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PAEASITES.

PAKASITES;

A TREATISE ON THE

ENTOZOA OF MAN AND ANIMALS,

INCLUDING

SOME ACCOUNT OF THE ECTOZOA.

T. SPENCER COBBOLD, M.D., F.R.S., F.L.S.,

HONORARY VICE-PRESIDENT OF THE BIRMINGHAM NATURAL HISTORY
AND MICROSCOPICAL SOCIETY.

LONDON:

J. & A. CHURCHILL, NEW EUKLlNGlO.N STREET.

1879.

PBEEACE

My introductory treatise on the Entozoa having long been
out of print, it occurred to me that instead of attempting
another edition it would be better to write an entirely new
work, employing only such fragmentary portions of the old
treatise as would harmonise with the far wider design I have
now in view. Whilst, therefore, I have freely utilised a
selection of the illustrations given in the elementary volume,
comparatively few of its pages have been incorporated in the
present work.

Dealing with parasites and parasitism after a manner not
hitherto attempted I have purposely omitted minute anatomical
descriptions, and, with rare exceptions, I have avoided the
introduction of clinical details. While bringing to a focus the
records of, and principal references to, a widely scattered, intri-
cate, and voluminous literature, it has been my chief endeavour
to supply abundance of original matter of a kind that cannot
be found in the columns of any existing treatise. Whether I
have succeeded or not the experienced helminthologist alone
can judge. He, at all events, will perceive that the summary,
though compressed within the space of a moderate-sized octavo,
can only have resulted from sustained effort.

This treatise is not professional, that is to say, it does
not concern itself with therapeutics or the curative treatment
of parasitic affections ; yet it introduces and helps to solve

PREFACE

many questions relating to epidemics, endemics, and epizootics
due to parasites. The medical man who only looks at the
phenomena of parasitism as displayed within the human terri-
tory must of necessity acquire a cramped, narrow, and distorted
conception of the role played by parasites in the production
of disease. Let it be freely granted that to the practising
physician, as such, it matters little how many beasts, birds,
reptiles, or fishes perish annually from parasitic affections ; yet,
when it is demonstrable that a large proportion of the strictly
human entozoa require a change of hosts — or, in other words,
need to pass through the bodies of the lower animals — then it
is evident that some acquaintance on his part with the entozoa
infesting animals becomes a practical necessity. Knowledge of
the kind here offered will often materially aid him in recom-
mending prophylactic measures. Moreover, the study of
comparative pathology, almost ignored in England, conveys
with it other lessons of high value in relation to the healing
art. The great mind of John Hunter comprehended all this
long ago, as any student of the beautiful preparations contained
in the museum of the Royal College of Surgeons may readily
convince himself ; and this is all the more noteworthy, since the
subject concerns the physician rather than the surgeon.

To the naturalist the second half of this book addresses
itself in a very direct manner. When engaged in his dissec-
tions, an appeal to its pages will often enable him to decide at
once as to the species of parasite accidentally encountered, and if
a full diagnosis be demanded it will guide him to better sources
' of information. Many hundreds of correspondents, not having
ready access to the systematic writings of Rudolphi, Diesing,
and Dujardin, have requested me to identify their " finds." I
have rarely or never failed to comply with their requests ; but
it is hoped that the present work may prove of ready service to
subsequent inquirers, and thus place a reasonable limit upon the
number of future applicants. Since the manuscript of this work
was completed I have received Dr von Linstow's Compendium

PREFACE

Vll

der Helminthologie, which, for the purposes held in view by the
author, leaves little to be desired.

Expressly to meet the requirements of the Sanitarian I
have dwelt upon the developmental phenomena exhibited by
those parasites that occasion fatal helminthiases ; and, in this
relation, I have not confined my remarks to the parasites that
are injurious to man in a direct manner, but have extended my
observations to the genesis of those entozoa that prove destruc-
tive to horses, to beasts of burden generally, and to other
creatures which, like cats and dogs, are in various ways sub-
servient to man's wants. It will be seen that in this way
several questions relating to the purity of water and flesh-
food, respectively, have been incidentally brought under notice.

In view of the magnitude of the task which my enthusiasm,
perhaps unwarrantable, has led me to undertake, I know full
well how considerately my foreign friends and correspondents
will deal with the errors of omission and commission that they
will certainly detect in these pages. If there be any educated
persons at home who still affect to despise the revelations of
helminthology, I can assure them that their prejudices are
misplaced. The study of the structure and economy of a
humble parasite brings to the investigator no slight insight
into the workings of nature. If these workings cannot at all
times be pronounced to be " good and beautif ul," they must at
least be characterised as " true." The knowledge of the true
— especially if that knowledge by its practical applications be
calculated to confer substantial benefits upon man and his
inferior fellow- creatures — ought to be held in high esteem; but,
apart from this purely utilitarian view, there remains for the
investigator the delight occasioned by the in-rush of new scientific
ideas. The average mind, being either essentially commercial
or ridiculously sentimental, as the case may be, is totally inca-
pable of comprehending the motive power that animates and
guides the votary of science. The late Professor Faraday, a

viii

PREFACE

man wholly untinged by the ambitions of wealth and power,
once remarked to me that there were no people so difficult to
instruct as those who were ignorant of their own ignorance.
It is just these very persons who, when placed in high positions
of social, political, or professional trust, most powerfully con-
tribute to check a nation's progress. There are too few
genuine workers at science in this country. As one of the
rank and file, I claim only to have honestly contributed my
mite. I should like to see a small army of helminthologists
rise up and lay siege to the fortresses at present securely held
by thousands of death-dealing parasites.

T. S. 0.

74, Portsdown Road, London
May, 1879.

SYNOPSIS OF CONTENTS.

PAGE

General Introduction . • . . 1

BOOK I.
PARASITES OF MAN.

Section I. — Trematoda (Flukes) . . . .14

„ II. — Cestoda (Tapeworms) . . .56

„ III. — Nematoda (Roundworms and Threadworms) . 149
„ IV.-

Part I. — Acanthocephala (Thornheaded Worms) . 256

„ II. — Suctoria (Leeches) .... 257

„ III. — Arachnida (Parasitic forms of) . . 259

„ IV. — Crustacea (alleged Parasitic forms of) . 268

„ V. — Insecta (Parasitic forms of) . . . 269

„ VI. — Protozoa (Parasitic forms of) . . 276

Appendix (Statistics) ..... 284

BOOK II.

PARASITES OF ANIMALS.

Section I. — Parasites op Mammalia.

Part I. — Parasites of Quadrumana . . . 289

„ II. — Parasites of Cheiroptera . . . 293

„ III. — Parasites of Insectivora . . . 295

„ IV. — Parasites of Carnivora . . . 297

„ V. — Parasites of Pinnipedia . . . 313

„ VI. — Parasites of Rodentia . , .315

„ VII. — Parasites of Edentata . . . 320

„ VIII. — Parasites of Ruminantia . . . 322

IX. — Parasites of Solidungula . . . 356

„ X. — Parasites of Pachydcrmata . . . 393

„ XI. — Parasites of Cetacea and Sirenia . . 416

„ XII.— Parasites of Marsupialia and Monotremata . 430

„ II. — Parasites op Aves .... 434

„ III.— Parasites op Reptilia . . . 451

„ IV. — Parasites of Pisces .... 457

„ V. — Parasites op Evertebrata . . . 480

Appendix (Hsematozoa) . . . .• . 485

Index . . . . . . .489

LIST OF BIBLIOGRAPHIES.

PAGK

General and systematic treatises

Minor treatises, general memoirs, and monographs

. 10

Literature of Fasciola hepatica in man .

. 17

Distoma lanceolatum in man

. 20

„ crassum ....

. 28

„ sinense ....

. 2d

,, conjunctum in man

. 33

„ heterophyes ....

. 35

„ ophthalmobium

. 36

10.

Tetrastoma and Hexatliyridium .

. 36

11.

Amphistoma hominis ....

. 38

12.

Bilharzia hcematobia ....

. 55

13.

Taenia mediocanellata and the beef-measle

. 84

14.

„ solium and the pork-measle

. 94

15.

„ tenella and the mutton-measle

. 99

16.

„ lophosoma ....

. 99

17.

„ nana .....

. 100

18.

Tapeworm varieties and monstrosities .

. 105

19.

Bothriocephalus lotus, B. cordatus, and B. cristatus

. 112

20 a.

General literature of hydatids (English)

. 141

Hydatids of the liver

. 142

,, „ and other organs together

. 143

Liver hydatids. American cases

. 144

Hydatids of the lungs and pleura

. 144

/■

„ of the kidney

. 144

„ of the spleen, omentum, and abdominal cavity

. 144

,, within the pelvic cavity

. 145

„ of the heart and blood-vessels

. 145

Jc.

,, of the brain and cranial cavity

. 145

„ of the bones

. 145

„ of the breast, muscles, and soft parts .

. 146

„ of uncertain seat .

. 146

„ of animals

. 147

P-

„ in man. Foreign literature .

. 147

21.

Trichina spiralis. English literature

. 174

„ Foreign literature

. .177

LIST OF BIBLIOGRAPHIES

PAGt

22.

Trichocephalus dispar

23.

Filaria Bancrofti (F. sanguinis hominis) .

90.O

Stipplement (Hsematozoa)

A OQ

. 4oo

24.

Filaria loa .

. 20b

25.

lentis ....

. 206

26.

labialis ....

. 207

27.

trachealis and F. bronchialis

28.

Eustrongylus (Strongylus) gigas

. 210

29.

Dochmius duodenalis ....

. 21b

30.

Dracunculus medinensis

. 21:4

ol.

Oxyuris vwmicularis ....

OQO
_• >-

32.

Leptodera (Anguillula) stercoralis and L. intestinalis

not?

. 235

33.

Ascaris mystax ....

nil

. 241

o4.

himbricoides ....

OKI

35.

Echinorhynchus gigas ....

. 257

36.

Sanguisuga medAcinalis and other leeches

. 259

37.

Pentastoma tcenioides and P. constrictum .

. 265

38.

Demodex, Sarcoptes, and other Arachnidan ectozoa

. 268

on
OV.

Gammarus pulex in man

A A
40.

Bugs, lice, and other insect parasites of man

. 275

A 1

41.

Pswospermia, Gregarina, and other protozoa

. 283

42.

Entozoa of monkeys ....

. 293

A O

43.

and ectozoa of bats

. 295

44.

of insectivorous mammals

. 297

45.

of carnivorous mammals

. 310

4b.

J»

of seals ....

. 315

a n

47.

of rodents ....

. 320

48.

of sloths and ant-eaters

. 322

A Q

of ruminants

. 352

and ectozoa of solipeds

ooo

. 389

51.

„ of elephants

. 400

■>-.

of rhinoceroses

. 402

CO
OO.

of the hippopotamus and tapir .

. 403 .

and ectozoa of swine .

. 414

00.

of whales, dolphins, and dugongs

. 429

00.

of marsupial animals .

. 434

0/.

and ectozoa of birds .

. 448

to
Oo.

of reptiles ....

. 456

oy.

and ectozoa of fishes .

. 477

60.

of insects, crustaceans, and mollusks

. 484

ERRATUM.

Page 296, line 24 from the top, for " in the glow-worm (Olomeris)," read "in a
myriapod {Olomeris) which is phosphorescent like the glow-worm."

PARASITES.

INTRODUCTION.

No person can derive advantage from the study of parasites
unless the subject be approached in a right frame of mind. In
other words, the student of helminthology must, as a primary
discipline, dispossess himself of all preconceived opinions what-
soever, and in an attitude of child-like simplicity seek truth
for its own sake. Unless the mind be absolutely free and
unfettered it cannot rightly interpret the facts of this peculiar
department of biological science. Those students who are
nervously anxious to reconcile the conclusions of modern science
with the ideas of their forefathers are certain to remain just as
ignorant of the true value and significance of nature-teachings
as all their fathers were.

Whether dealing with the external or internal forms, the
study of parasites of man and animals is practically one of
boundless extent ; and there is probably no department of
knowledge, possessing an equal value in relation to the welfare
of man and beast, that is so thoroughly misunderstood by those
who are directly concerned in the appreciation of its revelations.
This has arisen from a total misconception as to cause and
effect. Most people, not excluding even the votaries of the
healing art, following tradition, regard the internal parasites or
entozoa as creatures either directly resulting from certain
diseased conditions of their hosts or as organisms which
would not have existed if their bearers had been perfectly
healthy. Nothing can be more absurd. Such a conclusion is
utterly at variance with all logical deduction from known facts.
It is, however, quite on a par with multitudes of other popular

INTRODUCTION

delusions which, in spite of the advance of science, will pro-
bably never become wholly eradicated from the public mind.
People who hold these notions either cannot or do not desire
to reject a view which has for them a dominating power almost
equal to that of any known religious dogma. In conversation
I have repeatedly noticed this to be the case. These people are
the victims of educated ignorance and they will never allow
that parasites are natural developments, accomplishing ends or
parts of the orderly mystery which reigns everywhere. Some
of them still cling to the creed that the presence of parasites,
of internal ones at least, betokens evidence of Divine disfavor;
and their minds are troubled with all sorts of distressing and
childish conceptions. In the present age one would have
thought that such ridiculous ideas could not be seriously main-
tained ; but instead of being relegated to the limbo of similar
c( old wives' fables " they dominate the opinions of thousands
of our so-called educated people. The genuine searcher after
truth does not need to be told that all preconceptions of this
order hopelessly obscure the mental vision. They operate to
render a just and adequate understanding of the science of
helminthology impossible. The biologist may say what he lists,
but he knows perfectly well that the superstitious mind will
continue to ignore the precious and elevating results of scientific
research, and that it will perseveringly continue to persuade
itself that internal worms, parasites, and entozoa, of whatever
kind, belong to the category of "plagues" liable to be dis-
tributed as special punishments for human wrong-doing.

As remarked in my previous treatise, the best way of studying
the entozoa is to regard them as collectively forming a peculiar
fauna, destined to occupy an equally peculiar territory. That
territory is the wide-spread domain of the interior of the bodies
of man and animals. Each bearer or " host " may be viewed
as a continent, and each part or viscus of his body may be
regarded as a district. Each district has its special attractions
for particular parasitic forms ; yet, at the same time, neither
the district nor the continent are suitable as permanent resting-
* places for the invader. None of the internal parasites " continue
in one stay all have a tendency to roam ; migration is the
soul of their prosperity ; change of residence the essential of
their existence ; whilst a blockade in the interior soon termi-
nates in degeneration and death. I repeat it. The entozoa con-
stitute a specialised fauna. What our native country is to

INTRODUCTION

ourselves, the bodies of animals are to them. To attack, to
invade, to infest, is their legitimate prerogative. Their organi-
sation, habits, and economy are expressly fashioned to this end.
How remarkable and complex is their structure, and how pecu-
liar, diverse, and varied are their ways and wanderings, the
contents of this volume will, I trust, sufficiently explain. The
puerile horror which even some scientific persons affect to
display in regard to the subject is altogether out of place. To
the rightly balanced mind the study of these much abused
" worms " is just as attractive as any other section of zoology.
Helminthology opens up to our view many of the strangest
biological phenomena of which the human mind can take
cognisance ; whilst a profound and extended knowledge of the
subject, in all its bearings, is calculated to secure to the com-
munity a rich practical reward by enabling us to do effectual
battle with not a few of the many ills of life to which our flesh
is heir.

Further on the general advantages to be derived from the
study of parasites I cannot here dilate, and it becomes the less
necessary that I should do so, since I have entered upon the
subject very fully elsewhere. The character of the present work,
moreover, imposes brevity. If the plan which I now propose
to follow should not be deemed altogether satisfactory from the
purely zoological standpoint, it will nevertheless have the
advantage of simplicity and novelty ; and knowing full well the
difficulties that must surround any attempt to give a perfect
classification of the entozoa, considered as a natural group, I feel
sure that my helminthological friends will credit me with
exercising a wise discretion in selecting the simplest available
method of arrangement. My plan, therefore, is to devote
separate sections of this work to the parasites of the different
classes of vertebrated animals, including man, treating of the
various species in regular succession. This arrangement is
merely one of convenience and has no reference whatever to
conceptions of zoological equivalency as variously interpreted
and maintained by authors and investigators. The parasitic
groups will be taken up in the following order, quite irrespective
of their relative importance, and also without any attempt to treat
each group with equal fulness. In the matter of recent literature
only will the present record and summary make any approach
toward completeness, my hope being to render this treatise
indispensable and trustworthy as a ready means of reference.

INTRODUCTION

I. Flukes. Trematoda. — This group embraces several fami-
lies of parenchymatous worms. The various species exhibit
one or more suckers, which the older naturalists regarded as so
many mouths or perforations. Hence the ordinal title. The
term fluke is of Saxon origin, meaning anything flat. Thus,
it has been applied to sole-fish or flounders, to the flattened
halves of the tail of cetaceans, to the blades of anchors, and so
forth. Although the common liver fluke is flat, many species
of the order are round, biconvex, or even filiform organisms.
I recognise six families : — Monostomida, Distomidce, Amphisto-
midce, Tristomidce, Polystomidte, and Oyrodactylida. Most of
the species are entozoal ; but many adhere to the surface of
the body of piscine hosts.

II. Tapeworms. Cestoda. — This comprises not only the
tapeworms, but also the measles and other bladder-worms or
cystic Entozoa of the old authors (Cystica). The Greek word
hestos means a band or girdle ; hence the ordinal term above
given. The bladder- worms, including Hydatids, Cysticerci,
&c, are the larval stages of growth of various tapeworms.
The further reduction of this order into sub-orders or families
requires careful attention. At present we have Tceniada, Acan-
thotaniada, Dibothrida (=Both7-iocephalidce), Diphyllobothridce,
Tetrarhynchida, and Tetraphyllobothrida. All the genera and
species are entozoal. The proposal to separate the snouted
or proboscidiform tapeworms (Rhynchotceniada) from those in
which the rostellum is absent (Arhynchotceniada) does not
recommend itself to my judgment.

III. Roundworms. Nematoda. — This series comprises not
only lumbricoid or roundworms proper, but also threadworms.
The term derives its origin from the Greek word nema, signify-
ing a thread. It likewise includes the strongyles, the term
strongulos meaning round or cylindrical. This is a very extensive
group whose parasitic members are strictly entozoal, whilst the
non-parasitic forms are either entirely free or they infest plants.
Some of the so-called free nematoids live in the slime of
animals. The artificial classification by Schneider, based on the
muscular system, places these parasites in three well-marked
groups, but I think it a disadvantage to separate widely many
really closely allied forms. Thus, in his Polymyarii we have
the genus Eustrongylus, and in his Meromyarii the Strongyli
proper. Most of the genera may be fairly included in the
following families : — Ascaridce, Chciracanthidce, Gucullanida,

INTRODUCTION

Strongylidce, Trichinida?, Oxyurida, Trichoccphalida, Filarida,
Gordiidce, Anguillulida.

IV. Thornheaded-worms. Acanthocephala.. — This group
embraces a small series of parasites, which, in general appear-
ance, resemble the nematode worms. They differ, however,
essentially, being, as the term indicates, furnished with spine-
covered heads. They are, moreover, destitute of digestive
organs. The species are entozoal in habit, abounding particu-
larly in fishes and reptiles. At present, all the known forms
are included in one family (Echinorhynchidce), which also
comprises only a single genus.

V. Annelid parasites. Suctoria. — In this category one
must place all such suctorial annelids as affix themselves
to hosts for a longer or shorter period. Many of the leech-
like parasites (Glepsinida, and especially Mallacobdellida)
remind one of certain flukes (Tristoma, &c.) possessing ectozoal
habits ; whilst the leeches, properly so called, afford instances
of the passage from a semi-parasitic to what has been called the
free parasitic mode of existence. In tropical countries these
creatures very readily attach themselves to man and animals,
often creating severe distress. The genera Glepsine and
Hcemocharis attack mollusks and fishes respectively. The
species are all ectoparasitic and exceedingly numerous. They
cannot be described in this work.

VI. Arachnid parasites. Arachnida (part of). — The great
class of articulated, limb-jointed, or, more strictly, arthro-
podous animals, includes a variety of parasites. The mites,
true ticks, and such like creatures, belong to this group.
Some few of them are entozoal in habit, others are only par-
tially so, whilst the majority are entirely ectozoal. Of the
two great sections of Arachnida, namely, Pulmonaria and Tra-
chearia, the latter alone contains strictly parasitic forms. The
parasitic species belong to the following families : — Pentasto-
midoe, Pycnogonidce, Ixodidce, Acarida, Gamasidce, Hydrach-
nida, Solpugidce. The parasitism of some of the species is
very partial or slight. Thus, certain of the water mites, in their
juvenile state, dwell on aquatic insects only j and the tick-like
Gamasidce occur upon dung-beetles. The other ectozoal species
attack vertebrated animals, and several attach themselves to
man himself. The whale lice (Gyamidce) are here included in
the PycnogonidO', though often placed by zoologists with the
Crustaceans.

INTRODUCTION

VII. Crustacean parasites. Crustacea (part of). — A large
number of species belonging to various well-marked sections of
this great class of Invertebrates are parasitic in their habits, most
of them being comprised in the so-called haustellated group.
They are familiarly known to zoologists as Epizoa. As this
latter term implies, they are strictly ectozoal in character,
most of the species victimising fishes by attaching themselves,
not only to the general surface of the body, but also to the
eyes, and especially to the gills or branchise. The species for
the most part belong to the families Lemceidce, Caligida?, Diche-
lestidce, and Argulidce. In this category must likewise be
placed two other families belonging to the so-called isopodous
section of edriophthalmatous crustaceans. These are the Cymo-
thoidcp, which attach themselves to the tails of fishes, and the
Bopyridae, which occupy the branchial cavity of shrimps. The
nature of this work precludes any detailed notice of the
numerous members of this section.

VIII. Insect parasites. Insecta (part of). — The insects,
properly so called (that is to say, arthropodous, evertebrated
creatures, with six legs), are many of them essentially parasitic
in their habits. The most important of these are " bots "
and other larvae or maggots of various flies (Diptera).
The varieties of lice are also included in this group. Some
few of the insect parasites are strictly entozoal in habit, at
least for a part of their lifetime, being previously attached
externally for a short period only. Most of the forms are
essentially ectozoal. A very large number of insect tormentors,
although deriving nourishment from their victims, attach them-
selves to the animals for so short a time that they cannot be
classed as parasites under the ordinary acceptation of the term.
As examples of the so-called free parasitism, the autumnal
flies (Tabanida) and Stomoxys may be cited. Although embrac-
ing but few strictly parasitic forms we have the following : —
QjJstridce, Hippoboscidce (with Melophagus), and Nycteribiida.
In regard to the maggots of Muscidce and Sarcophagi, some of
them are parasitic on animals and man, whilst others are
parasitic upon insects themselves. The larvae of Gonopidce
attack humble-bees internally. Those parasitic insects, pro-
perly so called, which, like certain of the crustaceans, are some-
times spoken of as epizoa, comprise three well-marked families.
Thus, we have Pediculidce (the source of lousiness), Pliilo-
pteridce, and Liothoidce. Both of the latter embrace numerous

INTRODUCTION

species which for the most part content themselves with
devouring the feathers of birds and the hairs of quadrupeds.
In addition to these it may be added that some of the rat-tailed
larvee or Helophilus maggots (Syrphidce) are parasitic in man
and quadrupeds, as are also the larvse of the churchyard beetle
(Blajptida) . The closely allied Tenebrionidce and other coleo-
pterous families also supply various maggots possessed of para-
sitic habits. Fleas and bugs come under Van Beneden's category
of free parasites. This is equivalent to calling them non-
parasitic parasites, an expression which looks very like a con-
tradiction of terms.

IX. Protozoal Parasites. Protozoa (part of.) — This mis-
cellaneous assemblage of minute creatures embraces a num-
ber of parasites of very low organisation. In the present work
it is neither desirable nor necessary to hazard any statements
respecting their precise zoological position. It is sufficient to
say that the parasitic protozoa are for the most part entozoal
in habit, not a few of them possessing vegetable affinities.
The microscopic Bacterids, Qregarinidce, and Psorospermiae,
comprise a multitude of organisms which are strictly parasitic
in their habits, whilst amongst the Infusoria we find numerous
forms which, though dwelling in the intestinal canal of their
hosts, do not derive nourishment in a direct manner from their
bearers. Of this kind are Paramecium and Balantidium. The
separation of the psorospermise and gregaringe into genera
is attended with difficulty ; nevertheless, I have for convenience
long recognised various types under titles corresponding with
the names of the observers who first discovered them (Hesslingia,
Gubleria, Lindermannia, and so forth). Of necessity, the
protozoal parasites will only be incidentally noticed in this work.
In this category I place the falsely so called " cattle-plague
bodies." The micrococci and bacteria hardly come within the
province of the helminthologist.

Without prejudice to the foregoing restrictions I must at the
same time observe that the varied characters presented by the
above-mentioned groups show how impossible it is to treat the
subject of parasitism adequately, if one is obliged to confine his
remarks to the internal parasites or helminths proper. Many
creatures possessed of entozoal and ectozoal habits are parasites
in every legitimate sense of the term, and yet they do not belong
to the class Helmintha in its common zoological acceptation.
That class taken by itself may still be allowed to stand pretty

INTRODUCTION

much as I represented it in 1864; but in the present work I
cease to speak of the Entozoa as in any sense the zoological
equivalent of the Helmintha. I prefer to employ the term
Entozoa in its popular and wider acceptation. It conveniently
stands thus, moreover, in direct contradiction to the term
Ectozoa.

As this work treats of parasites only, I purposely refrain
from dealing with the Turbellarians, and certain other creatures
usually classed with Vermes. The vague term " worms," so
often employed as the equivalent of Helmintha, is misleading in
many ways. I should like to see it adopted only when speak-
ing of the Annelids proper. It would still have a sufficiently
wide application, seeing that it would include Leeches, Earth-
worms, Naids, Tubed-worms, Sea-lobworms, Sea-mice, Nereids,
and a host of other setigerous species. Notwithstanding the
remote connection subsisting between " intestinal worms"
and worms properly so called, the notion that an intimate
relation subsists between the lumbricoid helminths and earth-
worms will probably never entirely disappear from the popular
or even from the professional mind.

Since one of the principal features of this treatise is to afford
a handy means of reference to the rich and extended literature of
parasitism, I here subjoin a list of general and systematic
treatises. To most of these I shall constantly refer. Full
special references to detached memoirs will appear in the biblio-
graphies scattered throughout the body of the work.

Bibliography (No. 1). — Bremser, ' Ueber lebende Wiirmer
im lebenden Menschen/ Vienna, 1819 ; French edit., by
Grrundler, 1824. — Idem, ' Icones helminthium/ Vienna, 1824. —
Gobbold, T. 8., ' Entozoa, an Introduction to the Study of Hel-
minthology, with reference more particularly to the Parasites of
Man/ London, 1864; Supp., 1869. — Reviews in the 'Lancet,'
Sept. 24th, 1864, p. 353 ; in the ' Med. Times and Gaz./ Oct.
29th, 1864, p. 474; in the 'Athenaeum/ Oct. 15th, 1864,]).
493; in 'Cosmos/ Oct. 27th, 1864, p. 463 ; in the 'Reader/
Nov. 26th, 1864, p. 668 ; in the ' Edinburgh Vet. Review/ Nov.,

1864, p. 662; in 'Intellectual Observer/ vol. vi, 1864, p. 190 ;
in the ' Quarterly Journal of Science/ No. v, January, 1865,
p. 145 ; in the ' Quart. Journ. of Micr. Science/ New Series,
No. 17, Jan., 1865, p. 43; in 'Popular Science Review/ Jan.,

1865, p. 214; in the 'Veterinarian/ Feb., 1865, p. 97 ; in the
' Medical Mirror/ Jan., 1865, p. 23; in the 'Natural History

INTRODUCTION

Beview' for July, 1865; in the ' British and Foreign Medico-
Chirurgical Beview,' April, 1865, in the ( Edinburgh Medi-
cal Journal ' for April, p. 929; in the 'Social Science Eeview'
for Feb. 1, 1866, p. 169; in 'Dublin Quart. Journ. of Medical
Science' for Aug., 1867. — Davaine, G., ' Traite des Ento-
zoaires et des maladies vermineuses de l'homme et des animaux
domestiques,' Paris, 1860, 2nd edit., 1877-79.— Diesing, 0. M.,
' Sy sterna helminthum,' Vienna, 1850. — Bujardin, F., ' Histoire
naturelle des helminthes ou vers intestineaux,' Paris, 1845. —
Goeze, T. A. 8., e Versuch einer Naturgeschichte der Einge-
weidewiirmer thierischer Korper,' Blank enburgh, 1782. —
Kuchenmeister, F., 1 Die in und an dem Korper des lebenden
Menschen vorkommenden Parasiten,' Leipsic, 1855, 2nd. edit.,
1878-79; Eng. edit., by Lankester, 1857.— Be Glerc, D., 'A
Natural and medicinal History of Worms bred in the bodies of
men and other animals' {sic), Browne's edit., London, 1721. —
Beuckart, B,., ' Die menschlichen Parasiten, und die von ihren
herruhrenden Krankheiten,' Leipsic und Heidelberg, 1863-1876.
■ — Bedi, F., ' De animalculis vivis quae in corporibus animalium
vivorum reperiuntur, observationes ;' Coste's edition, AmstelaB-
dami, 1688. — Rudolphi, G. A., ' Entozoorum sive verinium intes-
tinalium historia naturalis,' Amsterdam, 1808. — Idem, ( Ento-
zoorum Synopsis,' Berlin, 1819. — Van Beneden, P. J., 'Animal
Parasites and Messmates,' London, 1876.

