Pinnipedia, Suborder, 300
Pithecis, 289

Pithecinae, Subfamily, 289
Pithecus, 290
Pitynys, 298
Pongidae, Family, 291
Ponginae, Subfamily, 291
Potamochoerus, 30 1
Praomys, 299
Presbytis, 290
Primates, Order, 289
Procavia, 301
Procaviidae, Family-, 301
Proechinys, 291
Propithecus, 289
Prosimii, Suborder, 289
Protoxerus, 293
Protungulata, Superorder, 301
Pteromys, 292
Pteromys group, 292
Pygathrix, 290

Rangifer, 302
Rattus, 299

319

Redunca, 303
Reduncini, Tribe, 303
Rei thro don, 297
Reithrodontomys, 297
Rhabdomys, 299
Rhinopithecus, 290
Rhipidomys, 297
Rodentia, Order, 290
Ruminantia, Suborder, 302
Rupicapra, 303

Saccostomus, 299
Scalopinae, Subfamily, 2®
Scalopus, 288
Sciuridae, Family, 292
Sciuroidea, Superfamily, 292
Sciuropterus, 292
Sciurotajriias, 29 5
Sciurus, 294
Sciurus ^roup, 292
Sciurus section, 293
Scutisorex, 288
Semnopithecus, 290
Si ^rnodon, 297
Simia, 290
Simias, 290
Sorex, 288

Soricidae, Family, 288
Soricinae, Subfamily, 288
Soricoidea, Superfamily, 288
Spermophilus, 29-5
Strepsiceros, 302
Strepsicerotini, Tribe, 302
Suidae, Family, 301
Sui formes. Suborder, 301
Suina, Infraorder, 301
Suinae, Subfamily, 301
Suncas , 288

Sus, 301

Sylviacapra, 302
Sylvilajus, 291
Symphalanjes, 291
Syncerus, 302

Talpidae, Family, 288
Tamias, 299
Tamias section, 295
Tamiasciurus, 295
Tatera, 297
Taterillus, 297
Taurotra^us, 303
Tayassu, 301

Tayassuidae, Family,. 301
Tayassuinae, Subfamily, 301
Tballomys, 299
Thamnoinys, 299
Theria, Subclass, 288
Thomasomys, 297
Thryonomyinae, Subfamily, 292
Thryonomys, 292
Tubulidentata, Order, 30 1
Tupaiidae, Family, 289
Tupaiinae, Subfamily, 289
Tupaioidea, Superfamily, 289
Tylopoda, Suborder, 301

Urosciurus, 295

Yulpes, 300

Xenomys, 297
Xerus , 29 5
Xerus section, 295

Zalophus, 300
Zelotomys, 299

320

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Full text of "The sucking lice. With the collaboration of Chester J. Stojanovich."

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