Several of the above works, while professing to deal with
human parasites only, cover more or less of the whole ground
of helminthology. Leuckart's work is invaluable in this
respect ; and in the matter of literary references of a profes-
sional kind Davaine' s treatise is itself well nigh exhaustive.
In any ordinary volume it is not possible to give a complete
bibliography of parasitism. I make no pretension to do so
here ; nevertheless, the large number of modern memoirs that
I have received from the distinguished writers themselves,
enables me to render this part of my book very useful. As
second only in importance to the above-mentioned works may
be added the following — whether minor treatises, memoirs,
monographs, comprehensive articles, or reports of a general or
special character, respectively. As such it will be seen that
some of them are sufficiently comprehensive, and their mere
enumeration will enable the beginner to realise something like
a fair estimate of the scope of helminthology. In the case of
my own works I have ventured to add references to reviews

INTRODUCTION

and notices, because many of the latter contain valuable original
suggestions made by the various anonymous writers.

Bibliography (No. 2). — Bastian, H. C, " On the Anatomy
and Physiology of the Nematoids, parasitic and free/' ' Philo-
sophical Transactions/ 1865 (see also Bibliog., No. 60). —
Cobbold, T. 8., ' Worms ; a series of lectures on Practical Hel-
minthology/ London, 1872; Italian edition by Tommasi. Milan,
Florence, &c, 1873. — Idem, ' The Internal Parasites of our
Domesticated Animals,'1 London, 1873; Italian edit, by Tommasi,
Florence, 1874. — Idem, ' Tapeworms (Human), their Sources,
Varieties, and Treatment/ London, 3rd edit., 1875. Reviews
(1st and 2nd edit., with ' Threadworms '), in ' Brit, and For.
Med.-Chir. Review ' for 1867, p. 433 ; in ' Edin. Med. Journ.'
for 1866-67, p. 107; in ' Lancet/ Nov. 10th, 1866 ; in ' Popular
Science Review/ Oct. 1st, 1866; in ' Intellectual Observer/
Oct. 1866 ; in ' Med. Press and Circular/ Jan. 16th, 1867 ;
again in the ' Lancet/ for March 13th, 1867 ; and in ' Dublin
Quart. Journ. of Medical Science ' for 1867, 3rd edit.; in the
'Field/ Sept. 25th, 1875; and in ' Popular Science Review'
for Jan., 1876. — Idem, ' Catalogue of the Specimens of Entozoa
in the Museum of the Royal College of Surgeons of England/
London, 1866; noticed in the ' Lancet ' for March 24th, 1866,
p. 321. — Idem, " On the best Methods of displaying Entozoa in
Museums/' ' Journ. Linn. Soc./ vol. viii, p. 170. — Idem, 'New
Entozootic Malady/ &c, 1864; popular brochure, reviewed in
the 'Lancet/ Feb. 4th, 1865, p. 128 ; in the ' Athenaeum/ Jan.
21st, 1865, p. 87 ; in the ' British Med. Journal/ Jan., 1865 ; in
the ' Veterinary Review and Stockowners' Journal/ No. 2, New
Series, Feb., 1865, p. 76 ; in the ' Reader/ Feb. 4th, 1865,
p. 142 ; in 'Med. Times and Gaz.' for June 2nd, 1865 ; in the
'Field' for March 18th, 1865. — Idem, "Parasites of Man,"
forming a series of articles contributed to the ' Midland
Naturalist/ 1878-79. — Idem, " Notes on Entozoa contained in
the various Metropolitan Museums," in 'Lancet/ May 13th,
1865, p. 503. — Idem, " Report on Plica polonica, in refer-
ence to Parasites," in ' Pathological Soc. Trans./ 1866, p. 419.

Idem, " Report on Experiments respecting the Development

and Migrations of the Entozoa," 'British Assoc. Reports'
(Bath Meeting) for 1864, p. Ill; and briefly noticed in
'Lancet' for Sept. 24th, 1864.— Idem,, Miscellaneous observa-
tions, including " Note on Parasites in the Lower Animals,"
in 'Dub. Med. Press' for Feb. 11th, 1863, p. 154— Idem,

INTRODUCTION

« Vegetables, Fruits, and Water considered as sources of Intes-
tinal Worms in the ( Popular Science Keview ' for Jan., 1865,
p. 163.— Idem (anonymously), <c On Comparative Pathology
and Therapeutics " (in relation to Entozootics) ; leading art. in
' Lancet ' for Dec. 9th, 1865, p. 652.— Idem, " List of Bntozoa,
including Pentastomes, from animals dying at the Zoological
Society's Menagerie, between 1857-60 inclusive, with descrip-
tions of several new species/' ' Proc. Zool. Soc./ 1861. — Idem,
" Eemarks on all the Human Entozoa," ' Proc. Zool. Soc./
1862 ; abstracts in ' Brit. Med. Journ.' for 1862, and in
'Edinb. New Phil. Journ./ vol. xvii, new series, 1863, p. 145;
in Report of the ' Proceed, of the Brit. Assoc. at Cambridge/
1862. — Idem, " Our Food-producing Euminants, and the Para-
sites which reside in them ; being the Cantor Lectures of the
Society for the Encouragement of Arts, Manufactures, and
Commerce/' delivered in 1871, and pub. in the ' Journal of the
Soc. of Arts' for that year. — Davaine, C, " hes Cestoides,"
in f Diet. Encycl. des Sci. Med./ Paris, 1876 —Merth, C. J.,
' Untersuchungen ueber Nematoden/ Leipsic, 1863. — Heller, A.,
" Darmschmarotzer," in Yon Ziemssen's ' Handbuch/ Bd. vii,
1876 ; and in the American edition of the same, 1877. — Jones,
T. B., " List of Entozoa of Greenland," taken from Krabbe ;
' Arctic Manual/ 1875, p. 179. — Krabbe, E., ' Helminthologiske
Undersogelser,' Copenhagen, 1865. — LeucJcart, R., 'DieBlasen-
bandwiirmer und ihre Entwicklung/ Giessen, 1856. — Moquin-
Tandon, A., " Epizoa and Entozoa," in Hulme's edit, of his
' Elements of Medical Zoology/ London, 1871. — Nordmann, A.
von, ' Mikrographische Beitrage zur Naturgeschichte der wirbel-
losen Thiere/ Berlin, 1832. — Ohson, P., "Entozoa, iakttagna
hos Skandanaviska hafsfiskar.," Lund, ' Univ. Arsskrift/ 1867.
— Owen, B., " Entozoa," art. in Todd's ' Cyclopaedia of Anat.
and Physiol./ London, 1839. — Idem, " Entozoa," ' Lectures
(iv and v) on the Comp. Anat. and Physiol, of the Invertebrate
Animals/ London, 1855. — Pagenstecher, H. A., ' Trematoden-
larven und Trematoden/ Heidelberg, 1857. — Bhind, W.} ' A
Treatise on the Nature andCure of Intestinal Worms, &c.,' London,
1829. — Rolleston, G., " Characteristics of Nematelminthes and
Platyelminthes," in his ' Forms of Animal Life/ Oxford, 1870.
— Hclineider, A., ( Monographie der Nematoden/ Berlin, 1866.
— Hicbold, C. von, " Parasiten," art. in Wagener's 1 Handwor-
terbuch der Physiol., &c.,' 1845. — Idem, " Helminthes," Book v,
in Burnett's edit, of Siebold and Stannius' ' Comparative Ana-

INTRODUCTION

tomy/ London and Boston, 1854. — Thomson, A., " Entozoa/' in
the art. " Ovum/' in Todd's ' Cyclop, of Anat. and Physiol./
London, 1859. — Van Beneden, P. J., fMemoire sur les Vers
Intestineaux/ Paris, 1858. — Idem, " Les Vers Cestoides/'
' Mem. de l'Acad. Roy./ Brussels, 1850.— Verrill, A. E., " The
External and Internal Parasites of Man and the Domestic
Animals," ' Rep. of Board of Agriculture/ Connecticut, U.S.,
1870. — Von Baer, K. E., ' Observations on Entozoa / in an
analytical notice of his article " Beitrage zur Kentniss der
niedern Thiere," from ' Nova Acta Nat. Cur./ torn, xiii, in the
'Zool. Journ/ vol. iv, p. 250, 1828-29. — Wagener, G. B., ' Bei-
trage zur Entwicklungsgeschichte der Eingeweidewiirmer/
Haarlem, 1857. — Weinland, D. F., ' An Essay on the Tapeworms
of Man/ Cambridge, U.S., 1858.

BOOK I.

PARASITES OF MAN.

Whatever notions people may entertain respecting the
dignity of the human race, there is no gainsaying the fact that
we share with the lower animals the rather humiliating privilege
and prerogative of entertaining a great variety of parasites.
These are for the most part entozoal in habit. As the parasites
are apt to cause suffering to the bearer, a superstitious age
sought to interpret their presence as having some connection
with human wrong-doing. We can now afford to smile at such
erroneous ideas. The intimate relation subsisting between
parasitic forms dwelling in man and animals, and their inter-
dependence upon one another, alone suffices to preclude the
idea that parasites have been arbitrarily placed within the
human bearer. It would seem, indeed, that our existence
is essential to the welfare and propagation of certain species
of parasites. Possibly it is only by accepting the hypo--
thesis of " Natural Selection " that we can escape the somewhat
undignified conclusion that the entozoa were expressly created
to dwell in us, and also that we were in part designed and
destined to entertain them. View the matter as we may, the
internal parasites of man and animals strictly conform to a few
well-known types of structure, but these types branch out into
infinitely varied specific forms. The vulgar mind sees nothing-
attractive in the morphology and organisation of a parasitic
worm, and common-place conceptions of the beautiful cannot
be expected to embrace within their narrow grasp the marvel-
lous harmony and order that pervade the structure and economy
of the individual members of this remarkable class of beings. ',

PARASITES OP MAN

SECTION I. — Trematoda (Flukes).

Fasciola hepatica, Linneus. — The first form I have to con-
sider is the common liver fluke. The part this entozoon plays
in the production of disease will be fully stated when treating
of the parasites of the sheep and other ruminants. About
twenty instances of its occurrence in the human body have been
recorded. It has been found beneath the skin in the sole of
the foot (Griesker), and also under the scalp (Harris) , and
behind the ear (Fox). Its more frequent seat is in the liver
and gall-ducts (Pallas, Brera, Bidloo, Malpighi) and gall-bladder
(Partridge). The alleged cases by Bauhin, Wepfer, and Chabert
are spurious, as is probably also that given by Mehlis. DuvaPs
case appears to be genuine, but the occurrence of the worm in
the portal vein was accidental. Dr Murchison has recorded a
case, occurring at St Thomas's Hospital, where a solitary
specimen was found in the liver. Dr H. V. Carter also met
with the worm in a young Hindoo.

In the second half of the present work I shall reproduce
Blanchard's admirable figure of the sexually mature worm
(Fig. 61), accompanied by a categorical statement respecting
the known facts of development. In this place, however, I
may observe that the cases recorded by Griesker, Harris, and
Fox had clearly pointed to the circumstance that the higher larvas
of this fluke must be armed cercariaa, otherwise they could not
have bored their way through the human skin. As we shall see,
Dr Willemoes-Suhm's investigations have furnished evidence
as to the truth of this supposition. For anatomical details I refer
to my introductory treatise. In the adult state the liver fluke
has been known from the earliest times. "We have clear
evidences that it was described by Gabucinus in the year 1547,
and also subsequently by Cornelius Gemma, who, in a work
published some thirty years later, refers to an epizootic disease
prevalent in Holland during the year 1552, and which was very
justly attributed to the parasite in question. After this date
many writers described the liver fluke more or less accurately,
and entire volumes were devoted to the consideration of the
formidable disease which it occasions. The nomenclature of
the parasite has been a subject of controversy. Amongst
naturalists in general the common liver fluke is often de-
scribed under the combined generic and specific name of

Tli EM A TOD A

Distoma hepaticumj but the title is both incorrect and inappro-
priate. The proper generic appellation of this parasite is
Faseiola, as first proposed by the illustrious Linneus (1767) and
subsequently adopted by F. Miiller (1787), Brera (1811), Eam-
dohr (1814), and others. Unfortunately Retzius (1786) and
Zeder (1800) changed the generic title without good cause,
and the majority of writers, following their authority, refused
to employ the original name, although a consideration of the
distinctive types of structure severally displayed by the genera
Distoma and Fasciola fairly demanded the retention of the
Linnean title. In later times M. Blanchard (1847) strongly
advocated the original nomenclature, and I have myself con-
tinually urged its adoption. On somewhat different grounds
Professor Moquin-Tandon followed the same course.

In the sexually mature state the liver fluke commonly
measures three fourths of an inch in length, occasionally reach-
ing an entire inch or even sixteen lines ; its greatest breadth
also varying from half an inch to seven or eight lines trans-
versely ; body very flat, presenting distinct dorsal and ventral
surfaces, frequently curled toward the latter during life ; upper
or anterior end suddenly constricted, produced and pointed
in the centre, forming the so-called head and neck ; posterior
extremity less acuminated, sometimes rounded, or even slightly
truncated ; margins smooth, occasionally a little undulated,
especially towards the upper part ; oral sucker terminal, oval,
rather smaller than the ventral acetabulum, which is placed
immediately below the root of the neck ; reproductive orifices
in the middle line, a little below the oral sucker ; intromit-
tent organ usually protruded and spirally curved; a central,
light-coloured space, covering two thirds of the body from
above downwards, marks the region of the internal male re-
productive organs, being bordered on either side and below by
a continuous dark band, indicating the position of the so-called
yelk-forming organs; a small, brown-coloured, rosette-like
body situated directly below the ventral acetabulum, marks
the limits of the uterine duct ; a series of dark lines, branching
downwards and outwards on either side, indicate the position of
the digestive organs ; general color of the body pale brownish
yellow, with a slight rose tint. The surface of the body, though
smooth to the naked eye, is clothed throughout with small
epidermal spines which diminish in size towards the tail.

If any argument were necessary to show how desirable it is

I'ARASITES OP MAN

to furnish full descriptions of the commoner kinds of parasite,
I could adduce numerous instances that have been brought
under my notice where professional men and others have been
entirely mistaken as to the essential nature of their parasitic
finds. Thus, I have known an instance where a great authority
on the diseases of dogs has persisted in asserting for the free
proglottides of a tapeworm a nematode origin; and, in like
manner, human tapeworm-segments have frequently been
mistaken for independent fluke parasites. One of the most
remarkable instances of this kind is that which I have else-
where described as an error on the part of Dr Chabert. My
reasons for so regarding his interpretation of the facts observed
by him stand as follows :

In the ' Boston Medical and Surgical Journal ' for the years
1852-53-54, Dr J. X. Chabert described several cases of Taenia,
and he averred that the tapeworms were associated with
numerous specimens of Distoma hepaticum. The passage of dis-
tomes by patients during life was even regarded by Dr Chabert
as indicative of the presence of Taenia within the intestines.
Surely, I remarked, Dr Chabert was mistaken. Are not these
so-called distomes the well-known proglottides ? Not willingly
doubting Dr Chabert's statements, but desirous, if possible, of
verifying the accuracy of his conclusions, I wrote to him (March
22nd, 1864) requesting the loan of a specimen, but I was
not fortunate enough to receive a reply. In the " Case of
Taenia " in a boy four and a half years old, given in the 49th
vol. of the journal, Dr Chabert writes as follows : — " In con-
sequence of his passing the Distoma hepaticum, I concluded he
must be afflicted with Taenia." Further on it is added, that
the administration of an astringent injection " caused the dis-
charge of innumerable small worms (Distoma hepaiicum)." I
think this is quite decisive. The idea of "innumerable" flukes
being expelled in this way is altogether out of the question.

The only genuine case in which any considerable number of
Distomata, of this species, have been observed in the human
subject is the one recently recorded by Dr Prunac. In this
instance two flukes were vomited along with blood immediately
after the administration of salines (sel de Seignette), and about
thirty were passed per anum. On the following day, some
tapeworm proglottides having been evacuated, both salts and
male-fern extract were administered. This caused the expulsion
of an entire tapeworm, and also about twenty more flukes. Not-

TUEMATODA

withstanding this successful treatment the heematemesis returned
in about a month, when, finally, three more flukes were vomited
and the bleeding ceased. Had not the parasites been sub-
mitted for identification to a competent observer (Prof.
Martins, of Montpellier), some doubt might have been enter-
tained as to the genuineness of this remarkable case. In refer-
ence to Dr Prunac's comments on the facts of fluke-parasitism
in man, I will only remark that Dr Kerr's Chinese cases, to which
he refers, were probably due to Distoma crassum and not to D.
hepaticum. The Chinese flukes will be noticed below.

Bibliogeaphy (No. 3) . — Full references to details of the cases
by Partridge, Fox, and Harris are given in Appendix B. to
Lankester's Edit, of Kuchenmeister3 's Manual. See also the
works of Davaine and Leuckart (I. c. Bibl. No. 1). — Garter,
H. V., " Note on Distoma hepaticum " (from a patient under the
care of Mr Pandoorung), ' Bombay Med. and Physical Soc.
Trans/ (Appendix), 1862. — Ghabert, J. X. (quoted above).
Murchison, G., ( Clinical Lectures on Diseases of the Liver, (2nd
Edit., Appendix), London, 1877. — Prunac, De la Douve ou
Distome hepatique chez l'homme ; in ' Gazette des H6pitaux '
for December, 1878 (p. 1147). For further references in this
work, see Bibliog. No. 49.

Distoma lanceolatum, Mehlis. — At least three instances of
the occurrence of this small fluke in the human body have been
observed. The authority for these cases rests, severally, with
Bucholz, who found them in the gall bladder in considerable
numbers at Weimar; with Chabert, who expelled a large
number from the intestines of a girl in France ; and with
Kiichner, who obtained forty-seven specimens from a girl in
Bohemia. Probably many similar instances have been over-
looked, and Kuchenmeister hints that Duval's parasites (above
mentioned) may have been this species. Although this worm
will again be incidentally noticed in connection with bovine
parasites (and its ciliated larvae will also be referred to when
discussing the characters of the embryo of Bilharzia), I here
subjoin a diagnosis of the characters of the adult parasite.
The lancet-shaped liver fluke is a small flat helminth, measur-
ing rather more than the third of an inch in length, and about
one line and a half in breadth, being also especially charac-
terised by its lanceolate form ; the widest part of the body
corresponds with a transverse line drawn across the spot
where the vitellaria terminate below, and from this point, on

PARASITES OF MAN

either side, the width of the animal becomes gradually narrowed
towards the extremities; both ends are pointed, but the inferior
or caudal one more obtusely than the anterior or oral end ; the

general surface is smooth throughout,
and unarmed ; the reproductive orifices
are placed in the central line imme-
diately in front of the ventral sucker,
and below the point at which the in-
testine bifurcates ; the oral sucker is
nearly terminal, and ~" in breadth, the
ventral acetabulum being about the same
diameter ; the testes form two lobed
organs placed one in front of the other
in the middle line of the body and
directly below the ventral sucker; the
uterine canal is remarkably long, form-
ing a series of tolerably regular folds,
which occupy the central and hinder
parts of the body, reaching almost to
the caudal extremity. The vitelligene
glands cover a limited space, on either
side of the centre of the body near the
margin. The foramen eaudale commu-
nicates with a contractile vesicle, which
passes upwards in the form of a central
trunk-vessel, early dividing into two
main branches ; these latter reach as far
forwards as the oesophageal bulb, oppo-
site which organ they suddenly curve
upon themselves, retracing their course
for a considerable distance backwards ;
the digestive canals are slightly widened
towards their lower ends, which occupy
a line nearly corresponding with the
commencement of the lower fifth of the
body; the ova are conspicuous within
the uterine folds, which present a dark brownish color in
front, passing to a pale yellow color below.

In reference to Kichner's remarkable case I reproduce an
abstract of it from Leuckart's account ('Die menschlichen
Parasiten/ Bd. i, s. 608), the original particulars of which were
communicated to Leuckart by Dr Kichner himself : —

Tig. 1.— The lancet-shaped fluke
(Disloma lanceolatvm), Bhowing
the disposition of the digestive
and reproductive organs inter-
nally. Viewed from behind ; mag.
about 12 diameters. After Blan-
chard.

TREMATOUA

" Dr Kichner's patient was a young girl, the daughter of the
parish shepherd at Kaplitz, having been accustomed to look
after the sheep ever since she was nine years old. The
pasture where the animals fed was enclosed by woods, being
traversed by two water dykes, and being, moreover, also sup-
plied by ten little stagnant pools. These reservoirs harboured
numerous amphibia and mollusks (such as Lymnceus and Palli-
dum), and the child often quenched her thirst from the half
putrid water. Probably she also partook of the watercresses
growing in the ditches. At length her abdomen became much
distended, the limbs much emaciated, and her strength de-
clined. Half a year before death she was confined to her bed,
being all the while shamefully maltreated by her step-mother.
Dr Kichner only saw her three days before her death, and
ascertained that she had complained of pain (for several years)
over the region of the liver. A sectio cadaveris was ordered
by the Government, when (in addition to the external evidences
of the cruel violence to which the poor creature had been sub-
jected) it was found that she had an enormously enlarged liver,
weighing eleven pounds, The gall-bladder which was very
much contracted and nearly empty, contained eight calculi and
forty-seven specimens of the Distoma lanceolatum, all of which
were sexually mature."

As I have remarked in a former comment on this singular
case, one can have no difficulty in arriving at the conclusion
that these parasites were obtained from the girl's swallowing
trematode larvae, either in their free or in their encysted
condition. Leuckart says it was not possible to ascertain
whether the parasites had any connection with the gall-stones,
or whether the two maladies, so to speak, were independent
of each other; yet this question might possibly have been
solved if the calculi had been broken up in order to ascertain
their structure. It is just possible that dead distomes may
have formed their nuclei, and if so, the circumstance would,
of course, point to the worms as the original source of the
malady.

So far as I am aware, the actual transformations undergone
by the larvaa of Distoma lanceolatum have not been observed.
The Planorbis marginatus has been confidently referred to as
the intermediate bearer of the cercarias of the common fluke
and Leuckart supposes that the same mollusk harbours the
larvae of this species. The ciliated embryos carry a boring

PARASITES OF MAN

spine or tooth, and it is most probable that the higher larva?
are similarly armed.

Bibliography (No. 4). — Kichner (see Leuclcart), quoted above.
— Cobbold, 'Entozoa' (p. 187). — The case byBucholz (reported
as one of Fasciola hepatica) is given by Jordens in his work
(quoted by Diesing and Leuckart) ' Entomologie und Heltnin-
thologie des menschlichen Korpers ' (s. 64, tab. vii, fig. 14),
1802. — Chabert's French case is quoted by Iludolphi in his ' Ento-
zoorum sive vermium/ &c. (loc. cit., Bibl. No. 1), p. 326, 1808.

Distoma crassum, Busk. — This large species was originally
discovered by Prof. Busk in the duodenum of a Lascar who
died at the Seamen's Hospital, 1843. It, however, remained
undescribed until 1859, when, with the discoverer's approval, I
gave some account of it to the Linnean Society.

Of the fourteen original specimens found by Mr Busk,
several have been lost. The one that he himself gave me
I handed over to Prof. Leuckart, and it is figured in his
work (f Die mensch. Par./ s. 586). A second is preserved in the
museum attached to the Middlesex Hospital, and a third is con-
tained in the Museum of the Royal College of Surgeons. This
last-named specimen is the best of the original set. It supplied
me with the few details of structure figured in outline in my
'Introductory Treatise' (fig. 42, p. 123), published in 1864;
and it also in part formed the basis of the description of the
species communicated to the Linnean Society in June, 1859
("Synopsis of the Distomidge/' p. 5, ' Proceedings/ vol. v).
The late Dr Lankester, it is true, was the first to give a dis-
tinctive title to this entozoon {Distoma Bushii) ; but as the
discoverer objected to this nomenclature, and as Dr Lankester's
proposed terms were unaccompanied by any original description,
I requested Mr Busk to suggest a new name for the worm,
which he accordingly did. As I subsequently pointed out, Von
Siebold had already employed the compound title Distoma eras-
sum to designate a small fluke infesting the house-martin
(Hirundo urbica) ; but for reasons similar to those which con-
tributed to set aside Dr Lankester's nomenclature, the title
adopted in my synopsis at length came to be recognised by
v Leuckart and by other well-known helminthologists. Before
this recognition took place, Dr Weinland, of Frankfort, had so
far accepted Lankester's nomenclature as to call the species
Dicroccelium Bushii. In my judgment there are no sufficient
grounds for retaining Dujardin's genus. Further, 1 may

TEEMATODA

observe that, in addition to the above-mentioned specimens,
two others are preserved in the Museum at King's College. Thus,
only five out of the fourteen specimens are still in existence.

No well-authenticated second instance of the occurrence of
this worm took place until the year 1873, when a missionary
and. his wife from China consulted Dr George Johnson re-
specting parasites from which they were suffering. After a
brief interval, both of Dr Johnson's patients were by an act of
courtesy on the part of this eminent physician placed under my
professional care. I need hardly add that Dr Johnson had
from the very first recognised the trematode character of the
parasites. From the patients themselves I ascertained that
they had been resident in China for about four years. During
that period they had together freely partaken of fresh vege-
tables in the form of salad, and also occasionally of oysters, but
more particularly of fish, which, in common with the oysters,
abound in the neighbourhood of Ningpo. From their state-
ments it appeared to me that to one or other of these sources
we must look for an explanation of the fact of their concurrent
infection. Fluke larvae, as we know, abound in mollusks and
fish ; but whether any of the forms hitherto found in oysters or
in fish have any genetic relation to the flukes of man, is a ques-
tion that cannot very well be settled in the absence of direct
experimental proof. I should add that it was not until after
their visit to the interior of the country, some 130 miles distant
from Ningpo, that the symptoms (which Dr Johnson in the first
instance, and myself subsequently, considered to have been due
to the presence of the parasites) made their appearance. Whilst
in the country the missionary and his wife freely partook of
freshwater fish, and on one occasion they received a quantity of
oysters that had been sent up from Ningpo. The husband
assured me that the fish were always thoroughly well cooked.

If it be asked what were the symptoms produced, I can only
furnish such few and hitherto unpublished particulars as the
missionary himself supplied. I need hardly say that he was a
highly cultured and intelligent gentleman, since only such
persons are chosen for missionary work in China.

From inquiries made by me on the 29th of January, 1875, I
learnt that they left Ningpo in November, 1872, and travelled
thence 130 miles into the interior of the country. In the
following September, or about ten months subsequently, the
missionary was attacked with diarrhoea, which persisted until

PARASITES OF MAN

expulsion of some of the parasites had occurred. Accord-
ing to the patient's statements this result, so far, was entirely
due to his having been placed on a milk diet; this course
of treatment having been recommended by Dr Henderson,
of Shanghae. The patient himself always suspected the pre-
sence of intestinal worms of some sort or other, although a
Japanese doctor laughed at the idea of such a thing. Some
other doctor treated this missionary for parasites, administering
both male-fern and santonine without effect.

It was not until several months had elapsed that his wife
was attacked with diarrhoea. In both cases there was more or
less flatus. The motions were white, and there were other indica-
tions implying that the liver was affected. Later on, symptoms
of indigestion, with heartburn, set in and became very severe.
Streaks of blood appeared in the fasces, but there was no
dysentery. For the most part these symptoms were attributed
to the effects of climate.

When, in the month of February, 1875, I saw the missionary
a second time, professionally, I found that all the old sym-
ptoms had returned. He had a foul tongue, the surface of the
body was cold, he felt chills, and the pulse, though regular,
registered ninety-six to the minute. Indigestion, nausea, head-
ache, and diarrhoea had reappeared. Notwithstanding these
febrile symptoms, so satisfied was the patient himself that all
his ailments were entirely due to the presence of parasites, that
I felt inclined to take the same view of his* case. Accordingly
my attention was principally directed to an effort for their expul-
sion ; and in this view I ordered an aloetic pill followed by a
castor-oil emulsion. This having no effect, I subsequently
prescribed aloes and assafoetida pills, followed by scammony
mixture. The action of the latter drug did not occasion griping,
but, although efficient, led only to negative results. I should
mention that in the patient's judgment none of the vermifuges
administered to him at any time had exerted any influence in
the expulsion of the flukes. He was still thoroughly impressed
with the notion that the milk diet, ordered by Dr Henderson,
was the sole cause of their expulsion.

As even a missionary could not live by milk alone I insisted
upon a more substantial diet. The milk, indeed, had occasion-
ally been supplemented by Liebig's extract of meat and by light
farinaceous food. When I last saw him neither he nor his wife
had passed any more flukes, but they did not feel satisfied that no

TlvEMATODA

more guests remained. Somewhat improved in general health,
the missionary resolved to go back to his duties in China. I
expressed my fears, however, that his strength would prove
unequal to the work.

From the size and almost leathery texture of the two flukes
which were in the first instance submitted to my notice, I at
once recognised the species ; but as they were spirit-specimens,
I requested that if any more examples were obtained they
should be sent to me in the fresh state. Fortunately others
were brought in a few days, when, from an examination
conducted whilst they were still fresh, I was able to make out
several details of structure which had hitherto escaped notice.
Altogether I secured seven specimens, three of them being in a
mutilated condition. In what way these mutilations (as shown
by my dried specimens) occurred I have not been able to make
out, either by personal observation or by questioning the
bearers. Two of the parasites look as though portions had
been carefully excised near the centre. The new facts I have
gleaned were derived from the examination of two compara-
tively small specimens, one of which, dried, has, by Prof.
Kolleston's desire, been deposited in the anatomical department
of the University Museum at Oxford. When I took occasion
to bring some of the new specimens under Mr Busk's attention,
he at once recognised them as referable to the species he had
long ago discovered.

The earliest literary notice of Distoma crassum appeared in
Dr Budd's classical treatise ' On Diseases of the Liver •/ and in
it the author correctly stated, from data supplied by Mr Busk,
that these human flukes were " much thicker and larger than
those of the sheep," being, it is added, from "an inch and a
half to near three inches in length." The longest of my recent
specimens, however, scarcely exceeds two inches, whilst the
smallest and most perfect (the one at Oxford) measures less
than an inch from head to tail. The greatest width of my
broadest specimen is little more than half an inch, or
None of the twelve examples that I have examined approach
the length of three inches; but Mr Busk assured me that,
judging from his recollection, some of his specimens were even
longer than that. I fear, nevertheless, that the estimate given
in my Synopsis is somewhat exaggerated • at all events it is so
for average specimens.

The new anatomical facts made out by me bear referenco

PARASITES OF MAN

principally to the reproductive apparatus. What else I have
observed is for the most part confirmatory of the statements
made by Mr Busk. In particular, his brief account of the
position and character of the digestive organs was not only
confirmed by my earlier examinations, but is now re-verified.

In the representation
given in my ' Intro-
duction ' I showed in
dotted outline two
large organs which I
supposed to be the
testes. I distinctly ob-
served radiating lines
proceeding from the
centre in each ; but
I could not discover
the slightest trace of
any limiting border to
either organ. I now
found in the same
position two nearly
circular flattened
masses with clearly
defined limits (i, k).
No doubt could be
entertained as to the
testicular character of
the lower organ (k).
In the original draw-
ing I further indi-
cated the presence of
a third and much
smaller globular mass,
which I termed the

Pi„ .2 —The large human fluke (Distoma crassum). a, Oral sucker ;
* intestine; c, ctecal end of same; d, reproductive papilla;
e uterine rosette (the folds of which are not branched) ; /, one of
the folds (in profile); g, vitellarium ; A, hernial protrusion (the
result of an injury to the specimen); i, upper testis; streaks
or layers of seminal fluid which have escaped by rupture
and assumed a branched appearance ; k, lower testis uninjured
(but Blightlv altered in outline from flattening) ; I, ventral sucker.
Magnified 2 diameters. Original.

ovary ; but what I
supposed to represent
this organ in the
particular specimen
from which the accompanying illustration was drawn turns out
to be merely a hernial protrusion resulting from injury (h).
The radiating, broad, and branching seminal ducts are beauti-
fully distinct in one of my specimens, forming the most attrac-

THNMATODA

tive feature of the parasite's organisation (fc). In consequence
of injury to the specimen which is here drawn, the upper testis
(i) displays no seminal tubes. I made out the female reproduc-
tive organs with more completeness. In the outline drawing
given in my introductory treatise I had indicated the probable
position of the uterine folds ; reducing the organ to the simplest
expression of what I concluded must obtain in the normal
condition. My conjecture was perfectly correct. The uterus
consists of irregularly folded tubes, which, though here and
there apparently branching from a central tube, are in reality
folded evenly upon themselves. The oviduct can be distinctly
traced to its outlet in the reproductive papilla, which, as usual
in true Distomes, is placed in the middle line, immediately
above the ventral sucker. In my examination of Mr Busk's
original specimens I could not find the slightest trace of
vitelligene organs ; but in my fresh examples I not only
obtained proof that these organs were largely developed,
but that their limitations could be fixed with accuracy (g g).
They consisted of two large elongated masses, one on either
side of the body, occupying about two thirds of the entire
length of the parasite. Their yolk-vesicles were distinctly
seen ; but the main efferent canals were only here and there
traceable. Clearly, the position and character of the yolk-
forming glands of this large human fluke are quite unlike those
of any of its congeners. This fluke is a remarkably fine
species, and, when viewed in the fresh state with a powerful
pocket-lens, presents a most striking appearance. I did not
observe any cutaneous spines. I found the eggs to present an
average long diameter of about ^5", by 3^" in breadth. They
are therefore somewhat smaller than those of the common fluke.
In the specimen preserved in' the Hunterian Museum there was
complete evidence of the presence of an excretory outlet at the
caudal extremity ; but I did not succeed in finding any trace of
the water- vascular system higher up. I have no doubt, how-
ever, that it exists.

As regards the affinities of Distoma crassum, it is clear that
this Trematode has little in common either with the liver-fluke
of cattle and sheep (Fasciola hepatica), or the still larger species
obtained by me from the giraffe (Fasciola gigantea). The
simple character of the digestive tubes obviously connects it
more closely with the lancet-shaped fluke {Distoma lanceolatum) ,
the last-named parasite being, as already shown, an occasional

PARASITES OP MAN

resident in the human liver, where its presence, moreover,
undoubtedly contributed towards the production of the fatal
result.

In my remarks on the missionary's diet it is hinted that
the Ningpo oysters may have played the role of intermediary
bearers to the parasite in question ; and as tending in some
measure to strengthen this notion, it should be borne in mind
that Mr Busk's original fluke-bearer came from the east. It
is not improbable that the Lascar host may have partaken of
the same particular species of fish or shell-fish that the
missionary and his wife partook of. Be that as it may, the
frequency of the occurrence of Trematodes and their larvae in
marine mollusks is well known. According to Woodward,
several species of oyster- are sold in the Indian and Chinese
markets. Thus, it would require the skill of a malacologist to
determine the particular species of Ostrea to which the Ningpo
oysters should be referred.

Mons. Giard is of opinion that the singular larvae known as
Bucephali attain sexual maturity in sharks and dog-fishes ;
therefore it is extremely unlikely that the Bucephali should have
been in any way concerned in the infection of our missionary
and his wife ; nevertheless there remains the probability that
these human bearers swallowed other kinds of Trematode
larvae when they consumed the Ningpo oysters. Moreover, if
it should happen that none of the other larvae occurring in
oysters are capable of developing into flukes in the human
territory, it yet remains highly probable that some one or other
of the various encysted (and therefore sexually immature)
Trematodes known to infest marine fishes will turn out to be
the representative of our Distoma crassum. In this connection
we must not forget that the flesh of the Salmonidee forms the
probable source of human Bothriocephali ; and there is some
likelihood that salt-water fishes, if not actually the primary,
may become (after the manner explained by M. Giard) the
secondary intermediary bearers of fluke-larvae. At all events,
I am inclined to look to the Ningpo oysters, or to some
other of the various species of marine shell-fish sold in eastern
markets, as the direct source of Distoma crassum; for, in
addition to the bucephaloid cercarians, we have abundant
evidence of the existence of other and more highly developed
fluke-larvae in marine bivalve mollusks.

In this connection I will only further observe that we possess

TliEMATODA

very little knowledge of the parasites which take up their abode
in the viscera of savages. This ignorance results partly from
the fact that these untutored races, as proved by the statements
of Kaschin and others, actually, in the matter of severe sym-
ptoms, suffer much less from the presence of intestinal worms
than their civilised fellow-men do. The subject is worthy of
further attention, but no one, so far as I am aware, has cared
to institute the necessary inquiries in a methodical way. I
strongly suspect that several of the human parasites which we
now consider to be rare would be found to be abundant if by
means of post-mortem examinations and other methods of in-
vestigation we could be made acquainted with the facts of hel-
minthism as they occur amongst the raw-flesh and fish-eating
savage tribes. Of course any person, notwithstanding the
utmost care and cleanliness, as in the cases before us, may
contract a noxious parasite ; nevertheless, speaking generally,
it may be said that the measure of internal parasitism affecting
any given class of people bears a strict relation to the degree
of barbarism shown by such persons in their choice of food and
drink, and in their manner of eating and drinking. This
statement, if true, is not destitute of sanitary importance ;
moreover, it applies not alone to ourselves, but also to all the
domesticated animals that serve our wants. Cleanliness is just
as necessary for their welfare as for our own.

In the spring of 1878 my patients returned from China.
They had experienced fresh attacks from the parasite ; more-
over, one of their children, a little girl, was also victimised by
the same species of fluke. Thus, in one family I have en-
countered three cases of fluke-helminthiasis due to Distoma
crassum ! One of the worms passed by the little girl joer anum
is now in my possession. It not only shows the upper testis
perfectly, but also the many times transversely folded, simple,
uterine rosette which is certainly not branched. There are
also traces of an organ which I take to be the cirrhus-pouch ;
but I have never seen the penis protruded externally.

For the purposes of diagnosis I subjoin the following cha-
racters. The Distoma crassum is a large, flat helminth vary-
ing from an inch and a half to two and a half inches in length,
and having an average breadth of five eighths of an inch ; it is
especially also characterised by its uniform and considerable
thickness, combined with the presence of a double alimentary
canal which is not branched ; the body is pointed in front, and

28 PARASITES OF MAN

obtusely rounded posteriorly ; the integument being smooth and
unarmed ; the reproductive orifices placed immediately above
the ventral sucker ; the testes form two large rounded organs,
situated below the uterine rosette, and disposed in the middle
line, one in front of the other; the uterine folds occupy the
front part of the body ; near the lateral margins there are two
large vitelligene glands, one on either side of the intestinal
tube ; the excretory organ probably consists of a central trunk
with diverging branches, opening below.

Bibliography (No. 5). — Budd, original notice in his 'Dis-
eases of the Liver/ 2nd edition, quoted by Lankester in Appen-
dix B to Kiichenmeister's ' Manual of Parasites/ p. 437, 1857.
— Cobbold, T. 8., " Synopsis of the Distomidce," in ' Journ. of
the Proceed, of the Linnean Soc./ vol. v, Zool. Div., 1860
(original description p. 5). — Idem, ' Entozoa/ p. 193, 1864. —
Idem, " Remarks on the Human Fluke Fauna, with especial
reference to recent additions from India and the East," the
' Veterinarian/ April, 1876. — Idem, " On the supposed Rarity,
Nomenclature, Structure, Affinities, and Source of the large
Human Fluke (D. crassum) ," ' Linn. Soc. Journ./ vol. xii,
Zool. Div., 1876, p. 285 et seq. — Idem, " Observations on the
large Human Fluke, with notes of two cases in which a mis-
sionary and his wife were the victims," the 'Veterinarian/
Feb., 1876. — Idem, " The new Human Fluke," in a letter
published in the 'Lancet/ Sept., 1875. — Leidy, in 'Proceed.
Acad. Nat. Sciences of Philadelphia / see also Dr McConnelPs
paper quoted below (Bibl. No. 6). — LeucJcart, 1. c, Bd. I, s. 560.
— Weinland, 1. c. (Bibl. No. 2), Appendix, p. 87.

Distoma Sinense, Cobbold. — The discovery of this species is
due to Prof. J. F. P. McOonnell, who " on the 9th of Sept., 1874,
found a large number of flukes in the liver of a Chinese, ob-
structing the bile ducts." The species measures T75" in length,
by \" in breadth, the eggs being gj," by Dr McConnell

showed in his original memoir that the worm cannot well be
confounded with Fasciola hepatica, with Distoma lanceolaturn ,
or with D. conjunctum. In this conclusion he was supported
by Dr.T. R. Lewis, who examined the specimens with him.
In a letter communicated to the 'Lancet/ quoted above, I
proposed the nomenclature here given; but Prof. Leuckart,
unaware of this step, afterwards suggested the terms Distomum
spatulatum. Later on I received numerous specimens from
Calcutta, the examination of which enabled me to confirm the

THEMATODA

accuracy of the original description. As regards the male
organs in the subjoined figure, it will be seen, by comparing the
lettering and references, that I have in-
terpreted the facts of structure .some-
what differently from Prof. McOonnell.

In the month of December, 1874, a
Chinese died in the Civil Hospital at
Port Louis, Mauritius, whilst he was
under the care of Dr William Mac-
gregor, chief medical officer of the
Colony of Fiji. The post mortem revealed
the presence of a very great number of
flukes in the bile- ducts- Dr Macgregor
described these parasites with great care,
and having favored me with a copy of
his manuscript I at once recognised the
worms to be identical with the species
discovered by McConnell. I also re-
ceived through Dr Henry Clark, of Glas-
gow, two Mauritius specimens, which
when compared with the Calcutta exam-
ples proved to be specifically identical.
Dr Macgregor's paper, communicated to
the Glasgow Medico-Chirurgical Society,
gives full particulars of the helminthiasis
associated with this parasite, whilst both
his and Prof. McConnell's account of the
structure of the worm are remarkably
complete in details, and well illustrated.

It is not a little CUrioUS to notice that Fi g. 3.— The Chinese fluke (Dis-
1,1 i i • i • toma Sinense). a, Oral sucker ; b,

although these parasites were obtained m
countries far removed from China, they
were in both instances taken from
Chinese ; moreover, from the statements
of Macgregor, it appears very probable
that the parasites in question are a com-
mon source of liver disease. Without doubt oriental habits
are eminently favorable to fluke infection, for we are now
acquainted with four species of flukes whose geographical range
is limited to eastern parts.

Bibliography (No. 6). — McConnell, J. F. P., " Eemarks on
the Anatomy and Pathological relations of a new species of

oesophageal bulb ; c, intestine ;
c', caeca! end ; d, veutrnl sucker ;
e, genital pore ; f, uterine
folds ; g, ovary ; h, vitellarium ;
t, vitelligene duct; k, upper
seminal reservoir ; /, testes ;
tn, lower seminal pouch ; o,
vas deferens; p, pulsatile vesi-
cle; p', water vessel. After
McConnell.

PARASITES OF MAN

Liver-fluke/' ' Lancet/ Aug. 1875 j repr. in the ' Veterinarian/
Oct., 1875; also in the ' Lancet/ March 16th, 1878, p. 406. —
Macgregor, W., " A new form of Paralytic Disease, associated
with the presence of a new species of Liver Parasite (Distoma
Sinense)," 'Glasgow Med. Journ/ for Jan., 1877 ; also in the
' Lancet 'for May 26th, 1877, p. 77$.— Cobbold, T. S., in a note
to the ' Lancet/ Sept., 1875, and in the Appendix to Macgre-
gor's paper, p. 15, 1877. — LeucJcart, B., 1. c, Bd. ii, s. 871, 1876.

Distoma conjunctum, Oobbold. — The little fluke which I first
discovered in the gall-ducts of an American fox (Oanis fulvus)
was fourteen years afterwards obtained from pariah dogs in
India by Dr. T. E. Lewis (1872) ; but it remained for Prof.
McConnell to show that this entozoon also invades the human
subject (1874). A second instance of its occurrence in man
was recorded in 1876. We all figured the worm, and in
respect of general details our descriptions for the most part
agreed (fig. 56). The worms from the dog and fox gave an
average of \" in length, but the majority of those found by
McConnell in man were fully §" from head to tail.

Writing in the spring of 1876 Dr McConnell says : — " In
the t Lancet ' for the 21st of August, 1875, I published the
description of a new species of liver-fluke found in the bile-
ducts of a Chinaman (sic) who died in this hospital. Dr Spencer
Cobbold has very kindly interested himself in this discovery,
and proposed the name of Distoma Sinense for the new fluke.
This discovery (in September, 1874) has stimulated me to pay
still greater attention to the morbid conditions of the biliary
canals in our post-mortem examinations ; but, although more
than 500 autopsies have been conducted since that date, I have
not met with another instance of distomata in the liver until
within the last fortnight. On the 9th of January, 1876, in exa-
mining the liver of a native patient who had died in the hospital,
I again found a large number of flukes in the bile-ducts, and
having carefully examined many specimens, I recognise the
species as the D. conjunctum of Cobbold. Dr Cobbold dis-
covered this fluke in 1858 ; but, as far as I am aware, the human
liver has never hitherto been found infested by these parasites,
and this will give general interest and importance to the follow-
ing case."

" Jamalli Khan, a Mahommedan, aged twenty-four, admitud
into the hospital on the 25th of December, 1875. He is a
resident of Calcutta, and an ordinary labourer (coolie). He

TKEMATODA

states that he had been suffering from ' fever ' for the last two
months, at first intermittent in character, but for the last seven
days more or less continued. He is much emaciated and reduced
in strength. Complains of pain on pressure over the liver and
spleen ; the latter can be felt much enlarged, reaching down-
wards to nearly the level of the umbilicus ; the lower border of
the liver, however, can only just be felt below the ribs. Tem-
perature on evening of admission 101° F. Conjunctive are
anseniic, but not jaundiced. Has also a little bronchitis. The
fever continued with slight remissions for ten days (January 4th,
1876), the highest diurnal temperature (in the afternoon) vary-
ing from 103° to 104° F.; it then abated, but dysentery set in.
He began to pass six or eight stools in the twenty-four hours,
attended with much griping, and containing varying quantities
of blood-tinged, gelatinous mucus. These became more frequent,
in spite of treatment, during the next three days, and on the
8th of January he was manifestly sinking ; passed his evacua-
tions into the bedclothes, became cold and collapsed, and died
in this state that same evening.

" A post-mortem examination was made on the following
morning, thirteen hours after death. All the organs of the
body were found more or less anaemic, but exhibited nothing
remarkable with the following exceptions. The lungs towards
their posterior margins and bases were dark, but still spongy
and crepitant. The spleen was found greatly enlarged, heavy ;
capsule tense and stretched ; substance soft, reddish brown,
irregularly pigmented ; weight 1 lb. 13 oz. The liver was of
about normal size ; its surfaces smooth, the capsule slightly
hazy looking. Hepatic substance firm, but abnormally dark,
and the bile-ducts particularly prominent and thickened.
Numbers of small distomata escaped from the incisions made
into the organ, and could be seen protruding from the dilated
biliary canals. The gall-bladder was filled with thick greenish -
yellow bile, measuring about an ounce and a half, but contain-
ing no parasites, and no ova even could be detected on micro-
scopical examination of this bile and of scrapings from the
lining membrane of the gall-bladder. The cystic duct was
free from obstruction. The condition of the common choledic
duct could not so well be ascertained, as the liver had been
removed from the abdominal cavity before anything extra-
ordinary had been detected in its condition, but, so far as it
could be examined, it was found patent j the duodenal mucous

PARASITES OF MAN

membrane was well bile-stained, and there was evidence of
biliary colouring matter in the faecal contents of the bowels.
On carefully dissecting out, and then laying open, the biliary
ducts in a portion of the right lobe of the liver (the rest being
preserved entire), numbers of distomata were found within them,
lying singly, flattened, and generally with the anterior extremity,
or " oral sucker," directed towards the periphery of the organ,
the posterior extremity towards its centre ; or in twos, threes,
or even little groups of fours, variously coiled upon themselves
or upon each other. The lining membrane of the biliary canals
was found abnormally vascular, its epithelial contents abundant
(catarrh ?), and, among these, ova could be detected under the
microscope. Sections of the liver, hardened and then examined
in glycerine, showed fatty infiltration of the lobular structure,
but not to any advanced degree ; the bile ducts considerably
dilated, their walls thick and hypertrophied, but nothing else
abnormal, or in any way remarkable. The weight of the liver
was 3 lbs. In the transverse and descending colon numerous
indolent -looking, shallow, pigmented ulcers were found, and in
the rectum others evidently more recent and highly injected.
The submucous tissues throughout were abnormally thickened.
The intestinal contents consisted of only about three ounces of
thin yellowish (bilious) faacal fluid, with small bits of opaque
mucus. This was carefully washed and examined, but no flukes
were discovered. About a dozen distomata escaped from the
liver on making the primary incisions, and quite twice this
number was found subsequently within the biliary canals. Only
a portion of the right lobe has, as I have said, been dissected,
so that it may be confidently stated that probably not less than
a hundred of these flukes must have infested this liver. All
were found dead, but it must be remembered that the autopsy
was performed thirteen hours after the death of the patient.
It is remarkable that in this case, as in the one before described
by me, no distomata were found in the gall-bladder. The
presence of these parasites in the bile-ducts seems to have led to
catarrhal inflammation of their lining membrane and abnormal
thickening and dilatation of their walls, but there is no evidence
of their having caused sufficient obstruction to produce cholasmin,
as in the case just referred to, and no marked pathological
change could be detected in the lobular structure of the liver."

After referring to the anatomical descriptions of the worm,
as recorded by myself (in ' Entozoa ') and by Lewis (in the

TREMATODA

memoir quoted below), Professor McConnell further observes
that the addition of a few more particulars seems necessary for
the determination of the identity of the species. He then gives
the following characters :

" Body lanceolate, anterior and posterior extremities pointed,
the latter obtusely. Surface covered with minute spines or
hairs. Average length §" (three eighths of an inch) ; average
breadth T'5". ' Ventral ' sucker slightly smaller than ' oral/
Eeproductive papilla or genital orifice placed a little above and
to one side of the former. Alimentary canal double and
unbranched. Uterine folds and ovary placed in the median
line, and above the male generative organs, the latter consist-
ing of two very distinct globular bodies or testes. Ova of the
usual type, i. e. oval in outline, having a double contour, and
granular contents ; average length ; average breadth ygg/'.
The only point of note is that the average length of these
flukes is greater than that of the same species found by the
authors above referred to. The D. conjunction in the American
fox, and in the pariah dog, has an average length of \"i only
two or three specimens of this size were found in this liver, and
these showed evidences of immaturity ; a few were also found
\" in length; but the great majority exactly §". The ana-
tomical characters are otherwise precisely identical."

Professor McConnell concludes his communication by a remark
in reference to the common source of infection shared by
mankind and dogs in India. The occurrence, however, of this
entozoon in an American red fox points to a very wide geo-
graphical distribution of the species. It is hardly likely that
the fox, though dying in the London Zoological Society's
Menagerie, should have contracted the parasite in England. In
the second half of this work I shall reproduce my original drawing
(fig. 56) from the ' Linnean Transactions / but I may refer to
my Manual (quoted below) for a reproduction of McConnell' s
figure. In my original specimens the integumentary spines had
fallen, probably as a result of post-mortem decomposition.

Bibliography (No. 7).—Cobbold, T. S., " Synopsis of the
Distomidse" (1. c), 1859; and in "Further Observations on
Entozoa, with experiments," 'Linn. Trans./ vol. xxiii (tab.
33, p. 349), 1860. — Idem, "List of Entozoa, including Pen-
tastomes, from animals dying at the Zool. Soc. Menagerie
between the years 1857-60," ' Proceed. Zool. Soc./ 1861.— Idem,
'Entozoa/ p. 20, pi. ii, 1864; and in "Manual of the Internal

PARASITES OF MAN

Parasites of our Domesticated Animals/' p. 81, 1873.— Lewis,
T. It,., and Cunningham, D. D., in a footnote to their ' Micro-
scopical and Physiological Researches/ Appendix C, 'Eighth
Ann. Rep. of the San. Comm. with the Govt, of India,' p. 168,
Calcutta, 1872.— McDonnell, J. F. P., "On the Distoma con-
junctum," in the ' Lancet' for 1875-76, quoted above ; reprinted
in the ' Veterinarian/ 1876 ; also (a second case) in the
' Lancet ' for March 30th, 1878, p. 476.

Distoma heterojphyes, Yon Siebold. — This minute parasite,
measuring only f of a line in length, was discovered by Dr
Bilharz, of Cairo, in the intestines of a lad, post-mortem, in
the year 1851. A second similar instance occurred, when
several hundred examples were collected and afterwards dis-
tributed amongst the helminthologists of Europe. Through
the kindness of Leuckart two of the worms eventually reached
myself. From one of these the accompanying figure was drawn.
For the purpose of supplying a full diagnosis I have else-
where described this worm as presenting an oblong, pyriform
outline, attenuated in front, and obtusely rounded behind ;
body compressed throughout, the surface being armed with
numerous minute spines, which are particularly conspicuous
(under the microscope) towards the head ; oral and ventral
suckers largely developed, the latter being near the centre of the
body, and about twice the diameter of the former ; pharyngeal
bulb distinct and separate from the oral sucker, and continued
into a long oesophagus, which divides immediately above the
ventral acetabulum ; intestinal tubes simple, gradually widening
below and terminating near the posterior end of the body ; repro-
ductive orifices inconspicuous, but evidently placed below and a
little to the right of the ventral sucker, at which point they are
surrounded by a special accessory organ, resembling a super-
numerary sucker; uterine folds numerous and communicating
with small but conspicuously developed vitelligene glands ;
testes spherical and placed on the same level in the lower part
of the body ; ovary distinct ; aquiferous system terminating
inferiorly in a large oval contractile vesicle, the latter opening
externally by a central foramen caudale.

Apart from its minuteness, moreover, this trematode is
especially characterised by the possession of a very remarkable
apparatus surrounding the reproductive orifices. It consists of
an irregularly circular disk, measuring in diameter, and
having a thick-lipped margin, which supports seventy fish-

TEEMATODA

Fig. 4— The small Egyptian
fluke (Distoma heterophyes),
viewed from behind. The
large ventral sucker, supple-
mentary disk, uterus, testes,
simple divided intestine,
vitellarium, aud pulsatile
vesicle are conspicuous.
Original.

basket-like horny ribs comparable to the claw-formations seen
in the genus Octobothrium. According to Bilkarz these ribs
give off five little branches from their sides,
but Leuckart could not see them in his
specimens. Leuckart estimated the length
of these horny filaments to be jig", whilst
their breadth was . On the whole we
may regard this organ as a complicated
form of "holdfast" designed to facilitate
or give efficiency to the sexual act. I may
here also state that this structure is by no
means unique ; for, if I mistake not, it
exists in an equally developed degree in
the young trematode which Dr Leared
found infesting the heart of a turtle.
Leared believed that he had found an ordi-
nary distome ; an opinion to which I could
not give my assent, seeing that the organ
described by him as a "folded, ventral
sucker " presented a very different aspect
to the oral sucker displayed by the same
animal. Without doubt, however, the organ in his so-called
Distoma constrictum is analogous to the supplementary " hold-
fast " existing in Distoma heterophyes. The views which I
originally advanced as to the source and condition of the parasite
are probably correct.

As regards the structure of Distoma heteropJiyes, I have only
to add that a special set of glandular organs is situated on
either side of the elongated oesophagus, but the connection
between these organs and the digestive apparatus has not been
clearly made out. Leuckart compares them to the so-called
salivary glands found in Distoma lanceolatum, and says, " The
presence of such a glandular apparatus is also indicated by the
more ventral position of the oral sucker, and the development
of the cephalic margin." The conspicuous contractile vesicle
terminating the excretory system is developed to an unusually
large extent, exhibiting in its interior multitudes of the well-
known active molecular particles. Lastly, I have only to add
that the eggs of Distoma hetcrophyes measure — ' in length by
-555" transversely.

Bibliography (No. 8). — Bilharz, " Beitrag zur Helminth,
humana," ' Zeitsch. fur wissenschaftl. Zool./ s. 62, 1851.

PARASITES OF MAN

Cobbold, 'Entozoa/ p. 195, 1864. — Kiichenmeister, F., ' Para-
siten/ 1855, s. 210, Eng. edit., p. 276, 1857.— Beared, "Descrip-
tion of Bistoma constrictum," ' Quarterly Journal of Micros.
Science/ new series, vol. ii, 1862. — Leuchart, B., 1. c, s. 613,
1863. — Moquin-Tandon, on the Genus Fasciola, 1. c., 1861. —
Weinland, on Bicrocmlium, 1. c, p. 86, 1858.

Distoma ophthalmobium, Diesing. — There is every reason to
believe that the small flukes found by Gescheid
and Von Amnion in the human eye were sexually
immature worms, but since it cannot be decided
as to what adult species they are referable I pre-
fer to notice them under the usual title. Possibly
these eye-worms may be referred to D. lanceo-
latum, as suggested by Leuckart. However

Via 6.— The eye fluke , , -T n • ,

(Distoma opktkaimo- that may be, 1 deem it unnecessary to repeat the
suckers and intestinal details recorded in the treatises quoted below.
Auimon. The largest examples measured only half a line

or about one millimetre in length.
Bibliography (No. 9). — Gobbold, e Entozoa/ p. 191. — Ges-
cheid (D. oculi humani), in Von Amnion's ( Zeitsch. f. Ophth./ iii,
and also in Amnion's ' Klin. Darstell. d. Krankheit d. Menschl.
Auges./ vols, i and iii. — Kiichenmeister, Eng. edit., p. 287. —
Leuchart, 1. c, s. 610. — Nordmann (Monostoma lentis), " Mikr.
Beitr.," Heft, ii, 'Vorwort/ s. ix, 1832.

Tetrastoma renale, Chiaje ; Hexathyridium pinguicola, Treut-
ler ; and H. venarum, Treutler. — Whether these forms are good
species or not, the fact that they were genuine parasites cannot,
I think, be disputed. The first-mentioned measured five lines
in length, and was found by Lucarelli in the urine. The
second, eight lines long, was found by Treutler in a small
tumour connected with the ovary. The third, measuring three
lines in length, was twice found in venous blood, and twice in
the sputum of patients suffering from hemoptysis.

Bibliography (No. 10). — Delle-Chiaje, 'Elmintografia Umana,'
1833.— Bremser (1. c, Bibl. No. 2), s. 265, 1819.— Cobbold,
' Entozoa ' (p. 204, et seq.). — Bujardin (1. c, Bibl. No. 2),
s. 265, 1819. — Treutler, ' Obs. Path. Anat. ad Helm. Corp.
Humani/ p. 19, 1793. — Zeder, ' Anleitung zur Naturg. der Ein-
geweidewiirmer/ s. 230, 1803.

Amphistoma hominis, Lewis, and McConnell. — The original
account of this species is based upon two finds. The first
series of specimens was procured from Dr J. O'Brien, of

TREMATODA

Gowhatty, and the second set from the Pathological Museum of
the Calcutta Medical College. Dr O'Brien and Dr Curran
together procured their specimens, post-mortem, from an Assa-
mese. There were hundreds of worms present in the vicinity of
the ileo-colic valve. The museum specimens were procured from
a patient who died at the Tirhoot gaol hospital in 1857. They
were (say the authors) presented to the museum by Dr Simp-
son, and in the catalogue their history was briefly recorded as
follows :

Fig 6 — The human amphistome (Jmphistoma hominis). Longitudinal section, a, Oral sucker ;
b, pharyngeal bulb; e, nerve ganglia; d, oesophagus; e, genital pore; f, vagina; g, ductus
ejaculatorius; h, ventral nerve cords; i, intestinal canal;,;, upper testis; k, water vessel;
I lower testis (ovary according to Lewis); m, principal ducts of the vitellarium ; n, branches
of the vitellary ducts ; o, ventral pouch or bursa ; p, caudal sucker. Magnified 12 diameters.
After Lewis.

" The caecum of a native prisoner who died from cholera in
the Tirhoot gaol hospital, with a number of peculiar and,
probably, hitherto unrecognised parasites, found alive in that
part of the intestinal canal." {Presented by Dr Simpson through
Professor E. Ooodeve.)

In continuation of their narrative, Drs Lewis and McConnell
go on to say that, " with reference to this preparation, the follow-
ing very interesting particulars from the ' Annual Jail Report of
Tirhoot ' for 1857 have been very kindly placed at our disposal by
the Surgeon-General, Indian Medical Department. The prisoner,

PARASITES OF MAN

Singhesur Doradh, aged 30, was attacked with cholera on the
13th, and died on the 14th of July, 1857. Had not been in
hospital previously, and was employed in cleaning the jail."

The post-mortem examination was made three hours after
death : — " Colon externally livid, contracted ; contains a little
serous fluid with flakes of mucus. Mucous membrane healthy
except venous injection. In the ceecum and ascending colon
numerous parasites like tadpoles, alive, adhering to the mucous
membrane by their mouths. The mucous membrane marked
with numerous red spots like leech-bites from these parasites.
The parasites found only in the caecum and ascending colon,
none in the small intestines." This description is by Dr Simp-
son, who adds, " I have never seen such parasites, and apparently
they are unknown to the natives. They are of a red colour,
size of a tadpole, some young, others apparently full grown,
alive, adhering to mucous membrane, — head round, with circular
open mouth, which they had the power of dilating and contract-
ing. Body short and tapering to a blunt point."

Drs Lewis and McConnelPs description of the worm is too
long to be quoted in full. The parasites measure ~" to in
length, by \" to 5" in breadth. Science is much indebted to
these eminent observers for having unearthed the museum
specimens and for recording the facts they could gather.
From a zoological point of view the most interesting fact con-
nected with Lewis's amphistome is the existence of a gastric
pouch. This structure brings these human Masuri into close
relation with the equine parasite which I have named Gastro-
discus Sonsinoii, and which will be found illustrated in this
work (fig. 62). In short, Lewis's worm appears like a transi-
tion form ; the absence of gastric supplementary suckerlets
separating it from the new generic type.

Bibliography (No. 11). — Lewis, T. B.} and McOonnell, T. F. P.,
" Amph. hominis ; a new parasite affecting Man," f Proceedings
of the Asiatic Society of Bengal/ Aug., 1876.

Bilharzia hamatobia, Cobbold. — This remarkable parasite
was discovered by Bilharz in 1851. It was subsequently
found by myself in an ape (1857) other species of the same
genus having since been detected by Sonsino in the ox and
sheep (1876). The human examples were originally obtained
from the portal system of blood-vessels. Afterwards they were
obtained by Bilharz, Griesinger, and others, from the veins of
the mesentery and bladder. It was shown that they were not

TJIEMATOPA

only associated with, but actually gave rise to a formidable and
very common disease in Egypt.

In 1864 Dr John Harley made the interesting announcement
that he had discovered specimens of this singular genus in a
patient from the Cape of Good Hope. He also showed that
the entozoon was the cause of the hematuria known to be
endemic at the Cape. Harley believed his parasites to represent
a new species (Distoma capense), but in this view I showed that
he was mistaken. His admirable contribution, nevertheless,
served not only to establish the wide range of this parasite on
the African continent, but also to throw much light upon the
subject of endemic helminthiasis. As this worm forms an
almost altogether exceptional type of fluke- structure, it became
necessary to supersede the original no-
menclature proposed by Bilharz and
Von Siebold (Distoma haematobium).
Accordingly I proposed the term Bil-
harzia, whilst other helminth ologists
subsequently proposed various titles
(Gyncecophorus, Diesing ; Schistosoma,
Weinland; Thecosoma, Moquin-Tandon) .
On various grounds, and chiefly on
account of priority, most writers have
at length definitely accepted the nomen-
clature which employed the disco verer's
name for generic recognition.

The Bilharzia hamatobia may be
described as a trematode helminth
in which the male and female repro-
ductive organs occur in separate indi-
viduals; the male being a cylindrical
vermiform worm, measuring only half an inch or rather more
in length, whilst the female is filiform, longer, and much
narrower than the male, being about four fifths of an inch from
head to tail ; in both, the oral and ventral suckers are placed
near each other at the front of the body; in the male the
suckers measuring 7ol5", in the female in diameter; in either,
the reproductive orifice occurs immediately below the ventral
acetabulum. The comparatively short, thick, and flattened
body of the male is tuberculated and furnished with a gyrueco-
phoric canal, extending from a point a little below the ventral
sucker to the extremity of the tail j this slit-like cavity being

Fig. 7 — The blood fluke (Bilhartia
htematobia). The lower end of the
female is withdrawn from the gynse-
cophoric canal of the male. After
Kuchenrneister.

PARASITES OP MAN

formed by the narrowing and bending inwards of the lateral
borders of the animal, the right side being more or less com-
pletely overlapped by the left margin of the body ; caudal
extremity pointed j intestine in the form of two simple blind
canals. Female with a cylindrical body measuring only ~ of
an inch in thickness in front of the oral sucker ; lodged in the
gyneecophoric canal of the male during the copulatory act ;
thickness of the body below the ventral acetabulum being about
and at the lower part — surface almost smooth through-
out ; intestinal canals reunited after a short separation to form
a broad, central, spirally twisted tube extending down the
middle of the body ; vitelligene and germigene canals combining
to form a simple oviducal canal, which is continued into a
simple uterine tube, finally opening near the lower margin of
the ventral sucker; eggs pointed at one end, or furnished with
a projecting spine near the hinder pole.

The study of the structure and formation of the contents of
the ova possesses great interest. When fully developed the
eggs are oval, measuring from ~" to in length, with an
average transverse diameter of 3^". Some are a trifle larger,
others smaller. Occasionally one encounters narrow specimens,
and also aberrant forms presenting a pear-shaped outline. I
have met with eggs not exceeding — ' in their long diameter,
and gig" transversely, whose yolk-contents had already arrived at
an advanced stage of segmentation.

The shell is transparent, of a brown colour, and free from any

markings, lines, or sculptur-
ing. One pole of the shell is
invariably narrower than the
other, and usually presents a
more or less pointed extremity
(fig. 8). This narrow end
commonly displays a sharp,
projecting, beak-like spine,
which, at its base, constantly
rests upon the centre of the
pole of the shell, but occa-
sionally it is eccentrically

Fig. 8.— Two eggs of Bilharzia. a. With the yolk " "

coarsely segmented ; b, with the yolk granulated and placed, (tig. O a) . In SOme I6W
the spine wanting. Original. , ,

examples the spine is re-
moved to a little distance from the actual extremity of the
shell ; but even in these instances, so far as my observations go,

TREMATODA

its apex always projects beyond the level of the curved end
of the pole. Now and then the spine is altogether absent (fig.
8fc); and when present it is, as already hinted, very
unequally developed. In size the spine ranges from a mere
point, having an extreme length of only up to the

comparatively large magnitude of lengthways.

According to the best evidence there is no good ground for
asserting the existence of any specific differentiation between
the parasites coming from the Cape and Egypt respectively.

Taking a more extended view of the significance of these
singular chorional spines, I think we may here recognise the
early efforts of Nature, so
to speak, to form or evolve
a special organ, which, in
the eggs of certain other
parasites, becomes capable
of attaining a relatively
prodigious degree of deve-
lopment. To me it seems
that the little process in
question is a kind of rudi-
mentary holdfast ; and, as
such, it may be reckoned
as the homologue of a
variety of egg-appendages.
Eleven years ago Mr Ed-
win Canton discovered some curious ova attached to the
conjunctiva of a turtle's eye. I had no hesitation in pronounc-
ing them to be referable to some ectozoon or entozoon
belonging to one or other of the allied genera Polystoma,
Tristnma, Octobothrium, and Dactylogyrus. Now, whilst the
Bilharzia ova display only a solitary and imperfectly deve-
loped holdfast, placed at one end of the shell, the singular
eggs described by Mr Canton develop organs of anchorage at
both extremities. Parasitic ova exhibiting analogous pro-
cesses, spines, and filamentary appendages at both poles,
have been observed in various species of parasite — as, for
example, in Monostoma verrucosum infesting the fox, in Tania
cyathiformis infesting the swallow, in Taenia variabilis of the
gambet, in Octobothrium lanceolatum attached to the gills of
the common herring; and in Polystoma ajpjpendiculata, from
the branchiae of various marine fishes. Eggs of parasites which,

Fig. 9. — Two eggs of Bilharzia., with eccentrically placed
spines. That to the left shows mulberry cleavage of
the yolk ; the other having lost its embryonal contents
by rupture. Original.

PARASITES OF MAN

like Bilharzia, are furnished with a single appendage, may like-
wise be seen in the ova of different species of Dactylogyrus in-
festing the gills of the pike. In the more strongly pronounced
developments it is easy to perceive how admirably these out-
growths are adapted to the necessities of the different species of
parasite to which they are severally referable ; and, even in the
case of Bilharzia, the trifling amount of anchorage furnished
by a projecting point is not absolutely thrown away. The
resistance will also be greater where the spine is situated a. little
on one side of the pole of the egg, which seems to need steady-
ing during the violent struggles of the embryo to escape from
its temporary abode.

Fig. 10.— Free ciliated embryo Fig. 11.— Two ciliated embrvos of Bilharzia; showing sar-
of Bilharzia, with pear-shaped code spherules in their interior. That to the left has

rudimentary organs below the recently escaped the shell. Original,

head. Original.

When any number of ova are removed from the urine and
examined, it will be found that a large proportion of them contain
embryos in an advanced stage of larval growth. The struc-
tural appearances presented by the embryos whilst still in the
eggs are remarkably uniform ; since, in all, the yolk appears to
have resolved itself into a mass of rounded sarcode-globules, one
or two of these particles being conspicuously larger than the rest
(fig. 12). At this stage, except towards the cephalic division of
the larva, no tendency to differentiation is perceptible; but

TEEMATODA

4<S

some time after the embryo lias escaped, one may notice
elongated masses of sarcode formed by the coalescence of the
globules. Whilst still in the egg, one end of the primitive
embryonal mass becomes gradually narrowed, cilia at the same
time appearing. This part becomes the future head, eventually
acquiring the form of a cowl. Whatever form the body of the
embryo may display after extrusion from the shell, the head
retains its conical shape, the cone itself being narrowed
or widened only when the larva is subjected to abnormal
conditions (fig. 14). Whilst the head is undergoing develop-
ment within the shell, one, two, or sometimes three, pyri-
form masses make their appearance within the cone ; and
after the embryo has escaped, these structures become more
marked (fig. 10). The sarcode-globules refract light strongly;
and, when the larva is not compressed in any way, they move
freely within the somatic cavity. In well-
developed embryos, whilst still in the egg,
the cilia are observed to clothe every part
of the larva except the oral papilla. This
minute nipple-like projection measures
about the ~m of an inch transversely, form-
ing a very simple kind of unarmed pro-
boscis. When the head of the free embryo
is viewed from above, the proboscis looks
like a central ring surrounded by a series
of regular folds, which radiate outwards
like the spokes of a wheel. The ridges
thus formed support numerous cilia, these
latter projecting at the circumferential mar-
gin of the cephalic cone in such a way as to
present the figure of a star. Dr Harley
has admirably represented this character,
which is shared by many other parasitic
larvae. Throughout the greater part of the time, whilst the
embryo is still resident within the egg, the broad neck or base
of the cephalic cone forms a fixed point of resistance by its firm
attachment to the inner wall of the shell ; and this structural
union, so long as it remains intact, enables the embryo to move
not only its head and body from side to side synchronously, but
also each part independently. When the time for final escape
is drawing near, the vigorous movements of the cone-shaped
head seem chiefly concerned in loosening the membranous

Fig. 12.— Eg<? of Bilharzia,
with contained embryo and
free sarcode globules. Ori-
ginal.

PARASITES OF MAN

connection just referred to ; and when, at length, the ciliated
animalcule has succeeded in overcoming this first difficulty, it is

ludicrous to witness its frantic efforts to find
an opening in the shell. While thus par-
tially liberated, it will rush to and fro from
one pole of the egg to the other, performing
a series of summersaults, and at the same
time occasionally rolling itself over laterally.
This activity becomes gradually more and
more violent, until at length its excitement is
worked up into a sort of frenzy. I have many
times watched these performances, which,
however, are only to be seen within those ova
whose shells, for some reason or other, refuse
to yield to the earlier aud ordinary efforts of
the prisoner. In all cases where these phe-
nomena are witnessed the eye readily detects
a number of small free globules between the
embryo and the inner wall of the shell (fig.
13). These minute particles are likewise
tossed about tumultuously during the rapid
rotatory movements of the imprisoned larva.
Except as regards their size, these globules
do not differ in character from the sarcodic
contents of the animalcule. They are pro-
Fl^VoVIXSatB?ighay bably superfluous detachments from the pri-
tSSfi&lSfi&iiS mitive yolk-mass, but it is just possible that
developed, original. ttey may affora. some aid in the final break-
ing up of the shell. Whilst the embryo remains fixed its
tail is usually directed towards the narrower or spine-bearing
pole of the egg, but in a few instances I have seen this position
reversed. As regards the precise mode of emerging from the
shell, and the time occupied by the larva in freeing itself, there
are several points of interest. Speaking generally, the purer
the medium into which the ova are transferred, the more rapid
will be the movements of the larvae. To give an example of
observed facts in relation to the rapidity of development, I cite
the following : — " On the 20th of August, 1870, I placed twelve
eggs of Bilharzia under the microscope. The medium in which
they were immersed consisted of eight parts of ordinary
drinking water to one of urine. At the expiration of seventeen
minutes the first-born made its escape. In the course of

TREMATODA

another minute two more emerged. In twenty-six minutes the
fourth, in twenty-eight the fifth, in thirty-two the sixth, in
thirty-four the seventh, in thirty-seven the eighth, in thirty-eight
the ninth, in forty the tenth, in forty-three the eleventh, and
in forty-six minutes the twelfth, respectively made their
appearance/''

Now, this rapid mode of birth and emergence from the shell
is very much more striking in the case of eggs which are placed
in perfectly pure water ; for, whilst the eggs are still in the
urine, there appears to be neither the power nor the inclination
on the part of the embryo to escape ; but, on isolating and
placing them in suitable conditions, their behaviour is even more
remarkable. In a space of less than two minutes I have
repeatedly seen the hitherto motionless embryo alter its shape
by contractions, become violently agitated, and burst out of its
shell in the condition of a free-
swimming animalcule. More-
over, it is worthy of remark
that the eggs and larvae of Bil-
harzia soon perish in stale urine.
" On the 16th of August, 1870,
I placed about a thousand eggs
in a quart of fountain- water, to
which only a drachm or rather
less of urine had been added.
At the expiration of forty-eight
hours not a single living em-
bryo could be found. I subse-
quently ascertained that I could
not keep the embryos alive for
twenty-four hours in any water
in which I had introduced the
smallest trace of mucus, blood-
corpuscles, urinary crystals, or
decomposing matters of any kind.
All sorts of reagents speedily
killed the larvae. Mere discolora-
tion by carmine solution, or by
the addition of a drop of the solu-
tion of permanganate of potash,
instantly caused them to assume grotesque and unnatural shapes
(figs. 13 and 14), death sooner or later following as a result

Fio H.— Ciliated embryos of Bilharria, deformed
by the application of reagents. Original.

PARASITES OP WAN

of the disintegration and resolution of their delicate bodies into
mere sarcode-masses. Still more rapidly poisonous effects
were produced by the addition of a little sherry or alcohol.
In solutions where the amount of spirit did not exceed one part
of spirit, proof strength, to fifty parts of water the effect was
the same."

The development of the larva is equally well accomplished in
distilled water, in well-water, and in brackish water: In pure
sea-water the process goes on less satisfactorily. It was found,
indeed, that the addition of slightly saline water to ciliated
embryos, which were on the point of expiring in fresh water,
had the effect of reviving them for a time. These facts have
an important practical bearing.

I have thus shown that the escape of the embryo is by no
means the slow process that Bilharz has described. Almost
invariably the shell bursts by a longitudinal slit extending over
fully two thirds of its long diameter, the first point of rupture
being commonly situated midway between the spine and the
centre of the shell. In normal births, so to speak, the head of
the animalcule emerges first ; but occasionally the animal
escapes sideways, and I have even seen the embryo extricate
itself tail foremost. Not unfrequently it has a difficulty in
detaching itself from the shell, in which case the egg is whirled
round and round by the half -freed prisoner (fig. 15). The
lodgment of the spine, however, against any foreign substance
affords the necessary leverage for ensuring escape.

The larva never displays its proper elongated, spindle-shaped,
or cylindro- conical figure, until some short time after its escape
from the shell ; and, as a consequence of this, its powers of
locomotion are less marked at first than they are subsequently.
At the time of extrusion the larvae are commonly more or less
hour-glass shaped (fig. 11) ; this particular form being some-
times retained for many minutes or even for an hour. Usually
the larvee have a tendency to acquire their normal shape imme-
diately after quitting the shell; the oval, pear-shaped, and
variously contracted forms gradually merging into the charac-
teristic cone-shaped animalcule (fig. 10). In their fully devel-
oped condition, they exhibit the most lively movements ; and to
witness several hundreds of them rushing about with unceasing
activity is a curious sight. The phenomenon, moreover, loses
none of its interest from the consideration that only a few hours,
or it may have been minutes, previously, these now actively

T1JEMAT0DA

gyrating animalcules were lodged in ovo within the blood-vessels
of their human host. From persons who are infested, myriads
of these eggs of Bilharzia daily make their escape during the
act of micturition ; and, when this act is accomplished by the
host out-of-doors, it is easy to perceive how readily the ova may
be subjected to conditions favorable to the development of larvee.
The direct passage of the urine into any considerable receptacle
of natural or fresh water would in a few minutes ensure the
hatching of all the eggs ; and in the absence of any such direct
aid to development, the accidental occurrence of a shower of
rain would, in all localities where the Bilharzia disease is
endemic, readily transfer the ova into ditches, ponds, rivers,
lakes, and ultimately, perhaps, even into the sea itself.

Fig. 15.— Egg of Bilharzia, with the shell still adhering Fig 16 —Ciliated embryo of Bilharzia in
to the escaped ciliated embryo. Original. the act of dying from the escape of its

sarcodic contents. Original.

The behaviour of the embryo under the action of reagents of
various kinds is remarkable. Thus, when on the 5th of Sept.,
1870, I placed some ova in brackish water, of the strength of
two parts of fresh water to one of pure sea- water, their contents
were readily developed, though the escaping embryos did not
swim vigorously. When again I placed some other eggs in
pure sea-water, their contained embryos became instantly trans-
fixed, the vibratile cilia of the head being rigid and motionless.
At first I naturally concluded that the embryos were killed
outright j but, to my great surprise, the shock passed away in
about half an hour, when they revived and were soon afterwards
hatched. One of the larvae thus set free carried off several of
the loose intra-chorional globules which had, during the period
of transfixion, become firmly adherent to the ends of the caudal
cilia. Here I may remark upon a decided difference observable

PARASITES OF MAN

between the cilia of the head and body respectively. The former
are at all times vibratile, active, and conspicuous, whilst the
latter are more delicate, capable of comparatively little motion,
and partaking more of the character of fine setae. In length
their general measurement varies from to '. The action
of pure sea-water on the free animalcules, previously immersed
in fresh or brackish water, was equally striking. All, without
exception, immediately became paralysed and almost motionless ;
nevertheless, on again adding fresh water, several entirely reco-
vered. It is worthy of notice that in these cases the cephalic
cilia furnished the first indications of returning viability. I was
particularly struck with the behaviour of one embryo, which,
under the stimulus of- the sudden shock, retracted its cone-
shaped head almost entirely within the general cavity of the body
(fig. 14, lower specimen) . In their moribund condition, whatever
shape the embryo retained, the sarcodic contents gradually faded
away ; the outline of the creature, however, becoming more
marked (fig. 16). Usually the body of the animalcule became
elongated whilst expiring in sea- water. Under other circum-
stances the embryo frequently bursts ; the sarcodic contents
escaping in the form of amceba-like bodies and the cilia retain-
ing their powers of movement long after all traces of the
sarcode have disappeared.

The larvse of Bilharzia closely resemble those
of Fasciola hepatica, which latter may be appro-
priately noticed in this place. The ciliated em-
bryo of the common liver fluke has the form of
a long cone inverted ; the anterior end or head
being flatly convex. In the centre is a short
proboscis- like papilla destitute of cilia (fig. 17).
The general covering of cilia rests on a well-
defined granular epidermis ; this latter being
succeeded by a dense peripheral layer of large
nucleated cells, each of them measuring about
slm" in diameter. The epidermis measures g^g"
in thickness. In the central mass of paren-
chyma no internal organs are recognisable, but
Leuckart observed indications of a canal which
he thought might open at the tail, though the
opening itself was not actually visible.
As long as the ciliated covering remains intact the embryo,
like other animalcules, displays great activity, whirling round

Fig. 17.— Cilioted em-
bryo of Fasciola he-
patica, Bhowing the
so-called eye-spot.
After Leuckart.

TREMATODA

and round on its own axis, and also describing gyrations and
circles of different degrees of range in the water, the latter
movements being accomplished by bending the body upon itself
to a greater or lesser curvature. The embryos of Bilharzia and
other infusoria exhibit the same behaviour, and, as Leuckart
observes, when these embryos knock against any obstruction,
they pause after the blow, as if to consider the nature of the
substance they have touched. As in the case of fluke embryos
generally, the ciliated covering eventually falls off and the
embryo reassumes a more or less oval figure, at the same time
changing its swimming mode of progression for the less digni-
fied method of creeping. In the free ciliated condition the
embryo of the common liver-fluke measures, according to
Leuckart, j^" in length, the anterior broad end being — ' . The
cilia have a longitudinal measurement of y^g".

According to the observations of Dr Willemoes-Suhm, the
cilia of the embryos of the Distoma megastoma are limited to
the anterior pole of the body. This is also the arrangement, as
Leuckart first pointed out, in Distoma lanceolatum (fig. 18).
On the other hand, Pagenstecher has shown that the embryos
of Distoma cygnoicles and Amphistoma (Dijplodiscus) subclavatum
are ciliated all over, an observation which, as regards the latter
species, has been confirmed by Wagener and others. Dr
Pagenstecher* s original statement to the effect that "intracho-
rional germs of trematodes offer no distinctive characters,"
must, therefore, in the present state of our knowledge, be
accepted as a general conclusion admitting of many exceptions.
In the early stages of development the embryo of Distoma
lanceolatum occupies the centre of the egg, and according to
Leuckart has its conical head invariably directed towards the
upper pole of the shell, or, in other words, to that end of the
egg which is furnished with a lid-like operculum. Leuckart
describes the embryo itself as " finely granular and armed at
the tip with a dagger-like spine, which, with the simultaneous
displacement of the adjacent granular mass, can be pushed
forward and drawn back again." Besides this so-called cephalic
granular mass, there are within the embryonic body two other
granular masses widely separated from each other, but occupy-
ing the posterior half of the embryo. These Leuckart supposes
to be the rudiments of a future brood, to be developed at the
time when the free embryo shall have lost its ciliated swimming
apparatus, shall have bored its way by means of the cephalic

PARASITES OF MAN

spine into the tissues of a mollusk, and shall have become
metamorphosed into a sac-like larva (Nurse, Sporocyst, or Redia,
as the case may be). Whatever be the full significance of these
internal developments, we have at least satisfactory evidence
that the complete and free embryo is a globe-
shaped animalcule, having the anterior third
or cephalic end of the body covered with cilia,
and armed with a central boring spine. In
consequence of this limitation of the ciliated
covering, its swimming movements are less
fig. i8.-cniated em- vivacious than those of the embryo of Fasciola
Siatl^Sriiut- hepatica ; it will, therefore, probably take up
kart- its residence in a less active host than that

chosen by the embryo of Fasciola, selecting one of those mollusks
which either move slowly or are prone to keep at the bottom
of the water. The mature eggs have a length of B^ to ^ of an
inch, and a breadth of ggg.". The long diameter of the free
embryo varies from ~" to ggg", the transverse diameter being
tsW "• Whilst the embryos were still in the egg Leuckart could
see no ciliary, motion. With most observers, both the ciliary
apparatus and the boring spine appear at this stage to have
altogether escaped observation.

As regards the intimate structure of the ciliated embryo of
Bilharzia hcematohia, I have further to observe that, shortly
after its extrusion from the shell, the hitherto loose, globular
sarcode particles coalesce. This is apparently a preliminary
step towards the subsequent differentiation process. Respect-
ing the pedunculated blind sacs formed within the head, 1
think that we must regard the largest one as representing the
stomach of the larva in its future cercarian stage. Under
the objective I distinctly recognised, in the cavity of the
central blind sac, numerous highly refracting granules, the
diameter of which averaged not more than j^mo" '. The rudi-
mentary stomach is often traceable whilst the larva is still within
the egg. It measures about in length, including the
peduncle, and ' in breadth. The width of the narrow stalk
does not exceed ^o5". The other two-stalked bodies appeared
to have the character of lemnisci. They were occasionally well
seen whilst the embryo was still within the egg. As regards
the integument, it is easy to recognise two layers. In careful
adjustments of the focus the inner wall of the transparent
dermis presents a beaded appearance. These minute and

TREMATODA

regular markings do not undergo alteration during the con-
tractions of the body of the larva.

A highly developed water-vascular system exists in these
little animalcules. On many occasions I saw traces of this set
of vessels, and in several instances I obtained a most satisfactory
view of the entire series of branches.
Anxious to receive confirmation of my
discovery, I demonstrated the existence
of these vessels to a skilled microscopist
—the late Mr J. G. Pilcher, of H. M.
Army. In the briefest terms it may
be said that the water-vascular system
of Bilharzia, in the larval condition,
consists of two main stems, which
pursue a tortuous passage from head to
tail, and which, in the course of their
windings, give off several anastomosing
branches (fig. 19). As also obtains in
the corresponding larvas of Diplodiscus
subclavatus, there is no excretorv outlet
visible at the tail.

Encouraged by the experiences and
determinations of Pagenstecher, Filippi,
Wagener, Leuckart, and others, I sought
for the intermediate hosts amongst
fresh- water mollusks and small Crus-
tacea. Failing of success in these, it
occurred to me that the larvas of Bil-
harzia might normally reside in fluviatile
or even in marine fishes. This latter
idea seems also to have struck Dr

Aitken. In an appendix tO his ' ReDOrt Fra ^.--Outline representation of
j ■ r » ,r t. , _ ^ a c^ted embryo . of Bilharzia,

to tne Army Medical Department for sll0wing the arrangement of the

1 QAQ > 1 i. J £ "vr ii r,T „ *'H.te.r vessels and the vacuoles.

l«o«, dated from Netley, Nov., 1869, 0ris"^-
he gives a figure of a nurse-form, which he terms a cercaria,
from the tail of a haddock— suggesting for Bilharzia some
genetic relation. Dr Aitken also extends his views in reference
to certain larval trematodes alleged to have been found in the so-
called Delhi boils and Lahore sores. These parasitic forms have
however, been shown by Dr Joseph Fleming to be nothing more
than altered hair-bulbs ('Army Med. Reports » 1868 69)

In regard to the flukes from the haddock, I have satisfied

PARASITES OF MAN

myself that these immature trematodes from the nerves of the
cod-tribe can have no genetic relation with Bilharzia ; and I
think it due to Dr Maddox to say that I accept his conclusion
respecting them. In his paper (f Micros. Trans./ vol. xv, 1867,
p. 87) he offers strong proof that the so-called Bistoma neuronaii
Monroii of the haddock (Morrhua ceglefinus) is the juvenile
condition of Gasterostoma gracilescens of the angler (Lophvus
piscatorius) .

I am sorry to have to state that all my experiments proved
negative. I tried to induce the ciliated embryos to enter the
bodies of a variety of animals, such as Gammari, Dipterous
larvae, Entomostraca, Lymnaai, Paludinae, different species of
Planorbis, and other mollusks; but neither in these, nor in
Sticklebacks, Roach, Gudgeon, or Carp, did they seem inclined
to take up their abode.

The very peculiar and formidable helminthiasis produced by
this parasite has been thoroughly investigated by Griesinger
and Bilharz, and it has been fully described in the standard
works of Kuchenmeister and Leuckart. My own case from
Natal also supplied many interesting clinical facts which
were published in my ' Lectures on Helminthology/ quoted
below. The comparative prevalence of this disorder in Egypt
is well established. Symptomatically, its principal feature
consists in a general disturbance of the uropoietic functions.
Diarrhoea and hasmaturia occur in advanced stages of the com-
plaint, being also frequently associated with the so-called
Egyptian chlorosis, colicky pains, anaemia, and great prostra-
tion of the vital powers. The true source of the disorder,
however, is easily overlooked unless a careful microscopic
examination be made of the urine and other evacuations. If
blood be mixed with these, and there also be a large escape of
mucus, a minute inspection of the excreta will scarcely fail to
reveal the presence of the characteristic ova of Bilharzia.
Besides the increase of mucus secretion, there may even be
an escape of purulent matter, showing that the disorder has
far advanced. The patient's constitution eventually becomes
undermined ; pneumonia often sets in, and death finally ensues.
On making post-mortem examinations the following pathological
facts come to light. In cases where the disease has not advanced
very far, minute patches of blood- extravasation present them-
selves at the mucous surface of the bladder, but in more
strongly pronounced cases the patches are larger or even

TREMATODA

confluent. In some instances there are villous or fungus-like
thickenings, ulceration and separation of portions of the
mucous membrane, with varying degrees of coloration, accord-
ing to the amount of the extravasation, which becomes con-
verted into grey, rusty-brown, or black pigment deposits. A
gritty or sandy deposit is often superimposed, consisting of
ordinary lithic-acid grains mixed with eggs and egg-shells. Eggs
are retadily detected in the urine, these having escaped from the
ruptured vesical vessels. The lining membranes of the ureters
and renal cavities are also more or less affected ; the kidneys
being frequently enlarged and congested. It must, however,
be borne in mind that in all these organs the true seat of the
disorder is the blood, which forms the proper habitat of the
Bilharzia ; and this being the case, the worms as well as their
escaped eggs may be found in any of the vessels supplying the
diseased organs. In one instance, quoted by Leuckart, Grrie-
singer found a number of empty eggs in the left ventricle of
the heart, and from this circumstance it was supposed that
they might be carried into various important organs, or
even plug up the larger vessels. As before stated, however,
the parasites are more particularly prevalent in the vessels of
the bladder, mesentery, and portal system. The effects upon
the intestinal mucous membrane are, in most respects, similar
to those occurring in the urinary organs. Blood extravasations,
with thickening, exudation, ulceration, and fungoid projections,
appear in and upon the intestinal mucous and submucous
tissues ; these appearances, of course, being more or less
strongly marked according to the degree of infection.

In regard to the treatment of the helminthiasis, I am pre-
cluded from entering into details here ; nevertheless, I am glad
to perceive that the principles which I long ago enunciated
have received approval both at home and abroad. As stated
in my ' Lectures ' our object should be not to interfere with,
but to promote nature's curative efforts. If I read the patho-
logical facts correctly, she seeks to bring about this result by
erecting artificial barriers which serve to moderate the bleeding.
In this way, under vprdinary circumstances, the life of the
bearer is sustained, or held in the balance until the parasites
either perish or cease to be capable of causing active disease.
Depend upon it, this is the principle which should guide
physicians in their treatment of the Bilharzia disorder. If the
adult parasite were merely attached to the lining membrane of

PARASITES OP MAN

the bladder, then powerful diuretics and medicated injections
would probably prove serviceable ; but since the entozoa reside
in the blood we must be careful not to increase the patient's
troubles. In the case of intestinal worms the most powerful
parasiticides may be prescribed without let or hindrance ; but
that drug must be a truly subtle worm-poison which, when
taken into the system, shall kill the blood-flukes without
exerting any injurious effects upon the parasite bearer.

When, in 1872, I published my lectures on helminth ology, I
remarked that it was not improbable that, ere long, many
more cases of Bilharzia disease would be brought to light.
What has been added in this respect is chiefly due to the
researches of Sonsino, but a case of some interest has been
recorded comparatively recently by Dr W. K. Hatch, stationed
at Bombay. From the particulars furnished it seems evident
that the victim, an English gentleman, contracted the disease
by drinking water, either in Arabia or in Egypt, in which
latter country, however, he had only sojourned fifteen days.
From the patient's statements it appears that, hematuria is
frequent amongst the Arabs. Incidentally, Dr Hatch men-
tions that Dr Yandyke Carter had informed him that, so early
as the year 1862, he (Dr Carter) had detected the embryos of
Bilharzia in the urine of an African boy admitted to the
Jamsetjee Jejeebhoy Hospital. The treatment employed by
Dr Hatch was that recommended by Dr Harley in his well-
known memoir. Having myself energetically opposed Dr
Harley's views on pathological grounds, I am not surprised to
see it stated that Dr Harley's method of treatment effected " no
diminution in the number of the parasites." As I said in my
lectures (now out of print) it is evident that " nature " in view
of moderating the hematuria — by the formation of plugs at the
ulcerated points of the mucous surface — sets up the artificial
barriers above referred to ; therefore if you catheterise and
employ medicated injections you do more harm than good.
As to the administration of belladonna internally, in view of
retarding development, or of destroying the parasite, no good
can be expected from this source. I certainly obtained better
results with buchu and bearberry (Arctostaphylos).

In the matter of sanitation it is quite evident, from the
foregoing data, that the danger of infection cannot arise from
the drinking of impure water, as ordinarily understood. The
embryonal larvae would be killed by an admixture of sewage.

TREMATODA 55

It is obvious that infection can only occur from swallowing free
cercarias or freshwater mollusks which contain the higher larval
forms in their encysted or pupa condition. Slow running
streams or stagnant pools with sedgy banks are eminently
favorable to the existence and multiplication of intermediary
bearers, and consequently their waters are dangerous if
employed for drinking purposes.

Bibliography (No. 12). — Bilharz, in Siebold and Koll.,
'Zeitsch. fur wissensch. Zool./ iv, 1851. — Idem, 'Wiener
medic. Wochenschrift/ 1856.— Oobbold, T. 8., "On some
new forms of Entozoa {Bilharzia magna)," 'Linn. Trans./ vol.
xxii, p. 364, 1859. — Idem, " Synopsis of the DistomidEe," in
'Proceed. Linn. Soc./ vol. v, Zool. Div., p. 31, 1860. — Idem,
" Eemarks on Dr J. Harley's Distoma capense," in ' Lancet/ also
in the ' Veterinarian/ and in ' Intell. Observer' for Feb. and
March, 1864. — Idem, " Entozoa," 1. c, p. 197, 1864.— Idem, " On
Blood Worms," Lecture xx in ' Worms/ 1. c, p. 145 et seq.,
1872 ; Tommasi's edit,, Vermi, p. 141, 1873. — Idem, "On the
Embryos of Bilharzia" ' Brit. Assoc. Rep./ 1864. — Idem, " On
the Development of Bilharzia hcsmatobia, together with Remarks
on the Ova of another Urinary Parasite occurring in a case of
Hasmaturia from Natal," 'Brit. Med. Journ./ July, 1872;
repr. in the 'Veterinarian/ 1872. — Idem, 'New Entozootic
Malady, &c/ (brochure), London, 1865. — Idem, " Helminthes,"
in Gunther's 'Record of Zool. Literature/ p. 617, 1865. — Idem,
" Entozoa in relation to Public Health and the Sewage Question,"
Rep. of the Proceed, of the Metrop. Assoc. of Officers of
Health, iu 'Med. Times and Gazette/ Jan., 1871, repr. in the
' Veterinarian/ p. 359, 1871. — Idem, "Verification of recent
Haematozoal Discoveries in Australia and Egypt," ' Brit. Med.
Journ./ June, 1876. — Idem, " On Sewage and Parasites, espe-
cially in relation to the Dispersion and Vitality of the Germs of
Entozoa," rep. in ' Med. Times and Gaz/ for Feb., and the ' Vete-
rinarian' for May, 1871. — Davaine, 0.,l. c, ' Synops/ and p. 312,
1860. — Diesing, C M., ' Revis. d. Myzelmith/ Vienna, 1858.
— Griesinger, " Klin, und Anat. Beobachtungen uber die
Krankheiten von Egypten," in ' Arch, fur physiol. Heilkunde/
1856. — Idem, ' Gesammelte Abhandiungen/ Berlin, 1872. —
Idem, 'Arch. d. Heilk./ 1866. — Barley, J., ' On the Hematuria
of the Cape of Good Hope, produced by a Distoma/ rep. in
'Lancet/ and ' Med. Times and Gaz./ Feb., 1864; also in
Ranking's 'Abstract/ p. 1 73, 1864, and fully in ' Medico- Chirurg.

PARASITES OP MAN

Trans./ 1865. — Idem, " On the Endemic Hematuria of the
South Eastern Coast of Africa/' 'Med.-Chir. Trans./ vol. liv,
1871. — Idem, in Hooper's ' Vade Mecum/ 1869. — Hatch, W. K.,
"Case of Bilharzia hamatobia," in ' British Medical Journal/ Dec.
14, 1878, p. 875. — Kiichenmeister, F., ' Parasiten/ 1855; Eng.
edit., p. 277, 1857.— Leuclmrt, E., 1. c, s. 617, 1863.—
Sonsino, P., " Richerche intorno alia Bilharzia haematobia in
relazione colla Ematuria Endemica dell' Egitto e nota intorno
un Nematoideo trovato nel Sangue Umano," ' Estr. dal Rend.,
del. R. Accad./ 1874. — Idem, ' Delia Bilharzia haematobia e delle
alterazione Anatomo-patologiche che induce nell' Organismo
Umano, loro importanza come Fattori della Morbilita e Morta-
lity in Egitto, con cenno sopra una Larva d'Insetto Parassita
dell' Uomo. Estratto dall' Imparziale/ Firenze, 1876. — Idem,
' Sugli ematozoi come contributo alia Fauna Entozooca Egi-
ziana,' Cairo, 1877. — Idem, "La Bilharzia haeniatobia, et son
role Pathologique en Egypte," ' Arch. Gen. de Medicine/ for
June, p. 650, 1876. — Idem, " Intorno ad un nuovo Parassita del
bue {Bilharzia bovis)," ' Estr. dal Rend. del. R. Accad. di
Napoli/ 1876.—Weinland, D. F., 1. c, p. 67, 1858.

SECTION II.— Cestoda (Tapeworms).

Tania medio canellata, Kiichenmeister. — This cestode is fre-
quently spoken of as the unarmed or beef tapeworm. In general
appearance it is very similar to the armed form. It is, how-
ever, a larger and broader animal, being at the same time rather
stouter. It varies usually from fifteen to twenty-three feet in
length, but specimens have been described as attaining thirty
feet. It is called the unarmed tapeworm in consequence of the
absence of any coronet of hooks on the head ; and consequently,
also, from there being no prominent rostellum or proboscis.
The place cf the last-named structure, however, is supplied by a
small rudimentary disk, which I have seen protruded on pres-
sure (fig. 20). Usually this disk forms a more or less conspicuous
cup-shaped circular depression, which has been compared to and
described as a fifth sucker. That it is not, in any structural
sense, comparable to the true suckers, I have had abundant
opportunity of ascertaining ; nevertheless, I do not doubt that it
is to a slight extent capable of being used by the parasite as a
supernumerary holdfast. The anchorage thus secured, however,
is by no means equal to that obtained by the armed species.

CESTODA

This explains the comparative difficulty we find in procuring a
specimen of the armed tapeworm with the head attached.

Fig. 20.— Head of Tania mediocanellata. Showing the calcareous corpuscles, suckers, rudimentary
proboscis, and water vessels. Highly magnified. Original.

The establishment of this species as distinct from T. solium
is due to Kiichenmeister ; but it is curious to observe how
accurately this determination was foreshadowed by the shrewd
naturalist and theologian, J. A. E. Goeze, who clearly indicated
two forms of the common tapeworm, remarking (1. c, Bibl. No.
1, s. 278): — "Die erste ist die bekannte grosse, mit langen
dicken und gemasteten Gliedern, die ich Tania cucurbitina,
grandis, saginata, nennen will." The same author (s. 245)
pointed out the resemblance subsisting between the tapeworm
of the cat (T. crassicollis) and the vesicles (" Krystallblasen ")
and their contained " erbsformige Blasen " [Gysticercus fascio-
laris ) of the mouse. Thus the celebrated pastor of St Blasius,
in Quedlinberg, almost contemporaneously with Pallas, early

PARASITES OF MAX

arrived at the conclusion that the hydatid- measle was a kind of
tapeworm.

Respecting the organisation of this worm I may observe that
the mature joints have a more complicated uterine organ than
obtains in Tania solium, presenting nearly double the number

of lateral branches. They
are more closely packed,
running outwardly in an
almost parallel manner.
The first sexually ma-
ture proglottis occurs at
about the 450th joint,
but whereas, in the pork

Fig. 21.— Free proglottides of T,enia mediocaneilata. After tapeworm, Onlv SOmG 200
Leuckart. r > J

subsequent segments share
this perfect character in the beef tapeworm, according to Leuc-
kart, as many as 360 or even 400 mature joints may be present.
The joints are very liable to form monstrosities ; these abnor-
malities sometimes affecting the reproductive organs, which
become doubled or even trebled. In the Hunterian collection
there is a- proglottid showing twenty-two sexual orifices. Dr
Cullingworth, of Manchester, has described a specimen in which
the joints are curiously tripartite.

As already hinted the true source of this parasite has been
proved by experiment ; the first successful worm-feeding having
been accomplished by Leuckart. Mosler's, and subsequently
my own feeding experiments, immediately followed. Other
successful experiments with this species have been conducted
by Zurn, Probstmayer, St Oyr, Perroncito, Masse and Pour-
quier, and Zenker. As will be again mentioned below Dr
Oliver, R.A., whilst stationed at Jullundur, successfully reared
the adult tapeworm in a Mohammedan groom and in a Hindoo
boy. It will also be seen that Prof. Perroncito reared the worm
in a student in fifty-four days. In my own experiments on
animals I was assisted by Professor Simonds. The feeding
materials were tapeworms expelled from my own patients.
We obtained the following interesting results :

jfcp, 1— A calf. First feeding, Dec. 21st, 1864. Marked
symptoms. Slaughtered April 3rd, 1865. Result positive.

JExp.2. — A calf. First feeding, April 13th, 1865. Second, third,
and fourth feedings in May and June. No symptoms. Died
on Sept. 3rd, 1865, after thirty-six hours' illness with " cattle

OESTODA

plague." Result stated to have beeu negative as far as the
muscles were concerned. Viscera not examined.

jSkp, 3 _A Dutch heifer. First feeding, March 3rd, 1865.
Three subsequent feedings. Symptoms only slight. Slaugh-
tered April 4th, 1866. Result positive. Measles especially
numerous in the diaphragm, but all had undergone calcareous
degeneration.

Exp. 4.— A calf. Fed May 27th, 1872, with ripe proglot-
tides. Marked symptoms set in on June 7th, which began to
abate on the 12th, and had nearly disappeared by the 20th of
the same month. The record of the post-mortem result has
been lost ; but the animal was infected.

j^p. 5. — A calf, which had been made the subject of a
" glanders experiment." First fed on Oct. 17th, 1872, and
thrice in the following year, Jan. 1st and 11th, and March 8th.
No symptoms having appeared the animal was kept for six or
eight months after the last feeding. Seeming to be free from
disease of any kind, it was sold as a sound heifer.

Exp. 6. — A young heifer calf, of six months. Fed Oct. 18th,
1873, with the mature proglottides of a large beef tapeworm.
No symptoms. Slaughtered several months afterwards. Result
stated to have been negative. Unfortunately I was not present
at the autopsy.

Exp. 7. — A young heifer. First fed May 19th, 1874, with the
joints of a tapeworm, and again on June 12th. No apparent
ill effects resulted, but the animal died in October. At the
post-mortem examination, made by Prof. Simonds, no parasites
were observed. Subsequently I found calcareous specks in the
liver which proved to be degenerated measles.

Exp. 8. — A calf. Fed on or about March 24th, 1875, with
sexually mature joints. The calf was put to and remained
with a foster mother until it died from disease of the larynx on
the 15th of the following July. The animal was ill-treated by
its foster parent, and at the post-mortem I observed a large
intercostal cicatrix, evidently the result of injury. In this case
I devoted several hours to the exploration of the muscles aud
viscera. Not a trace of the Oysticercus bovis could be found in
the muscles or connective tissues, but the liver contained scores
of perfectly developed measles, besides hundreds of others in
various stages of calcareous degeneration. On comparing some
of the latter with those I had obtained from the precediug
experiment the pathological appearances were at once seen to be

PARASITES OF MAN

identical. It was easy to find and pick out the measles in their
cysts from the naturally friable liver. I also detected four
Cysticerci in the lungs, two of which had degenerated. Micro-
scopic examination confirmed my interpretation of the naked-eye
appearances.

Fig. 22.— Section of the heart of a calf infested by cestode larvsc. After Mosler.

Fragmentary as the above data are, they serve to show that
we have hitherto been too hasty in concluding that beef and veal
measles reside only in the voluntary and striated muscles of
their hosts. The facts here recorded prove that the liver of a calf
may be extensively invaded by cysticerci, and yet the animal will
exhibit no sign of constitutional disturbance. The cestode
tuberculosis may come and go without any diagnostic symptom,
whilst a few months suffice for the natural death and decay of
the parasite by calcareous degeneration. Thus it becomes
extremely probable that many experiments hitherto regarded
as negative in their results have really been positive ; the
pathological evidences having been either misinterpreted or

CESTODA

altogether overlooked. Every pathologist is familiar with
gritty particles in the various viscera of man and animals, but
few are probably aware how constantly these are dead and
degenerated Cysticerci. The gritty particle itself may be
reduced to the merest point, no larger than the receptaculum
capitis of the Cysticercus itself, and in course of time it will
disappear entirely. Practically it is satisfactory to have experi-
mental evidence of the fact that cattle, as well as other animals,
however extensively measled they may have been, can become
thoroughly cleansed of the disorder by nature herself. It is
only necessary that the diseased animals be separated from
infectious influences.

Although the beef measle has never yet been found in
man, I have for convenience sake introduced the facts of
larval parasitism in this place. The sanitary bearings of this
subject are far too important to be dismissed in a summary
manner. I have shown that the prevalence or rarity of the
beef tapeworm in man is strictly dependent upon the habits of
the people ; this same cause operating to produce healthy or
diseased meat-food, according to the degree of civilisation. In
this connection the oft-quoted statements of Kas chin respecting
the prevalence of tapeworms among the Burates, and the well-
known frequency of this entozoon in Abyssinia, need only be
alluded to.

When discussing the food question in my ' Manual/ I freely
availed myself of facts privately communicated by Dr. Joseph
Fleming, and I especially referred to the published labours of
Lewis, Hewlett, Veale, and other observers stationed in India.
Beef measles are extremely common in the cattle of the north-
west provinces of India, so much so that severe restrictions
have been imposed upon the consumption of ration beef. The
presence of a few measles in the flesh of cattle has been deemed
a sufficient excuse for condemning and burying entire car-
cases. The measle is easily distinguished from that of
mutton and pork by the fact that its head is not furnished
with hooks, whilst in the place of a rostellum there is a small,
centrally placed, retractile disk, which assumes the appearance
of a supplementary sucker as in the adult worm. The four
true suckers are also comparatively large. The measle usually
varies in size from the fourth to the half of an inch in length,
but my cabinet contains a specimen nearly an inch long. This
was contributed by Dr J. Fleming, who mentions having seen

PARASITES OF MAN

a measle which, when unrolled, measured nearly an inch and
a half in length. Although thousands of these bladder
worms must exist in the cattle of England, up to the pre-
sent time not a single instance has been recorded of the
occurrence of these cystic parasites in the United Kingdom,
except in our experimental animals. Notwithstanding my
inquiries, I have not yet found a butcher, flesher, meat-
inspector, or veterinarian, who has encountered this parasite in
any animal slaughtered for the market. Several butchers have
denied their occurrence in meat sold by themselves. Even so late
as June, 1874, the presence of measles in the flesh of cattle was
denied before an assembly of French savans ; yet for many years
past I have constantly exhibited measly beef and veal in the
lecture room of the Royal Veterinary College. (See the discussion
of the Societe de Therapeutique, recorded in the ' Bulletin Gen.
de Ther/ for June 30th, 1874, and also the ' Jour, de Ther./ No.
14, for July, p. 556, where, however, special remarks on this
head have been omitted ; see also the ' Lond. Med.. Record ' for
July 29th, 1874, p. 472, and the ' Lancet ' for Dec, 1874, p. 794.)
Quite in contrast with the statements referred to are those of
recent Italian observers.

Some few years back Professor Gr. Pellizzari communicated
to the Medico-Physical Academy, at Florence, the results of a
series of experiments conducted by himself, with the assistance
of Dr Tommasi, in regard to the temperature necessary for the
destruction of cysticerci in measled meat. An account of these
experiments is published in Tommasi's edition of my ' Manual/
The researches were made in relation to certain sanitary
measures effected by the Municipal Commission of Florence,
the express object of these measures being to prevent the
injurious distribution of measly meat, especially that of swine.
Signor Bosi, the superintendent of the public slaughter-
houses, granted every facility in his power. In a previously
published memoir by Professor E. Perroncito it was .stated
that measly meat (panicatura degli animali) required a higher
temperature than that of boiling-point for the destruction
of the bladder worms in question. In this opinion Signer
Bosi shared. According to the original memoir of Perroncito
we are told that " about twenty specimens of Cysticerci were
collected by the author, and placed in boiling water. After
twenty minutes' boiling, not one of the parasites appeared to
suffer. The head continued to be drawn into the body, and

CESTODA

when the Cysticerci had their heads drawn out one by one they
still appeared to possess all the elasticity of living bladder
worms, displaying those movements of extension which are
proper to parasites not yet dead. The hooks were observed
regularly disposed on the proboscis, where they formed a double
crown, the suckers remaining intact." Perroncito remarked,
however, that the Cysticerci showed a coloring tendency
towards brown, and he added that " with the aid of two needles
it became easy to lacerate the body of the Cysticercus, which
appeared to be swollen, and possessed of diminished cohesion
of its parts." It was evident to all eyes, observed Professor
Pellizzari, that these statements involved clear contradictions.
Yet again, at page 28 of the memoir, Professor Perroncito
wrote : — " During the past winter I introduced some little slices
(fettuccie) of muscle-flesh (8 to 10 millimetres in thickness),
infested with Cysticerci into a vessel {cassolina) containing fat
at the temperature of 190 to 200° Cent. (374 to 400° Fahr.).
At the expiration of ten or fifteen minutes the slices of meat
were fried, and the Cysticerci lying at the surface had acquired
a light brownish colour, as if they were roasted. By breaking
up the slices one could still see the small reddish muscular
bundles, whilst the Cysticerci in the middle remained entire
and well preserved. Their heads displayed the hooks and
suckers regularly distributed." It is certainly singular, as
Pellizzari observes, that these Cysticerci, having been thoroughly
fried and roasted, should still remain alive and in their normal
state ; but the ultimate conclusion at which Perroncito arrived
was still more startling, and one which, if it were true, would
not fail to create a considerable stir among our officers of health.
On reviewing the whole matter Perroncito says : — " It appears
to me that the melted fat alone of hogs (maiali grandinosi)
should be utilised, and I am pleased to reckon the illustrious
Gerlach and all other distinguished practitioners to be of the
same opinion. Permit me, therefore, being well satisfied also
with the results of many other experiments, once more to
advance the conclusion that, if it is not certain that the
Cysticerci die at from 80 to 100° Centigrade (176 to 212° Fahr.),
we are quite sure that they dry up and become completely
mummified at 125, 130, and 150° Cent. (257, 268, and 302°
Fahr.), temperatures which we could easily produce by means
of a properly constructed apparatus."

After remarking upon the serious nature of the conclusion

PARASITES OP MAN

which Perroncito sought to establish, Professor Pellizzari
makes further use of quotations which bear upon the question
as to whether the quality of the vessels in which the fat of
diseased hogs is melted down may not largely affect the degree
of high temperature sought to be obtained (in view of a perfect
destruction of the Cysticerci). Perroncito repeatedly witnessed
the operations of pork-butchers ; and when portions of meat
were introduced, with water, into the cauldrons, he always saw-
that the temperature " was maintained between 97° and 98°
Centigrade." However, this part of the question may be dis-
missed in a very few words, since Perroncito himself finally
allows that " the different composition of the vessels cannot
elevate the temperature of* the fat by many degrees."

With the praiseworthy intention of either verifying or refuting
these conclusions, Pellizzari, with the approval of Bosi and
with the assistance of Tommasi, instituted a fresh series of
experiments at a private laboratory. The details of these
experiments are exceedingly interesting ; but as their record
occupies several pages of Tommasi's appendix already referred
to, I must content myself with a general statement of the results
obtained. Professor Pellizzari found that Cysticerci, so far
from requiring a temperature of upwards of 100° Centigrade
for their destruction, die at a temperature of 60° Centigrade
(140° Fahr.). He had, it appears, previously taken the initia-
tive in recommending certain measures to the Florentine
municipality, in view of protecting the public health, and he
had now the satisfaction of more than confirming the wisdom of
these sanitary precautions. In excessively measled animals the
fat is removed and boiled in suitable cauldrons, and has potash
mixed with it to render it useful for industrial purposes. By the
various measures adopted the entire animal is utilised, and with
proper precaution there seems little chance for the measles to
arrive at the tsenioid or sexually mature condition.

In the next part of his communication Pellizzari touches upon
the question of measles in beef, referring especially to the
experimental labours of Leuckart and myself. Finding addi-
tional support from our views Pellizzari declared the proposi-
tions of Dr Perroncito as of no value whatever. " But how is
it," he adds, " that notwithstanding that so low a temperature
suffices to kill these cysticerci, yet cases of Taenia are continu-
ally occurring 1" The answer to this question will appear in
the sequel ; but meanwhile it will be as well to refer to the

OESTODA

recent brochure by Dr Giacomini. This author appears to have
had no opportunity of perusing Pellizzari's communication
already cited, and consequently it is not surprising that he should,
in common with others, have accepted the original conclusions
of Perroncito. Dr Giacomini clearly perceives that, whatever
precautions of a hygienic character are suitable for the preven-
tion of disease arising out of the consumption of measly pork,
the same, or at all events similar, measures ought to be adopted
with the view of checking tapeworm affections arising from the
ingestion of other kinds of meat, especially veal and beef. Like
Pellizzari, he is satisfied as to the human origin of the small
bladder worms found in cattle, and establishes this position
not only from the oft-quoted experiments of Leuckart and
Mosler, but also from those conducted by myself and Simonds
in England, and by Professor P. Saint-Cyr in France. From
a careful review and consideration of all the facts of the case,
he recommended a more complete supervision over the flesh of
oxen before it is employed commercially, and greater precaution
when employing veal as food, by causing it to be subjected to
a high temperature, in order that the parasites may be killed
before it is ingested. It is evident that Giacomini thinks that
a temperature exceeding that of boiling-point is necessary for
the destruction of the beef and veal measles, since he immedi-
ately adds, " Though experiments have not been made with the
object of ascertaining the amount of resistance of heat which
the unarmed cysticercus can bear, yet, judging by those con-
ducted by Professor Perroncito on the measle of the hog, we
are in a position to say that a temperature of 135° Cent. (275°
Fahr.) is necessary for the destruction of an isolated Cysticercus,
whilst the heat should be raised from 150° to 200° Cent. (302°
to 392° Fahr.) for ten or fifteen minutes, in order to ensure the
complete destruction of the Cysticerci encapsuled in the interior
of a piece of meat." I have abridged this portion of Giacomini^s
text, because his statements are pretty much the same as those
already quoted from Perroncito (as cited by Tommasi). But,
in the next place, Dr Giacomini is in error when he states that
experiments had not been performed on the Cysticerci of the
ox. So far from this being the case, similar experiments
had long previously been conducted by Dr Lewis in India;
and these researches had quite as much to do with the
measles or Cysticerci of beef as they had with those of the
hog, if not more. Naturally but few foreign investigators can

PARASITES OP MAN

have had access to the work in which Lewis's experiments were
originally recorded, and to which, therefore, I must call their
attention. Thus, Dr Tommasi has fallen into the error of
supposing that the investigations of Lewis were made in England.
It is of very little moment where the experiments were carried
on, but Tommasi's statement (appendix, loc. cit., p. 161),
wherein he says that Pelizzari's experiments, in which he
himself took part (ai quali io stesso ho assistito), are even more
complete than those made in England by Dr Lewis, and
in Germany by Dr Kiichenmeister, cannot be allowed to pass
unchallenged. If Tommasi had enjoyed the opportunity of
consulting Lewis's original memoir, he would not have under-
estimated our countryman's labors. The memoir by Lewis
is singularly complete, and well-nigh exhausts all the facts
that can have any interest in relation to the question of
public health. Towards the close of his essay he expressly
states, as the result of investigation — " (1) That exposure to a
temperature of 120° Fahr. for five minutes will not destroy
life in Cysticerci, but that they may continue to manifest indica-
tions of life for at least two or three days after such exposure ;
(2) that exposure to a temperature of 125° Fahr. for five
minutes does not kill them ; but (3) after being subjected to
a temperature of 130° Fahr. for five minutes, they may be
considered to have perished. After exposure to this and higher
temperatures, in no instance have I been able (he adds) to
satisfy myself that the slightest movements took place in their
substance when examined even under a high power. At least,
it may be confidently asserted that, after exposure for five
minutes to a temperature of 135° to 140° Fahr., life in these
parasites may be considered as absolutely extinct" (p. 139).
Thus the statements of Lewis and Pelizzari were in perfect
accord; and seeing that their conclusions were alike the result
of very careful and independent inquiry, it seemed as if
the question at issue was finally solved. These investigations
made it perfectly clear that Cysticerci of all kinds, whether
found in veal, beef, or pork, could not retain their vitality when
exposed to a temperature of 60° Centigrade, or, in other words,
140° Fahr.

The rather severe strictures made on Perroncito's earlier
experiments induced the Turin professor to go over the subject
more carefully, when he obtained excellent results. He finally
ascertained that Cysticerci perished at a temperature below

CESTODA

50° C. (122° Fakr.). In May, 1877, Dr Perroncito furnished
me with an account of his researches. With the exception of
a few verbal alterations, for which I am responsible, Perroncito
wrote as follows :

" In order to resolve the highly important question of the
tenacity of life of the Helminths and corresponding larval
forms, I made since 1871 a very long series of experiments on
the Gysticercus cellulosce, which were published almost at the
same time with others of the same kind, made by Dr Lewis in
Calcutta. Towards the end of 1874 Mr Pellizzari, of Florence,
disputed the results of the investigations which I had made
known two years before, i.e. in 1872, and agreed with Dr Lewis,
who had stated already that the Gysticercus exposed to a tem-
perature of 55° C. can be held for dead after five minutes, and
also with Dr Cobbold, who thought the temperature of 60° C.
quite enough to kill it. But the characters he (Mr Pellizzari)
relied upon, needing the exactness and precision required to
enlighten and persuade in the most important scientific questions,
gave rise to a mistrust in the most scrupulous amongst the men
devoted to biological pursuits and to several hygienic measures
on the part of the sanitary inspectors with regard to in-
fected pork. Therefore, my conclusions, argued from the
experiments made in 1871-72, were still those followed by the
most important Italian cities, and approved in principle by the
superior Board of Health in 1873. I expressed doubt ther>
about the Gysticercus dying at a temperature lower than 100° 0.,
and some person misconstrued these doubts, saying that I
had contradicted myself in my work. However, as I could not
assert they died at 80° — 100° C, I only noticed the alteration
of color and cohesion which happened in the Gysticercus
exposed to various degrees of temperature, to the end that I
might contribute usefully to the solution of the difficult ques-
tion, and concluded that * if we could not be sure of the
Gysticercus dying at 80° — 100° C, it was certain at all events
that they perished at 125° or 130° C Not wishing to pre-
judice the question, I never said that they did not die at 80° —
100° C., but simply stated that at this temperature we could
not be certain of their death.

" Now, after a large number of experiments, I have been
able to ascertain with exactness the lowest degree of temperature
required to kill infallibly the Gysticercus and other parasites of
animals. The means I made use of for this kind of investiga-

PARASITES OF MAN

tion were Mr Schulze's heating table, the neutral tincture of
carmine, the tincture of heematoxylon, and breeding experiments.
" My method is founded essentially —

"(a) On the fact that the Oysticercus when it is fresh and is
stretched and conveniently prepared in pure water, or in chloride
of soda veiy much diluted, and afterwards brought gradually
from the temperature of the ambient air to that of the body of
higher animals and to degrees of heat still more elevated, until
life is extinct, keeps moving to and fro with more or less
energy throughout its body, using especially its suckers and
proboscis.

" (b) On the greater imbibing power of the dead tissue
generally, which is undoubtedly far more apparent in insects
and plathelminths.

" (c) On the experiments made to ascertain the value of the
two above- stated facts.

" If, after having prepared a Gysticercus, newly extracted
from a pig in the way we have pointed out, we examine it with
a microscope on M. Schulze's heating table, we find that
usually it begins to move after 30° or 35° 0., and each moment
with greater activity, especially after 38°, 40°, 42°, 44°, 45° C.
The temperature being raised progressively, we see that the
Oysticercus cellulosa puts a stop to its movements occasionally
at 45 — 46° C, seldom at 47° 0., more frequently at 48° C,
sometimes at 49° 0. ; and, in fifty and more experiments, only
one Gysticercus was able to live on beyond 49° C, standing still
at 50° C.

"As soon as it stands still the parasite is dead. In fact, if
we lower again the temperature gradually to that of the ambient
air, and if afterwards we raise it a second time, we pass through
all the intermediate temperatures without the Gysticercus show-
ing the least signs of life.

" But a more convincing proof of the death of the parasite is
got from the greater imbibing power of the tissue when life is
extinct, the same over the whole body of the plathelminths, and
their larval forms. If we dip the Gysticercus alive with its
head stretched in the neutral tincture of carmine or hjBinatoxylon
we can leave it there even two, four, eight, ten, or twelve hours
and more, without the head coloring or a real imbibition taking
place ; this begins only after the Gysticercus is dead, so that if
the Oysticercus is brought first to a temperature hot enough to
kill it (with M. Schulze's tables to one of 48°, 49°, 50° C.) and

CESTODA

dipped afterwards in the above-mentioned tinctures, it colors
intensely in less than 45°, beginning from the head, and onwards
to the extremity of the cyst of the tail. The head colors
more intensely and rapidly than the neck, as it is covered with
very numerous calcareous corpuscles, which are not met with
so frequently in the remaining part of the body.

" Cysticercus celluloses of the pig, and that of the ■ Taenia
mediocanellata of the calf, brought gradually to a final tem-
perature, the first of 50° C, and the second of 44°, 45°, and 47°
C, and then swallowed alone, or with a piece of butter or
crumb of bread, never produced the Taenia in the valiant
students who voluntarily undertook to make the experiment of
swallowing them.

" My investigations were extended to other kinds and forms
of Helminths, and the results were always the same, so that,
abiding by the same principles, I was able to ascertain that —

"1st. The Cysticercus cellulose of the pig dies sometimes at
45° C, more frequently at 47° C, ordinarily at 48 C, very
seldom reaches alive 49° C, and is quite an exception when it
resists for a few moments the temperature of 50° 0., so that we
can say that the Oysticercus brought gradually up to this tem-
perature most assuredly dies if it is kept there longer than one
minute.

" 2nd. A Cysticercus cellulosae, extracted by Professor Ray-
mond from the conjunctiva of a child's eye, died between 45°
and 46° C.

"3rd. The Cysticercus of the Taenia mediocanellata dies
sometimes at 44° C, very often at 45° C, and does not resist a
temperature superior to 46° C.

" 4th. The Cysticercus pisiformis of the rabbit, like the
celluloscB, dies sometimes at 45° and 46° C, but generally stands
still and perishes at 47° and 48° C.

" 5th. A Cysticercus tenuicollis died at 49° C.

" 6th. The scolici of the Caenurus cerehralis of a sheep died at
42° C.

" 7th. The scolices of the cysts of Echinococcus polymorjphus
die generally between 47° and 48° C, and in no case amongst
those I have experimented on did it reach 50° C. alive.

" 8th. The Taenia cucumerina died, one at 43° C, and a
second parasite at 45° C.

" 9th. A few individuals of Taenia scrrata of the dog died at
50° C.

rARA SITUS OF MAN

" 10th. Two individuals of Tania perfoliata of the horse died,
the first at 45° 0., the second at 50° C.

"11th. The embryos of the Filaria microstoma of the horse
began to stand still at 46 — 47°, and all died at 48° C.

"12th. The embryos of the Filaria megastoma of the horse's
stomach died at 47° C.

" 13th. The Trichina spiralis, both free and in a cyst/ in
several experiments always died at 48° C.

" 14th. The embryos of the Strongylus filaria of the sheep
stood still at 50° 0.

" 15th. Probstmayer's viviparous oxyurids, the infusoria of
the colon and caecum of the solipeds, and the psorosperms of
the liver of the rabbit did not stir at all.

" Each experiment lasted about ten minutes, and the tem-
perature rose from 8 — 10° C. to 45 — 46° C. in six to eight
minutes ; and from 46° to 50° in one minute. These experi-
ments have a great value, both scientific and practical, as they
show, on one side, which is the lowest intensity of heat suffi-
cient to kill always the Oysticercus, the Trichina, and other
parasites, reducing thus by far the tenacity of life generally
attributed to a large number of Helminths and corresponding
larval forms. They assure us, moreover, of the harmlessness of
the flesh infected by the above-mentioned parasites, when it is
cooked in such a manner as to reach the temperature of 50° C.
over all points of the pieces, even though it be kept at such a
degree of heat not longer than five minutes.

" In a piece of leg of pork the Oysticerci were found alive in
all places not yet putrefied twenty-nine days after the animal
had been slaughtered. On the other hand, in the dry muscles
of a calf the Cysticerci of the Tcenia mediocanellata were all
found dead fourteen days after the slaughtering of the animal.
I have ascertained that putrefaction of the flesh is fatal for the
two larval forms of these different kinds of helminths."

In a subsequent communication received from Professor
Perroncito towards the close of the year 1877 he writes :

" At the last meeting, held on April 23rd, I made a state-
ment to the Medical and Surgical Society of Turin, of the
results of other experiments tried by heating at M. Schulze's
table and by the imbibitions with the neutral tincture of car-
mine, through which I came to the conclusion that the Cysticerci
of the Tcenia mediocanellata die sometimes at 44° C, now and
then at 45° C, and always at 46° C. I therefore concluded

CESTODA

that they could in no case survive at 47° C. and 48° C. when
they were maintained at this temperature at least five
minutes. But to the end of more fully corroborating the facts
I had thus communicated, I, contemporaneously with these,
made some breeding experiments with the same Gysticerci on
bold and courageous students who generously offered themselves
for the benefit of science.

" Consequently I am now enabled to state that neither Mr
Gemelli nor Dr Eagni contracted the Taenia, though each of
them had eaten a Cysticercus of the Taenia mediocanellata pre-
viously, and respectively subjected to a temperature of 45° C.
and 47° C. The larvae were properly prepared and submitted
to gradual heating on the above-mentioned table, and swallowed
when they no longer gave signs of life. In like manner no
generation of the Taenia took place in the body of Mr Martini,
who ate the Cysticercus brought to a temperature of 44° C. It
was maintained at this degree of heat during a period of about
three minutes, and swallowed whilst a very slight movement
was still visible in a portion of its neck.

" In another student, on the contrary, who ate a living Cys-
ticercus of the Taenia mediocanellata, the tapeworm reached its
maturation in fifty-four days and eliminated the two first pro-
glottides. It threw off two more on the fifty-eighth day, and
thirty on the sixtieth. Sixty-seven days after swallowing the
Cysticercus this courageous young man, having, like his three
companions, taken some kousso and castor oil, emitted the strobila.
It was furnished with 866 rings, but destitute of the neck and
head. Its measurement afforded a total length of 4*274 metres.

" Adding now to the 866 proglottides the thirty-four already
eliminated, 900 would be the number of the segments ; and
reckoning the length of each of the latter to be fourteen milli-
metres, we should have had the strobila (deprived of the head
and neck) reaching a length of 4" 75 metres. Further, calcu-
lating the head and neck to be eight millimetres long, a total
length of 4*83 metres would be the result.

" From all these facts we may conclude that the Taenia has,
in our instance, reached an approximative length of seventy-two
millimetres a day, affording a daily production of 13'43 pro-
glottides."

In relation to requirements of state medicine I have
thought Perroncito's researches sufficiently valuable to be quoted
at some length ; but their chief interest culminates in the worm-

PARASITES OF MAN

feeding experiments. Excellent in all respects as was the
conduct- of the medical students who, with Professor Perroncito's
approval, swallowed living specimens of the Gysticercus bouis, the
intentional ingestion of beef measles is by no means a novelty.
Bight or ten years back Dr Oliver (after explaining to one
of the selected victims the possible consequences of the experi-
ment) induced a Mahommedan syce or groom and a Hindoo boy
to swallow perfectly fresh and living beef measles. In this
way Dr Oliver successfully reared the Tcenia mediocanellata in
India, and he was thus enabled to fix the amount of time
necessary for the full growth of the strobila. Many other
persons have diplayed an equal amount of zeal in the cause of
helminthology, by partaking of the larvae or germs of other
parasites. Thus, at the risk of repetition, I may state that
Moller many years ago swallowed the slender-necked hydatid
(Gysticercus tenuicollis) in the hope of infesting himself with
Taenia marginata. Several persons have defiantly swallowed
trichinised flesh. Professor Leuckart and some of his pupils
also courageously swallowed the eggs of Oxyurides, and they
had the infinite satisfaction of noticing the young worms in
their feecal discharges some fifteen days afterwards. Dr Crisp
ate part of the cooked flesh of an animal that had died of
cattle plague, and I myself partook of moderately cooked meat
which I knew to be swarming with psorosperms. These
obscure organisms were by some persons considered to be
either a cause or product of the rinderpest. They will be
noticed in my account of the Protozoal parasites.

For the purpose of advancing science and the welfare of the
people, there are scores of persons always to be found ready
to make personal sacrifices of the kind undertaken by Drs
Ragni, Martini, and Gremelli. Unfortunately for English
science there are not wanting people in this country who are
prepared to threaten with fines and imprisonment any savant
who may think it desirable to perform a similar set of feeding
experiments on animals. Invaluable for good as our experi-
mental investigations have already been, it would seem as if it
were the deliberate aim of these sentimental obstructives to
put a stop to the acquisition of all useful knowledge in the
future.

In reference to the rate of growth of tapeworms, Professor
Perroncito's determinations are useful, inasmuch as they verify
certain ascertained facts with precision and confirm the general

CESTODA

conclusion that had been drawn by practical helminthologists
from various sources of information. In regard to the number of
proglottides proper to a sexually mature tapeworm, the circum-
stance that Perroncito's calculation was made without the head
and a portion of the neck of the worm being present shows that
it cannot be relied on absolutely ; nevertheless, as far as it goes,
it tends to confirm what Leuckart had long previously stated.
I have possessed myself of upwards of thirty perfect beef
tapeworms expelled from my patients, and in some of the
specimens it was noticed that the segmentation-rings in the
region of the neck were far more crowded together than they
were in others. I also possess a perfect Tcenia medio canellata,
removed post mortem. Though the rate of growth may be the
same from day to day, yet experience has shown that the number
of proglottides actually cast off varies exceedingly. Kiichen-
meister's estimate of the average number agrees in the main
with what we have ourselves observed (five to twenty daily) ;
and here again Perroncito's investigations serve to verify the
general correctness of our previous determinations.

To return to Pellizzari's researches, one of the most important
questions is that which relates to the prevalence of tapeworm.
In this connection he first brings forward some very interesting
and instructive data that had been previously communicated to
the Medico-Physical Academy of Florence by Professor Marchi.
On the occasion referred to Marchi had stated that, out of thirty-
five Taeniae which he had examined, only one belonged to the
species known as Tcenia solium ; all the other thirty-four being
of the unarmed type, or Tcenia mediocanellata. Reflecting on
this striking fact, and also on the circumstance that he had in vain
begged his colleagues to send him specimens of Tcenia solium,
Marchi seems to have missed the very palpable explanation of
this otherwise strange phenomenon. " How does it happen,"
exclaimed Marchi, " that, notwithstanding the occurrence of
13,000 kilogrammes of the flesh of measled hogs in the public
butcheries, I have seen but one specimen of Taenia solium,
whilst thirty-four cannot have originated from the pig V " The
wherefore is obvious enough," replies Pellizzari, " because our
hygienic regulations demand that the flesh of the hogs be raised
to a temperature of 60° Cent. (140° Fahr.) f and he then himself
immediately proceeds to ask another question, namely, as to how
it happens that the TcEnia solium is so frequently seen in other
places. To his own question Pellizzari responds by remarking

PARASITES OP MAN

(1) that there are not so many precautions (of a sanitary kind)
taken in other places ; and (2) that the people elsewhere
consume more slightly salted or uncooked meat, as sausages and
so forth (come salame giovane, salciccia e via dicendo). Pellizzari,
having explained that Marchi's thirty-four tapeworms must
all have arisen from the consumption of the Cysticercus of the
ox, then goes on to speak of the prevalence of tapeworm in
Florence, even in little children. This last-named feature, he
says, is due to the circumstance that raw meat is frequently
employed as a restorative (come cura ricostituente) . " Thirty
years ago," remarks Professor Pellizzari, "it was just as
difficult to find a single Taenia mediocanellata as it is now easy
to find a great number of these worms ; and all because it is
nowadays customary to eat the flesh of the ox either insuffi-
ciently cooked or raw. This absolute inversion of the facts of
the case affords proof of the correctness of the position sus-
tained by me, to the effect that the cooking of meat up to the
degree of temperature necessary for ebullition ensures the
destruction of the Cysticerci." Notwithstanding this state-
ment of his own, Pellizzari thinks that the interference of
inspectors may be pushed too far, and thus serve to bring
about the very disasters which it should be their supreme
object to prevent. Thus, he argues against the suggestions of
those who would entirely prevent the sale of measly meat, and
who would only permit, as obtains in the province of Modena,
the melting down of the fat of hogs. Very strict measures of
this sort would, as he says, constitute a radical means of entirely
stamping out Taenia, but he also very judiciously reminds the
sanitarian (igienista) that " such a step would be a serious
thing for the tradesman, bringing injury not only to the muni-
cipal administration, but also proving an encouragement to
smuggling. In this way the public health would sustain worse
injury by the inducement held out to the owners of infected
animals to slaughter them in secret butcheries, thus little by
little withdrawing the meat from the superintendence of the
public officials. By the adoption of fraudulent measures there
would be a daily consumption of diseased meat ; and thus also,
while the public administration would suffer loss, the public
health, on the other hand, would gain nothing." In effect
Pellizzari says, if we advise the employment of more severe
and radical measures than those already in vogue in Florence,
we should overburden the tradesman, almost compel him to

CESTODA

defraud the exchequer by smuggling, and greatly injure the
public health.

The facts and explanations advanced by Italian writers
regarding the causes of the endemic prevalence of tapeworm,
are in perfect harmony with those previously obtained from
other sources. Respecting these causes there is much that is
both new and interesting. The eighth annual report of the
sanitary commissioner of the Government of India had already
made us acquainted with the fact that during the year 1869, out
of 13,818, head of cattle slaughtered in the stations of the
Upper Punjab, 768 beasts were found to be infected with
measle-cysts. This, as I have remarked (Tommasi^s edit.,
p. 54), " affords a rate of 5*55 per cent., being a considerable
diminution of the proportion observed in 1868, when the per-
centage gave a total of 6*12. The reduction was, without
doubt, due to the vigilance and enlightenment of the army
meat inspectors. The prevalence, however, of tapeworm does
not bear relation to the number of animals infested with Cysti-
cerci so much as to the actual number of Cysticerci developed
in infected animals. I have frequently pointed out the inad-
visability of condemning and burying the carcases of measly
oxen, whether there be few or many Cysticerci present, and I
have stated, on trustworthy evidence, that even the presence of
a few Cysticerci is deemed by some inspectors a sufficient reason
for rejecting the entire animal. Such a waste should never be
allowed. In regard to the numbers of ox-measles present in
particular instances, I have elsewhere adduced some remarkable
facts communicated to me by Dr Joseph Fleming, of the
Indian Army Medical Staff. None of my experimental
animals, though fed with scores of ripe proglottides, yielded
such an abundance of Cysticerci as Dr Fleming encountered in
Punjab cattle. In one pound weight of the psoas muscles
Fleming counted no less than 300 Cysticerci.-" From this it
follows that the flesh of a largely infested animal is capable,
under the circumstances of ration distribution and imperfect
cooking, of originating numerous tapeworms.

Not many years back the leading medical journal of this
country challenged me to produce evidence as to the injurious-
ness of beef and mutton from Cysticerci. The writer stated in
his article that I had " failed to produce a single specimen of
beef or mutton measles " which had not resulted from experi-
ments conducted " at the Royal Veterinary College j " and he

rARASITES OP MAN

said, further, u that butchers, fieshers, and veterinarians were
practically right in refusing to adopt the opinion of Dr Cobbold,
that measled beef or mutton is produced to any great extent "
independently. How palpably I endured a species of unjust
reproach for being somewhat in advance of the knowledge
current at the time may be gathered from the voluminous
evidence which has since cropped up from various parts of the
world. It was, indeed, mainly through experiments conducted
at the Royal Veterinary College, and reported in the ' Lancet/
that professional men in India first became acquainted with the
possibility of finding Cysticerci in beef.

The statements of Dr Joseph Fleming, who was one of the
foremost in discovering cystic disease in cattle, have since
received abundant confirmation. The Indian Government
Reports given in the February issue of the c Madras Monthly
Journal of Medical Science ' for 1873 are especially instructive.
Referring to the prevalence of Cysticercus in the ration beef at
Jullundur, in the Punjab, the Inspector General (India Medical
Department) reports as follows :

" Cysticercus was first noticed here in the beef tendered at
the Royal Artillery ration stand in May, 1868. For some two
years previous to this date condemnations of cyst-infected meat
had been frequent at Peshawur, Rawul Pindee, Meean Meer
and several other stations in the upper part of the Punjab, and
here I had often detected the parasite in meat exposed for sale
in the bazaars, but no trace of it had been observed in the
Commissariat beef, either by myself or any other medical officer
who had preceded me.

" From May, 1868, to November, 1869, f cyst ' was more or
less frequently found both at the Artillery and 92nd High-
landers' ration stands ; but since the latter date it has almost
entirely disappeared.

" The following table shows the quantity of meat destroyed
on this account during 1868 and 1869 :

Years.

Months.

Number of cattle
infected.

Weight of meat
destroyed.

*•

1868

May
June

September ...
October
November ...
December ...

4
1
1
10
14
12

412 lbs.
77 „

130 „
1,763 „
2,010 „
1,785 „

CESTODA

Years.

Mouths.

Xumbcr of cattle
infected.

Weight of meat
destroyed.

1869

January

February

March

April

May

June

July

August
September ...
October

21
16
14
5
2
1
1

Q
O

2
4

4,062 lbs.
2,341 „
2,209 „
856 „
220 „
122 „
194 „

218 „
615 „

Total

111

17,478 lbs.

"The whole of this meat was otherwise well fed and of
excellent quality. The waste of so much good food led me to
make inquiries ; 1st, as to the sources from which the cattle
obtained the Taenia ova, and the best means for preventing their
infection ; and 2ndly, as to whether or not any evil results fol-
lowed the consumption of this meat when properly cooked.

" From information obtained from the Commissariat Officer
I found — 1st. That the infected cattle had been purchased by
native dealers from various parts of the district, not from any
particular locality. 2ndly. That when brought in they were
lean, and on an average required from two to three months'
feeding at the Commissariat cattle yards before they were fit
for the shambles. 3rdly. That their food consisted of the grass
they could pick up on the grazing grounds of cantonments,
supplemented by such an allowance of grain and bhoosd as their
condition required.

" They were supposed to be watered at a trough with water
drawn from a well, but on closely inquiring as to this, it tran-
spired that they very frequently were taken to a large dirty
tank near the yard for their water. The question which
occurred to me was, were the cattle infected before their pur-
chase by the Commissariat, or was there anything in their
feeding to account for it after purchase ? I am inclined to the
latter opinion for several reasons, thus : — In the large number
of the diseased cattle, the Cysticerci were of remarkably small
size ; many of them having no capsules, except such as were
formed by the surrounding structures, and not being more
than i to £ of an inch in diameter. Although the dry food
given to the cattle was doubtless good, still much of the water
they got during 1808 was probably filthy. The tank pro-

FARASITES OF MAN

viously referred to was situated close to the huts of the camel
drivers. These men are all Mussulmans from Cabul, Peshawur,
or thereabouts, and many of them are infected with Taenia
mediocanellata. Human filth was often to be seen on the
banks of the tank, and microscopic examination of mud and
stagnant water taken from the margin exhibited Taenia ova.

" The conditions above shown must have been eminently
favorable to keepiug up a constant supply of ova, and the fact
that Cysticercus entirely disappeared from amongst the cattle a
few months after means had been taken to secure them a good
supply of well water, seems to confirm the view that this tank
must have been the source of a large amount of, if not all, the
infection.

" It has been suggested that Cysticercus can be detected
before the animal is killed by an examination of the tongue.
In exceptionably severe instances this is probably correct, bnt
then it would be equally observable in some other parts of the
body. Major Biggs, Commissariat Officer here, tells me of an
animal he saw at Rawul Pindee, in which immense clusters of
cysts could be felt at the root of the tongue and under the
skin in several parts. After examining a very large number of
tongues of ' cysted ' animals, my experience is that it is found in
the soft muscles and cellular tissues at the root of the tongue,
perhaps more frequently than anywhere else ; but I have never
seen a case in which there was a chance of detecting it before
death.

« The most common situations in which it has occurred in
the ration meat have been the gluteal, psoas, and lumbar
regions. In many instances only from, one to ten cysts have
been found on cutting the carcase into small pieces, and I have
no doubt that it often passed without detection.

" During 1868 and 1869 I from time to time obtained pieces
of beef badly infected with Cysticercus, and made some experi-
ments as to the results of its consumption under different con-
ditions.

"After explaining to them the possible consequences of
eating it, a buttock of beef studded with Cysticercus was given
to three natives of low caste. They all declared that they were
free from Taenia, or, to use their own term, "Kadhu dana."
The meat they cooked in their own way. These men were
under my observation for some six months. Two of them had
no symptom of Taenia, but the third, who was a low- class

CESTODA

Mahommedan syce, and had probably eaten the meat in a very
raw state, developed a Taenia rnediocanellata in about three
months.

" My own sweeper ate this cyst-infected beef regularly two
or three times a week for some months. He cooked it well
generally as an ordinary stew, and has never shown a sign of
having tapeworm.

" Into the food of a boy of low Hindoo caste, but who had
never eaten beef, two scolices of Cysticercus were surreptitiously
introduced, the result being that, between three or four months
afterwards, he applied for some tapeworm medicine."

[The two successful experiments here reported are evidently
the same as those that I have referred to (p. 72) as having
been performed by Dr Oliver, of the Royal Artillery, stationed
at Jullundur. The report continues as follows :]

" Taenia rnediocanellata is very common amongst the Mus-
sulman population of the Punjab, and from reliable sources I
am informed that the lower classes amongst them are in the
regular habit of eating half-cooked beef ; indeed, prefer it so,
and it is amongst these people that tapeworm is so prevalent.

" But it is not only thorough cooking that is required to
guard soldiers in India from the ill effects of eating measly
meat ; there is want of cleanliness in the general arrange-
ments Of the kitchens and serving of meals, which must offer
great facilities for the introduction into the food of Cysti-
cercus.

" Barrack cooks, unless constantly looked after, are utterly
careless as to the "washing of chopping blocks, tables, dishes,
&c. The dish or pot cover on which the meat is placed when
raw is often used without washing for serving the piece up
for dinner, and I have myself picked up a Cysticercus from the
table on which a cook was preparing food. The dangers too
of the parasite being conveyed by the cook's unwashed hands
to the plates in which meals are served, and the common
practice of using the same knife for cutting up meat, and
afterwards, without washing it, for other culinary purposes,
must not be overlooked. With good selection and careful
feeding there seems to be every probability that Cysticercus
would soon almost or completely disappear from our Commis-
sariat cattle. If they were entirely stall-fed and watered from
wells there could scarcely be a possibility of infection after
their purchase.

PARASITES OP MAN

" Perhaps with the trench system of conservancy, which
will necessitate the growing up crops, a sufficient quantity of
root and other green produce may be obtained from canton-
ment lands set apart for this purpose, to supply green fodder
for the cattle."

The important question as to whether the presence of cysts
detected at the root of the tongue could be made available for
the purposes of diagnosis was made the subject of special
report through the agency of executive Commissariat officers,
and they testified to its practical valuelessness in the following
terms :

" Jullundur. — .No appearance of cyst has been found at the
root of the tongues of any of the cattle. A medical officer was
asked for assistance in making search for the cysts, but he also
found none.

" Raivul Pindee. — It is utterly impossible to discriminate
before slaughter, from any outward symptoms, cattle that are
cyst infected.

" Every endeavour has been made to discover by close and
careful scrutiny before slaughter the cyst-infected cattle, but
the result has been in no way satisfactory.

" SeaUwte. — All endeavours to discover any symptoms of the
infection by examination of their tongues, while the animals
were living, have been unsuccessful.

" Mooltan. — The mouth and tongue of a large number of
living cattle have been examined before slaughter, but in no
single instance has the infection been so detected. >

" Dr. Ross's plan of examining the tongues of all animals at
time of purchase is not feasible, as they are usually very wild
and frightened, and often dangerous to approach.

" Peshawur. — In probably 99 cases out of 100 it is utterly
impossible to discover cyst infection in cattle previous to
slaughter by examination of their tongues. In only one
instance has it been so discovered, and that was from the
animal's having a number of small lumps over the body which
were also apparent on the back part of the tongue. When the
tongue is infected the ' cyst ' lies so far at the very root of it
that it cannot be seen in the live animal."

From Mooltan a specially interesting report was made by Dr
Alexander Neill, who says : —

" I have carefully examined the mouth and tongue of a large
number of living cattle, and of those slaughtered for issue as

CESTODA

rations, and in no single instance did I find such cysts. These
cattle were healthy.

" In a case that died, and in which cysts existed, I could
discover nothing abnormal in or under the tongue.

" If such * cysts ' exist, or if such enlargements of the sub-
lingual glands are found, I argue that they are not a diagnostic
sign of what is termed t cyst infection/ or more correctly ' Cys-
tica reus horns j for in the recent outbreak of cattle disease in
England, one most prominent symptom of that disease was a
bunch of grape-like swelling under the tongue, which in advanced
cases suppurated, and to a casual observer would have been
called cysts or 'bags of matter/

" If such swellings are found in a bullock that is sick, it is
merely symptomatic of an inflamed condition of the whole
mucous surface of the intestinal canal, and not of any localised
disease, such as Cysticercus, the above-mentioned swellings
being merely inflamed sublingual glands.

" In the pig the diagnostic sign of swellings of the glands or
' cyst ' under the tongue is not found in ' Cysticercus/ and the
disease called ' measles ' is not ' Cysticercus/ but a mere super-
ficial inflammation of the skin and a symptom of fever. 1 Cys-
ticercus ceMulosus,' as its name shows, infects the cellular
tissue only of the pig, and cannot be discovered in life by any
abnormal condition of skin.

" In ' measles ' these swellings are found, because intestinal
mucous membrane sympathises with eruption on the skin and
are then merely inflamed glands, not cysts."

Dr Neill concludes his report by remarking that the larvae of
the beef tapeworm can " only arrive at maturity in the mucous
membrane of horned cattle," and not in the cellular tissue.
This is an error on Dr Neill' s part ; but in adducing these
instructive extracts from the Government Reports my chief
object has been to show the prevalence of Cysticercus in the
North-West Provinces of the Indian Peninsula. I may say
that a large proportion of my tapeworm-infected patients have
been officers from the Punjab, and one of these victims told me
ijthat when he superintended the serving out of rations to the
troops, " he (and those who acted with him) sent the meat away
Ito be burnt, even when they only detected a single cyst in any
igiven carcase." It is needless to remark that such a waste of
Ivaluable food is altogether reprehensible.

Some people, including not a few of the profession, make

PARASITES OF MAN

light of the occurrence of tapeworm, and I have seen many
patients who had been told by their usual medical advisers
that the presence of the worms was of little consequence.
To account for this wide-spread error there is some basis in the
fact that by far the majority of infested persons suffer only the
trifling inconvenience arising from the passage per anum of the
proglottides ; moreover, the less civilised the tapeworm-bearers
happen to be, the less are they likely to suffer. The recorded
experience of Kaschin, before referred to, where 500 hospital
patients, in the Baikal district, had tapeworm, although
all of them were being treated for other disorders, affords
another argument tending to the same conclusion. On the
other hand, amongst Europeans only a small percentage of
tapeworm-patients suffer severely. But without trenching
upon the symptomatology and prognosis of tapeworm disease,
I may remark that I have (in my Manual) summarised the
the whole facts of cysticercal prevalence within the compass of
two brief propositions: — 1. The prevalence or the rarity of
Oysticerci in cattle in any given country must be determined
primarily by the habits of the people ; for since the beef measle
caxi only result from the ingestion by the ox of the eggs of the
Tania mediocanellata, it is clear that the degree of infection of
cattle will correspond with the facilities offered by egg-disper-
sion. 2. It may be affirmed that the frequency of this particular
species of tapeworm amongst the people occupying any given
area will bear a strict relation to the amount of underdone
measly beef consumed by the inhabitants.

Another question, and one of great interest to sanitary
science, is that which I have raised in reference to the period
that nature requires for the destruction of the Oysticerci, or, in
other words, for the performance of a natural cure by calcareous
degeneration of the parasites. I have shown that all kinds of
tapeworm larvae (measles, bladder-worms, ccenuri, and so forth)
have a natural life-epoch assigned to them, and in one of my
experiments on a Dutch heifer or young cow I demonstrated
that a period of ten months was more than sufficient to ensure
the perfect destruction of the Oysticerci of cattle. Moreover,
this law or process of natural cure is not limited to cestode
parasites, but affects all other kinds of internal parasites in one
or other of their juvenile stages of growth. In the flesh of my
experimental animal I estimated that there were not less than
] 2,000 of these degenerated Oysticerci. This positive contribu-

CESTODA

tion to our knowledge of the limits assigned by nature to the
epoch of larval activity is not merely one of abstract scientific
interest, but it has important practical bearings, inasmuch as it
points out in what way an entire herd of cattle (known to
be measled by the post-mortem examination of one animal
previously selected for the purpose, or for that matter, by
the rather barbarous act of excising and examining a frag-
ment of the muscle of a living one) may be freed of its
parasitic guests ; and it also shows how all risk of propagating
tapeworm, apart from the question of subjecting the flesh
to a certain temperature, may be effectually prevented. The
stockowner has but to remove his animals for six or eight months
to localities where no fresh infection can occur, when, at the
expiration of the time mentioned, all those Cysticerci that
existed in the beasts at the time of the transfer will have perished.
The flesh of the animals may then be eaten with impunity,
whether well cooked or raw. This is an important teaching
deducible from experimental inquiry, and I am rather surprised
that it has hitherto escaped the notice of persons who, though
they affect to ignore the value of scientific researches, are
particularly anxious to parade their practical knowledge, which,
unhappily, too often proves a mere cloak for ignorance.

The memoir by Giacomini already quoted (p. 65) affords inter-
esting details respecting a case in which there was a most un-
usual degree of infection of the human body by Cysticerci. Dr
Giacomini instituted a searching comparison between the human
measles procured by himself and those of the pig sent to him
by Professor Perroncito. In the human Cysticerci he noticed a
greater adherence of the capsule to the enclosed measle, and
he also observed that while the human measle-heads either
displayed thirty-two, or in some few cases thirty-four hooks, in
two differently sized circles of fifteen or sixteen each, the pig-
measles, on the other hand, carried only twenty-four hooks to
the double circle of equal circumference ; consequently the hooks
appeared to be more crowded together in the human parasite.
This fact, Giacomini remarks, does not of itself constitute
an essential specific difference, since variations of the kind not
unfrequently occur in Cysticerci occupying one and the same
host. Even the beef-measle is not necessarily confined to one
species of host, since Zenker has succeeded in rearing it in a
goat.

Although the substance of the above-recorded conclusions

PARASITES OF MAN

was originally communicated by me, anonymously, to a pro-
fessional periodical, I have considered this work a suitable
medium for a fuller discussion of the subject. Its importance
in relation to the public health and the supply of meat-food has
not received the attention it deserves.

Bibliogeaphy (No. 13.) — Balert, B., ' Die Bandwiirmer/ &c.
(pamphlet), 1877. — Bertolus, G., ' Diss, sur les metamorph. des
cestoides/ Montpellier, 1856.— Oobbold, T. S., " On the Pro-
duction of the so-called ' Acute Cestode Tuberculosis ' by the
Administration of the Proglottides of Taenia mediocanellata "
(with Mr Simonds), in ' Proc. of the Royal Society ' for May
4th, 1865; repr. in the ' Veterinarian 3 for 1865, p. 513. — Idem,
" Experimental Investigations with Cestoid Entozoa," in ' Linn.
Soc. Journ./. vol. ix, p. 170 ; also for July, 1865, p. 141. — Idem,
" On Beef, Pork, and Mutton, in relation to Tapeworms," in
'Brit. Assoc. Rep/ for 1865, p. 102, and in 'Appendix to
Treatise on Tapeworms and Threadworms/ 1st Edit., 1866, p.
73; also in 'Med. Times and Gaz/ for Sept. 23rd, 1865, p.
343. — Idem, " Remarks on Entozoa," in ' Brit. Assoc. Rep/
for 1865, p. 102 ; also on " Cystic Entozoa from Veal and
Mutton," in the 'Path. Soc. Trans/ for 1866, vol. xvii, p. 462.
— Idem, " Entozoa found in a Westphalian Ham ;33 report in
' Athengeum ' for March 27th, 1869, p. 442 ; also in 'Brit. Med.
Journ/ for March 20th, 1869. — Idem, "Note on Beef Measles
from a Cow," in ' Path. Soc. Trans./ vol. xvii, p. 463, 1866 ;
also in the ' Lancet 3 for Feb. and August, 1865, p. 249. — Idem,
'Entozoa/ &c, p. 235 et seq., 1864; and in ' Supp./ sections
iii, iv, v, 1869. — Idem, ' Tapeworms/ 3rd Edit, (with 100 cases),
1875, p. 11. — Idem, 'Manual of the Internal Par. of Domesti-
cated Animals/ chap, iii to vi, 1874. — Idem, ' Worms/ Lec-
tures i to xi, 1872. — Idem, " On the Parasites of our Food-pro-
ducing Ruminants (Cantor Lectures)," in the 'Journ. of the Soc.
of Arts/ 1871. — Idem, " On the Entozoa of Abyssinia" (Lec-
ture), in 'Lancet/ 1867. — Idem, "Remarks on Eighty Cases of
Tapeworm," 'Lancet/ June, 1874. — Idem,, "Revised List of
Entozoa, with notes and references (the beef tapeworm, No. 15,
and the beef measle, No. 25)," in the ' Veterinarian/ Dec, 1874,
and Feb., 1875. — Idem (anonymously), " Cysticerci, being a
review of the writings of Pellizzari, Tommasi, Perroncito,
Lewis, Giacomini, &c," contributed to the ' Loud. Med. Record/
1874, p. 642 et seq. ; repr. in the 'Veterinarian/ Jan., 1875. —
Idem, " Notice of a Discussion by Paul, Martineau, Crequy,

CESTODA

Delioux de Savignac, Trasbot, and others, respecting the Source
and Treatment of Tapeworm/' ' Lond. Med. Rec./ July, 1874,
p. 472. — Idem, " Review of the Writings of Oliver, Fleming,
Hewlett, Lewis, and others, on the Cystic Disease of Animals/'
'Lond. Med. Rec./ June, 1873, p. 339. — Idem, "Further Ex-
perimental Researches with the Eggs of the Beef Tapeworm/'
the ' Veterinarian/ Aug., 1875. — Idem, " Remarks on Perron-
cito's Researches/' the ( Veterinarian/ Dec, 1877. — Dardel, A.,
" Sulla frequenza della Tenia in Savoia," ' Giorn. d'Accad. di
Med./ 1868.— Davaine, G, ' Traite ' (1. c. Bibl. No. 1), I860.—
Idem, " Les Cestoides," in ' Diet. Encyclopedique des Sci.
Med./ 1875.— Fleming, J., 'Indian Med. Gaz./ 1869.-^0^, H.
G. A. L., ' De Lintworm en het middel om hem mit te drijven/
Utrecht, 1873. — Fritsch, G., " Zur differentiellen Diagnose von
T. solium and T. mediocanellata," ' Berliner Klinische Wochen-
schrift/ 1874. — Gamgee, J., " Entozoa in Veal and Beef " (Letter
on), ' Lancet/ 1865. — Giacomini, G, ' Sul Gyst. cell, hondnis e
sull Taenia med., contrib. alio studio dei Oestoidi Parrassiti dell'
TJomo/ Torino, 1874. — Heller, A., " Darmschmarotzer," in
von Ziemssen's ' Handbuch der speciellen Pathol, und Therapie/
s. 598 et seq., 1876. — Hewlett, 'Health Officer's Report/ Bom-
bay, 1870.— Krabbe, H, 'Beretning om 100 Tilfaslde af Bcen-
dellorm hos Menesket iagttagne her i Landet (Aftryk af Uges-
krift for Lasger)/ 1869. — Kiichenmeister, F., ' Ueber Cestoden
im Allgeraeinen und die des Menschen insbesondere, hauptsahlich
mitBerucksichtigungihrerEntwickelungsgeschichte, geographis-
chen Verbreitung, Prophylaxe und Abtreibung ; specieller Theil.
Zittau/ 1853.— Idem, ' Parasiten ' (1. c. Bibl. No. 1), 1855, Eng.
Edit., London, 1857. — Laboulbene, A., "Sur les Tsenias," 'Mem.
de la Soc. Med. des Hopit./ 1876.— Idem, 'Anat. Pathologique/
1879, p. 962.— Letheby, " On Diseased Meat/" Med. Times and
Gaz./ 1867.— Leuckart, R., ' Die Menschl. Par./ Bd. i, s. 285
and s. 747, 1864.— Levi, "Della freq. della tenia," &c, ' Giorn.
Veneto di Scienz. Med./ 1874. — Lewis, T. K., " A Report on the
Bladder Worms found in Beef and Pork" ('App. B. to 8th
Ann. Rep. of the Sanit. Commiss. with the Gov. of India '),
Calcutta, 1872. — Masse, F. et Pourquier, P., "Le 'Taenia inerme
et la lardrerie du Bceuf, Nouvelles Experiences," &c, in 'Mont-
pellier Med. Journ. Mens, de Med./ p. 220, 1876.— Mosler,
' Helminthogische studien und Beobachtungen/ Berlin, 1864.
Neill, A., " Letter, forming the fifth of a series of important
articles on Cyst-infected Cattle, and on the prevalence of Cys-

PARASITES OP MAN

ticercus in Beef/' reported by the Inspector General (I. M. D.),"
in the ' Madras Monthly Journ. of Med. Sci./ Feb., 1873; repr.
in the ' Veterinarian/ July, 1873. — Nitsche, H., " Untersuch-
ungen ueber den Bau der Tgenien," ' Sieb. und Koll. Zeitschrift/
1873. — Oliver, "Rejections of Ration Beef on account of Cystic
Disease " (1. c. supra), ' 7th Rep. of the Commiss./ p. 82, Cal-
cutta, 1871. — Perroncito E., " Delia panicatnra negli animali,"
' Annali della R. Accad. d'Agricolt. di Torino/ vol. xv, 1872. —
Idem, " Sulla morte del Cyst. cell, delle carni del majale ibid.,
1872. — Idem, "Ueber die Lebenszahigkeit des Cyst. cell, und
anderer Eingeweidewiirmer," ' Zeitsch. f . prakt. Veter.-Wissen-
schaften/ Bern, 1876. — Idem, ' Della Grandine o Panicatura
nell' Uomo e negli animali/ Torino, 1877. — Idem, "Esperi-
menti sulla produzione del cisticerco nelli carni del bovini, coll'
amministrazione di anelli della tania med. dell' uomo," ' Lo
Studente Yet./ Parma, 1876, p. 146. — Idem, "Sulla tenacita,"
&c, ibid., 1877, p. 194. — Idem, " Esperimenti sulla prod, del
Cyst, della T. med. nelle carni dei Yitelli," ' Estr. della Annali
d. R. Accad. d'Agric. di Torino/ vol. xx, 1877 — Idem, " On the
Tenacity of Life of the Helminths, and their corresponding
Larval Forms in Man and Animals," the ' Veterinarian/ July,
1877, p. 457. — Idem (with similar title, including notice of
experiments), the ' Veterinarian/ Dec, 1877; partly from 'Osser-
vatore Gaz. d. Cliniche di Torino/ and from ' Archivvo per
le Sci. Med./ vol. i, 1877.— Idem, "On the Tenacity of Life of
the Cysticercus in the flesh of Oxen, and on the rapid develop-
ment of the corresponding T. mediocanellata in the Human
Body," the ' Veterinarian/ Dec, 1877, p. 817. — Probstmayr,
' Jahrb. der Munch ener Thierarzneischule/ 1869. — Pochard,
" Note sur la frequence du Taenia mediocanellata en Syrie, et
sur la presence du cysticerque qui lui donne naissance, dans la
chaire musculaire des boeufs de ce pays," in ' Bulletin de l'Acad.
de Med./ 1877, torn, vi, p. 998.— Thudichum, J. W. L., " On
the Parasitic Diseases of Quadrupeds used as Food," ' Privy
Council Med. Officer's Rep.' 1865. — 8ommer, F., Ueber den
Bau und die Entwickelung der Geschlechtsorgane, von Tcenia
mediocanellata und T. solium/' in ' Siebold and Koll. Zeit-
schrift/ Bd. xxiv, s. 499, 1874.— Oyr, "Deux Experi-
ences," &c, ' Journ. de l'Anatomie, de Robin/ p. 504 ; and in
' Lond. Med. Rec./ by Higgs, vol. i, 582, 1873.— Tommasi, T.,
' Appendice (to Cobbold's) Parasiti Interni degli Animali Do-
mestice/ p. 161, Firenze, 1874. — Van Beneden, P. J., "Icono-

CESTODA

graphie des Helminthes ou des vers parasites de l'homrne "
(Vers Cestoides, pi. ii), Louvain, I860.— Welch, F. H., "Obser-
vations on the Anatomy of Tania medio canellata," ' Quart.
Journ. of Microsc. Science/ vol. xv, 1875. — Zenker, in ' S. B.
Soc./ Brlang. iv, s. 71. — Zum, ' Zoopathologisclie und physiol.
Untersuchungen/ 1872.

Tama solium, Linneus. — This cestode was formerly known
as the common tapeworm, but in England it is of far less fre-
quent occurrence than the beef tapeworm. In contradistinc-
tion it is best to speak of it as the pork tapeworm. Though
only one specimen is usually present, the bearer may entertain
several worms of this species at one and the same time. The
parasite has been known to science from the earliest times,
though possibly not earlier than the measles, or Cysticerci, from
which it originates. Hippocrates, Pliny, and Aristotle describe
the full-grown worm ; and, in regard to the larvse, some have
gone so far as to express their belief that the prohibition of
swine's flesh as food amongst the Jews and other Oriental
people, was dictated by sanitary considerations. Weinland has
suggested that the Mosaic commandment not to eat pork may
have originated in an old popular notion " of the fact that tape-
worm sometimes comes from this food." Weinland' s hypo-
thesis is probably correct, for if one supposes Moses to have
been supernaturally informed that pork would produce tape-
worm disease, one naturally asks why veal and beef should not
also have been prohibited, seeing that these meats also fre-
quently harbour tapeworm larvae.

A perfect pork tapeworm presents itself to the eye of the
observer as a long, soft, white, jointed strobile, which, when
alive, elongates and contracts itself with facility. Though
commonly spoken of as a single creature, it is a compound of
many individuals. These are variously called " cucurbitini,"
"zooids," "proglottides/' "segments/' "links," or "joints."
When fully grown the segments are capable of detaching them-
selves and of enjoying a free and independent existence. Very
annoying it is to the human bearer to be continually reminded
of his unwelcome " guests " as they seek to quit his interior.

The head of Tcenia solium is seldom seen in anatomical
museums, although the evacuation of pork tapeworms is not of
rare occurrence. Placed under the microscope, the head dis-
plays a quantity of dark, almost black, pigment granules,
which are abundant at the base of the rostellum and in the

PARASITES OP MAN

neighbourhood of the hook-fangs. They are equally present

and abundant in the pork ineasle proper, and in measles derived

from the human subject. The cephalic hooks of this cestode

are comparatively large, those of the greater circle individually

measuring y^/', whilst the smaller hooks have a length of about
i "

220 '

Fig. 23. — Head of Tarda, solium. Highly magnified. After Van Bcneden.

The male reproductive organ consists of a number of small
vesicles or sacs, in which filiform spermatozoa have been
detected, these latter, when ripe, being conducted by a vas
deferens into a seminal pouch, from which a canal passes
laterally into the penis ; the latter organ, in its retracted con-
dition, being lodged within a flask-shaped sheath or cirrhus-
pouch. The female organs are somewhat more complicated.
They consist of two masses of vitelligene glands occupying a
limited space, a small ovarium, a centrally -placed and largely-
developed branched uterus, canals of outlet leading from all
these organs, and enlargements of the main passages to form
internal seminal reservoirs ; also, a vaginal canal, which is
widened at its termination to form a receptaculum for the
curved penis.

CESTODA.

In addition to the above-named structures, the entire series
of joints from the head downwards are traversed by a set of
vascular canals, which are doubled in the region of the head.
These form the so-called aquiferous system. There are two
main channels, one passing down on either side of the worm,
both being connected by transverse vessels, which occur singly
at one end of every joint.

The eggs in their mature condition are globular, and contain
a six-hooked embryo. They present an average diameter of

of an inch, the shell itself measuring about in thickness.
In 1856 I observed that many of the eggs, whilst still within
the uterine branches, displayed an outer envelope, very delicato
in structure and totally dissimilar from the egg-shell proper.
This has since been more accurately described by Weinland,
Yan Beneden, and Leuckart. The outer membrane, according
to the last-named authority, constitutes the primitive yolk-
membrane, within which a part of the yolk-contents separates
to form the true egg and embryo by a process of daughter-cell
formation. The remaining part of the yolk forms a granular
mass, being probably concerned in the formation of the true
chitinous shell. The true shell displays a series of radiating
and circular lines ; the former, however, are more conspicuous
than the latter, being due, according to Leuckart, to the
presence of a series of fine rod-like chitinous elements, which
are formed on the external surface of the original true shell-
membrane. The enclosed embryo is furnished with six boring
spines, arranged in three pairs, its granular body being invested
by an extremely delicate skin-membrane, which is separated
from the inner surface of the shell by a clear transparent fluid.
The embryo measures in diameter.

The scolex or higher larval stage of growth forms the well-
known pork measle or Cysticercus (tela) celluloses of authors.
The smallest measles found by Leuckart measured i" in length.
They were obtained from the brain, liver, and intermuscular
substance of a pig fed with proglottides about thirty days
previously. Only those specimens, however, occurring in the
liver at this early period displayed an outer membrane proper
to the worm itself, the others being simply invested with
capsules formed out of the connective tissues of the host.
Many measle-masses in the same host were much larger,
presenting an average diameter of . The smallest already
displayed a smooth, transparent, homogeneous, outer, cuticular

PARASITES OP MAN

membrane, overlying a double, finely-granular corium, the
latter being traversed by a branched system of aquiferous
vessels. These vessels proceed from a central spot, which
marks the position of the so-called head-cone, or receptaculum
capitis. It is, in fact, the first well-marked indication of that
flask-shaped capsule within which the head, neck, and body of
the Cysticercus is formed, and which Goeze long ago very aptly
compared to a lantern. As growth proceeds, a central granular
mass forms the true foundation of the head, its upper or stalk-
like extension becoming the future neck and body. Further
changes result in the evolution of the internal water- vascular
system, the calcareous corpuscles, the marginal transverse fold-
ings of the body, the four suckers, the rostellum, and, in
particular, the double coronet of hooks. All these metamor-
phoses were minutely followed and described by Leuckart, who
found the development of the larva to be completed within the
space of ten weeks.

As regards the injurious effects of this parasite upon man,
it may be said to act prejudicially in three separate ways. I
have remarked in my ' Entozoa/ that this parasite may cause
disease and death both by its action in the larval and adult
states. It may likewise injure us by rendering the flesh of
swine unwholesome.

When one or more sexually- mature tapeworms have developed
themselves within the human intestine, they are apt to give rise to
a variety of unpleasant symptoms, more or less marked accord-
ing to the habit or irritability of the patient. According to
Davaine (p. 103 of his ' Traite '') the principal features are
" vertigo, noises in the ears, impairment of sight, itching of the
nose and anus, salivation, dyspepsia and loss of appetite, colic,
pains over the epigastrium and in different parts of the abdo-
men, palpitation, syncope, the sensation of weight in the
abdomen, pains and lassitude in the limbs, and emaciation."
In ordinary cases there is always more or less anxiety and
restlessness ; but in severe cases the sympathetic symptoms
are very strongly marked, showing themselves in hysterical fits,
chorea, epilepsy, and epileptiform seizures, attended by more
or less alarming convulsions.

Amongst some of the more interesting and remarkable cases
recorded in our English journals, I may instance that of Mr
Hutchings, where a complete cure followed the evacuation of
the worm which had produced convulsions. Mr Tuffnell records

CESTODA

a case where irritability of the bladder and stricture of the
urethra were entirely dependent on tapeworm, as proved by
the subsequent recovery. At a meeting of the Pathological
Society, in 1853, Dr Winslow mentioned his experience of
three or four cases of mania arising from tapeworm ; whilst on
the same occasion Drs Ryan and Davey each recorded a similar
instance. A case has also been previously published by Mr W.
Wood. At a meeting of the London Medical Society, held on
the 10th of April, 1837, Dr Theophilus Thomson (during an
interesting discussion on this subject) stated the facts of a
case where the presence of tapeworm had given rise to a
tumultuous action of the heart, this symptom entirely disappear-
ing after evacuation of the worm. Our journals likewise (anony-
mously) record a considerable number of cases from foreign
sources. Thus, in the ' London Medical Gazette ' for 1840, there
is the case of a lady, aged thirty- seven, who had convulsions
attended with a complete loss of consciousness, the separate fits
lasting an hour at a time. The passage of the worms effected
a complete cure. In the same journal for 1838, there is also
the case of a younger lady (aged twenty- seven) suffering from
epilepsy, in whom a complete cure had been similarly brought
about ; here, however, in addition to a single specimen of the
Tania solium,, there were two lumbrici present. This journal
also gives Bttmiiller's case, where eighteen tapeworms were
the cause of hysteria ; and likewise the case published by
Steinbeck, where the symptoms presented an altogether peculiar
character. More precise references to some of the above cases
will be found in the ' Bibliography ' below ; and I may also
refer to my published lectures on Helminthology and especially
to my separate work on Tapeworms, where particulars of one
hundred cases are briefly recorded. These were all average
cases occurring to me whilst in private practice. Davaine's
book also abounds with remarkable cases.

Whilst the adult worm is capable of producing serious
and even fatal mischief to the bearer, the larvae or measles
much more frequently prove fatal. The Cysticerci may
develop themselves in almost any situation in the human body,
but they occur most commonly in the subcutaneous, areolar,
and intermuscular connective tissue ; next, most commonly in
the brain and eye, and lastly, in the substance of the heart and
other viscera of the trunk.

In my f Entozoa ' I have stated that probably not less than

PARASITES OP MAN

one hundred cases have been observed where death had resulted
from Cysticerci in the brain. Griesinger alone collected
between fifty and sixty such cases. Mental disturbance occa-
sioned by the presence of measles in the brain may occur with
or without epilepsy. When Griesinger states that " the epi-
lepsy from Oysticercus is in all respects like cerebral epilepsy
and the psychical disturbances have nothing characteristic
about them," he tacitly admits the impossibility of correct
diagnosis during life.

Since the publication of Griesinger's well-known memoir on
Cysticerci of the brain, many similar cases have appeared, and
amongst the more recent of these is one by Dr Fredet in which
the victim was a young man twenty-two years of age. Though
apparently in good health he fell dead in the street; the
fatal result being due to the presence of a Oysticercus within
the pons Varolii.

Many other cases of earlier date are especially noteworthy.
Thus Mr Toynbee recorded a case where an hydatid (which I take
to have been the Cysticercus celluloses) situated in the middle cere-
bral fossa beneath the dura mater, but in this instance death
ensued from other causes. Mr Ottley gives the case of a woman
aged forty, where an undoubted Cysticercus in the brain gave rise
to distressing fits, convulsions, and death. Then, again, there was
Dr Burton's workhouse patient, only twenty years of age, who
was found dead in bed, but who at the time of admission
merely complained of pain in the head. After death, four
hydatids (Cysticerci) were found in the tuber ancillare at the
summit of the spinal marrow. M. Bouvier's similar case is
also reported in our periodicals. Of instances where Cysti-
cerci occupied the cavity of the eye, we have one or two
cases by Mackenzie of Glasgow, one by Mr Rose of Swaffham,
and others by Windsor, Logan, and Estlin. Amongst the
more peculiar cases, I may mention that described by Dr
Greenhalgh in the ' Lancet ' (1848), where the Cysticercus
was lodged within the substance of the lip. Five similar cases
are likewise recorded by Heller of Stuttgard. Then there is
Dupuytren's case of a Cysticercus ensconced within the great
peroneus muscle ; and also Fournier's, where several of these
scolices were said to have been found in a boil. The so-called
Trachelocampylus, discovered by Fredault in the human brain,
was neither more nor less than a common Oysticercus cellulosa.

It is worthy of remark, as Griesinger has also observed, that

CESTODA

Fig 24 — Head of a Cysticcrcus removed from the
brain. Magu. 6 diam. with detached hooks. Original.

in cases where the Cysticerci have taken up their temporary-
residence in the brain, they are usually found, post mortem, in the
grey cortical or peripheral substance of the cerebrum. The
particulars of such a case are given in my ' Bntozoa ' where the
victim suffered from epi-
leptic fits due to the pre-
sence of numerous Cysti-
cerci (fig. 24). The patient
was under Mr Hulke's care.

As regards infection by
the adult worm it is not
alone sufficient that we
avoid underdone meat, as
brought to the dinner-table,
but we must be especially
careful to have our sausages
well cooked. Under ordinary circumstances, we are safe for the
following reasons : — No respectable butcher will knowingly
supply us with pork or with sausages which are measled. Even
in the case of underdone meats, in whatever way prepared, it is
usually only a small portion which is unaffected by cooking.
As we have seen a temperature of 140 Fahr. is sufficient to
kill the Cysticerci.

The successful rearing of pork measles by experimentation
with the eggs of T. solium has been accomplished by many
helminthologists, amongst whom may be particularised Van
Beneden, Leuckart, Kiichenmeister, Haubner, Gerlach, and
Baillet. The converse experiment of rearing the adult worm
from the Cysticercus was first successfully undertaken by
Kiichenmeister on a condemned criminal ; Leuckart, Humbert,
and others having repeated this method with more or less
success.

The dangers arising from infection by swallowing the larval
worms or six-hooked embryos are not easily avoided. Our
flesh, like pork, thus becomes measled, although certainly
not to the spawn-like extent so often seen in the lower
animals. A single measle is sufficient to prove fatal ; and
this humiliating contingency, moreover, is one which we can
never be absolutely certain of avoiding. We become the
" host " or beare>r of the measle by swallowing the fully-developed
eggs of the Tarda solium. This we may do directly by hand-
ling fresh tapeworms, whose eggs, being concealed under our

PARASITES OF MAN

nails or in our clothing, may subsequently be swallowed, and
develop within us accordingly. Even a thorough washing of
the hands will not ensure absolute security. In like manner,
those who partake of choice salads, prepared from the stores of
the market-gardener, run a certain amount of risk. The
vegetables may have been manured with night-soil containing
myriads of tapeworm eggs, or they may have been watered
with fluid filth into which the eggs were accidentally cast. In
such cases, one or more tapeworm ova will be transferred to the
digestive organs, unless the vegetables have been very carefully
cleansed. In the same way, one perceives how fallen fruits
all sorts of edible plants, as well as pond, canal, and even
river water procured from the neighbourhood of human habita-
tions, are liable to harbour embryos capable of gaining entrance
to the human body. One individual suffering from tapeworm
may infect a whole neighbourhood by rendering the swine
measly, these animals, in their turn, spreading the disease far
and wide. As already remarked, measles sometimes occur
in great numbers in different parts of the body. Among the
more remarkable cases of the multiple Cysticerci are those
recorded by Delore (1864) and Giacomini (1874). In M.
Delore's case, about 2000 were obtained post mortem. Of
these, 111 occurred in connection with the nervous centres,
eighty-four being in the cerebrum, twenty-two in the membranes
of the brain, four in the cerebellum, and one within the substance
of the medulla oblongata. Dr Knox published a less notable
instance in the ' Lancet ' (1838); and in the year 1857, Dr
Hodges, of Boston, U.S., published a case where the cysts,
which in size he compared to rice grains and coffee beans, were
felt subcutaneously. The coexistence of Taenia and Cysticerci in
the same individual has also recently been observed in France
(' Lond. Med. Rec./ 1875). Besides these, several remarkable
instances have lately been reported by Davy, Tartivel, and
others.

To the literature already quoted in connection with the beef
tapeworm the following may be added :

Bibliography (No. 14). — Aran, in c Archives Gen. de Mede-
cine/ 1841. — Baillet, " Helminthes," art. in f Bouley and
Reynal's Diet. Yeterin./ torn, viii, 1869. — Becoulet and Qiraud,
" On Cysticercus in the Brain," ' Bullet, de la Soc. Med. de
Gand/ 1872 ; and in ' Lond. Med. Rec./ Feb., 1873.— BirMt, J.,
Cases, 1 Guy's Hosp. Rep./ I860.— Bouchut, "Cyst, in the Brain,"

CESTODA

'Gaz. des Hop./ 1857, and 'Journ. fiir Kinderkrankheit./ 1859. —
Bouvi&Tj ' Bullet. del'Acad./ 1840. — Burton, in ' Med. Times and
Gaz.' (supposed hydatids), 1862. — Cobbold, "On Measly Meat and
Measles in Man/' the ' Veterinarian/ 1876. — Czermack, " Cysti-
cerci causing Insanity/'' Corresp. — Blatt, 1838. — Dalton, J. C,
"Cyst in the Scrotum/' 'New York Journ. of Med./ 1857.—
Davaine (see his ' Traite ' for many additional references ; p.
676). — Davy, R., " Cysticerci in the Muscles/' ' Eep. of Lond.
Med. Soc./ ' Lancet ' for Nov., 1876—Estling, "Cases of Cysti-
cercus," 'Lond. Med. Gaz./ 1838-39 — Fredet, " Cysticercus
in the pons Varolii," in the ' Lancet ' for June 23rd, 1877
(p. 925), from ' Giornale Veneto de Scienze.' — Fournier, ' Journ.
des Connois. Med. Chir./ 1840. — Oriesinger, " On Cysticerci of
the Brain/' from ' Med. Jahrb.' in ' Med.-Chir. Eeview/ 1863.—
Harley, J., " Cyst, in the Brain," ' Lancet/ 1867. — Hodges, B.M.,
" Specimens of Cyst, cell., felt as small tumours just beneath the
skin, varying in size from that of a grain of rice to that of
a coffee bean," ' Rep. of Boston Soc. for Med. Improvement/
in 'Brit. Med. and Surg. Journ./ 1857.— Hogg, J., " Obs. on
Cysticercus," in his 'Manual of Ophth. Surgery/ 3rd edit.,
1863. — Holler, A., " Cyst, cell., im Gehirne einer Geistes-
kranken/' ' Allgem. Wiener Med. Zeitung/ 1878. — Logan, JR.,
" Probable Cases of Cyst, cell.," removed by Robertson, 1 Ed.
Med. and Surg. Journ./ 1833. — Mackenzie, W., " Cyst in the
Eye," 'Lancet/ 1848, 'Lond. Med. Gaz./ 1833.— Mazotti, L.,
" Caso di numerosi cisticerchi del cervello e delle meningi,"
' Rivista Clin, di Bologna,' 1876. — Megnin, P., "La Ladrerie
du pore et le Tcenia solium," ' La France Medicale/ 1876. —
Putz, H., " Ueber die Lebenszahigkeit des Cysticercus cellu-
loses," &c, 'Zeitsch. f. pr. Vet. - Wissenschaften/ 1876. —
Rainey, G., " On the Structure, &c, of Cyst, cell.," ' Phil.
Trans./ 1857. — Rizzetti, G., " Rendiconto Statistico dell' ufficio
d'igiene di Torino per 1' Anno 1873.' — Rudall, J. T., " Cyst,
in the Brain," 'Australian Med. Journ./ 1859. — Tartivel, De A.,
" Cysticerques multiples dans le tissu cellulaire sous-cutane et
dans certain visceres," ' Rec. de Med. Vet./ 1876. — Von Grafe,
A., in 'Arch, fiir Ophthal./ 1857. — Wells, S., Bourman's
Case, ' Ophth. BLosp. Rep./ 1860. — Windsor, J., " Cyst, in
the Eye," 'Brit Med. Journ./ 1861.

Tcenia tenella, Cobbold. — I have long been acquainted with the
fact that there is a comparatively small human tapeworm which
cannot bo referred to either of the foregoing species. In

PARASITES OF MAN

the absence of experimental proof, I incline to the belief that
the worm in question owes its existence to measly mutton.
The sheep harbours an armed Cysticercus (0. ovis), which 1
regard as the scolex of Tarda tenella. The specific name
(tenelLi) was originally applied by Pruner to a cestode six feet
in length, which he found associated with a larger tapeworm.
This latter he called Taenia lata. Whilst Diesing has pro-
nounced Pruner' s Taenia lata to have been a T. mediocanellata,
I, on the other hand, consider Pruner' s T. tenella to have been
a T. solium. Mr J. 0. Mayrhofer has suggested its identity with
Bothriocephalic tropicus. When, some years back, I applied
the term T. tenella to a new tapeworm (of which I possess
several strobiles) I was quite unaware than any similar nomen-
clature had been adopted by Pruner. From the few facts
supplied by Pruner and Diesing, I cannot suppose that our
cestodes are identical. Unfortunately my specimens are im-
perfect, wanting the so-called head. It is not possible to
estimate the length of the worm accurately, but the perfect
strobile must measure several feet.

On one slide I have mounted nine mature proglottides of a
worm which I procured on the 15th Dec, 1875. The segments
measure, on the average, exactly in length, and only ^" in
breadth. The uterine rosettes are all full of eggs, and their
branches so crowded together that I am unable to ascertain their
average number. The segments are perfectly uniform in
character, their reproductive papillae alternating irregularly at
the margin.

In the autumn of 1872 I caused a lamb to be fed with the
proglottides of a tapeworm which I referred to this species.
The animal was slaughtered on the 22nd of January, 1873,
when the result was stated to have been negative. As I had
no opportunity of examining the carcase, I cannot feel quite
sure that there actually were no Cysticerci present. On several
occasions I have detected measles in the flesh of animals, when
none were supposed to be present by those who either assisted
me or were professional on-lookers. Assuming my Taenia tenella
to be derived from the sheep's Cysticercus, I. think it fitting to
describe the mutton measle in this place. Even if T. tenella
be not actually the adult representative of the mutton measle
{Cyst, ovis), it is quite certain that the scolex in question
gives rise to an armed tapeworm, and it is almost equally certain
that the adult armed cestode resides in man. In Pruner's case,

OESTODA

which is by no means unique, we have seen that two distinct
species of cestode may coexist in the human bearer. It is
quite possible that some one may yet have the good fortune to
detect the beef tapeworm, the pork tapeworm, and the mutton
tapeworm, all together in one and the same host.

On five separate occasions I have detected measles in " joints "
of otherwise excellent and healthy mutton brought to my own
table, and supplied by the family butcher. On several other
occasions I have had these parasites brought under my notice ;
nevertheless, many persons are either unaware of, or actually
deny, the existence of these ovine parasites. Thus, MM.
Masse and Pourquier, in the ' Montpellier Med. Joura.' for
Sept., 1876, make the following statement : " The sheep, not
being subject to measles, it seems to us natural to employ the
raw meat of that animal whenever it is required for nourishment
in the treatment of diarrhoea, in weaning children, in phthisis,
and for anaemics." Clearly, if MM. Masse and Pourquier
could have brought themselves to believe that English lite-
rature is worth consulting on such matters, they would not have
made this statement. Incidently they also observe, when
speaking of beef measles : — " Un fait que nous avons remarque
et que nous tenons a signaler, c'est que nous avons trouve des
cysticerques nageant librement dans l'eau ou nous avions plonge
de la viande infestee de ladrerie." Certainly this is a novel
experience. That measles should not only get out of their
cysts, but should have the power of " swimming freely" in the
water is a phenomenon which requires explanation. There must
have been some error of observation.

It was in the year 1865 that I discovered the mutton
measle (G. ovis, mihi) ; but I am not prepared to say that the
parasite had never been seen before, since it is alleged that a
two-headed Cysticercus was obtained by Fromage from the liver
of a sheep (as cited by Davaine). Be that as it may, my dis-
covery was announced in a communication made at the Bir-
mingham meeting of the British Association in the autumn of
1865, and subsequently at a meeting of the Pathological Society
of London, on the 3rd of April, 1866 (f Path. Trans./ vol. xviii,
p. 463). After these dates further announcements and veri-
fications appeared, amongst which I can only refer to my
remarks " On Beef, Pork, and Mutton, in relation to Tape-
worms," forming an appendix to the first edition of my work
Ion Tapeworms, 1866 ; to the "Kemarkson Cysticerci from

PARASITES OF MAN

Mutton/5 contained in the fourth chapter of the Supplement to my
introductory treatise on Entozoa, where a figure of the parasite
is given, 1869, p. 27; to Dr Maddox's paper "On an Entozoon
with Ova, found encysted in the Muscles of a Sheep/5 recorded
in ' Nature/ May 15th, 1873, p. 59; to the 'Monthly Micro-
scopical Journal/ June, 1873, p. 245; to my further com-
munications in the 'Lond. Med. Record,5 Aug. 6th, 1873; to my
' Manual/ 1874, pp. 74 and 105, Ital. edit. ' Nota Dell5 Autore/
p. 133 ; and especially to the article headed " The Mutton
Tapeworm/5 contained in the 3rd edit, of my little volume on
' Tapeworms/ p. 12, et seq., 1875.

In regard to the measle itself, I spoke of it as smaller than
the common pork measle. The head is ^" in breadth, and is
armed with a double crown of hooks, twenty-six in all, the larger
hooks each measuring -j^" in length. The suckers are four in
number, each having a breadth of ~". The neck and head are
abundantly supplied with calcareous corpuscles, being at the
same time marked by transverse rugaa. The data on which I
founded my brief description of the scolex were chiefly based on
the examination of a specimen which had been procured by Prof.
Heisch from the interior of a mutton chop. Subsequently much
fuller details of the structure of the scolex were supplied by the
illustrated memoir of Dr Maddox (above quoted) . This excellent
microscopist, however, announced the presence of immature ova
within the Oysticerci themselves. As the notion of the exist-
ence of eggs in larval cestodes was altogether at variance with
what we know of the phenomena of tapeworm life, I suggested
that the author might have mistaken the egg-shaped calcareous
corpuscles (which I found so abundant in my own specimens)
for the ova. In the interests of truth I felt bound to charac-
terise certain of the conclusions arrived at by Dr Maddox as
simply incredible, but I regarded his memoir as forming " an
important contribution to our knowledge of the structure of the
mutton measle.55 I had no idea that in pointing to errors of
interpretation I should offend the excellent author. However,
a long letter appeared in the ' London Medical Record/ in which
Dr Maddox showed that he was much vexed that I should
have " impugned55 the " accuracy of his conclusions.55 He
defended his position with the support of no less an authority
than Dr Macdonald, F.R.S., the distinguished Assistant Pro-
fessor of Naval Hygiene at the Victoria Hospital, Netley. Dr
Maddox says : — " We were quite alive to the anomalous posi-

CESTODA

tion. Hence the exceptional) ility of the case rests on more
than my own evidence." In regard to this unfortunate dispute
I will only add the expression of my conviction that Drs
Maddox and Macdonald will eventually become satisfied that
no cestode scolex is capable of displaying either mature or
immature ova in its interior.

Bibliography (No. 15). Cobbold (1. c, supra), 1865-75. —
Idem, " On Measly Meat, &c./' the ' Veterinarian/ Dec, 1876.
— Idem, "The Mutton Tapeworm (T. tenella)," No. 16 in my
revised list of Entozoa, the 'Veterinarian/ Dec, 1874. —
Diesing, G. M. {Tcenia tenella, Pruner nec Pallas), in "Kevis
der Cephalocotyleen," ' Sitzungsb. der Math. -Mat. Class d. k.
Akad. der Wissenschaften/ Bd. xlix, s. 369, 1864. — Maddox
(1. c, supra), 1873. — Mayrhofer, J. C, ' Die helminth, des
Menschen/ Erlangen, 1854. — Pruner, ' Krankheiten des
Orients/ s. 245, 1847.

Tcenia lophosoma, Cobbold. — This is a good species not-
withstanding the doubts that have been expressed by Heller
and others regarding it. I have called it the ridged tapeworm
in consequence of the presence of an elevated line coursing the
whole length of the body, which measures about eight feet.
The reproductive papillse are remarkably prominent and uni-
serially disposed throughout the entire chain of proglottides.
It is quite an error to suppose that this species is a malformed
cestode, or that it has any resemblance to Kiichenmeister's
variety of tapeworm from the Cape of Good Hope. Neither
does it in the slightest degree resemble the remarkably mal-
formed T. mediocanellata described by Mr Cullingworth. Of
the distinctiveness of this parasite as a species, any one may
satisfy himself by an inspection of the nearly complete strobile
preserved in the Pathological Museum attached to the Middlesex
Hospital Medical College. From the examination of several
mature proglottides detached from this specimen, I find their
average breadth to be one fifth of an inch, by three quarters
of an inch in length. Their greatest thickness does not exceed
the ^th of an inch. The eggs resemble those of other tape-
worms, and offer a diameter of about ey from pole to pole.

Bibliography (No. 16). — Cobbold, "Parasites of Man," in
the < Midland Naturalist/ April, 1878, p. 98.— Idem, ' Tape-
worms/ 1st edit., p. 52, 1866; 3rd edit., p. 27, 187 '5.— Culling-
worth (see Bibl. No. 18). — Davaine, c Les Costoides/ 1. c, p. 573.
— Heller, 1. c, s. 594.

100

PARASITES OP MAN

Tania nana, Siebold. — As regards the dwarf tapeworm,
unless Spooner's case be genuine, there is but one solitary
instance on record of its occui'rence in the human body ; more-
over, we have no evidence of its having existed in any other
host. It was discovered by Dr Bilharz, of Cairo, at the post-
mortem examination of a boy who died from inflammation of
the cerebral membranes. Prodigious numbers existed. The
largest specimen measured only one inch in length. To the
naked eye these worms resemble short threads, and conse-
quently they might very readily be overlooked. The head is
broad and furnished with a formidable rostellum armed with a
crown of hooks. These hooks have large anterior root-processes,
which, extending unusually forward, impart to the individual
hooks a bifid character. By far the best account of this worm
is furnished by Leuckart, to whom I am indebted for a
specimen.

Bibliogeaphy (No. 17). — Cobbold, ' Entozoa/ p. 244. —
Davaine (1. c, Bibl. No. 2), p. 57 '4.— Heller, 1. c, s. 606.—
Kuchenmeister, 1. c, Eng. edit., p. 141. — Leuckart, 1. c, Bd. i,
s. 393. — Von Siebold and Bilharz, in Von Sieb. and Koll.
Zeitschr., Bd. iv. — Spooner, e Amer. Journ. Med. Sci./ 1873. —
Van Beneden, ' Iconographie/ 1. c, pi. iii, fig. 17. — Weinland,
' Bvplacanihus nanus,' 1. c, p. 85.

Tcenia Madagascariensis, Davaine. — This appears to be a
well-defined species although the head has not yet been seen.
It probably forms the type of a distinct genus. Dr Grenet,
stationed at Mayotte (Comores), twice encountered single
specimens passed by two young children, eighteen and twenty-
four months of age respectively. The proglottides have their
genital pores uniserially arranged, and they show, in their
interior, remarkable egg-capsules, from 120 to 150 in number
in all, each containing from 300 to 400 eggs. These give a
long diameter of for the outer envelope and for the inner,
or shell proper. The embryo measures only the ~ of an inch.

A full account of this parasite, with figures, is given by
Davaine (' Les Cestoides/ 1. c, Bibl. No. 2, p. 577 et seq.).

Tania marginata, Batsch. — Although I possess no certain
evidence of the occurrence of this parasite in its adult condition
in the human bearer, yet there is a tapeworm in the Edinburgh
Anatomical Museum referable to this species, which was said
to have been obtained from the human body. This worm is
very common in the dog.

OESTODA

101

The principal evidence demonstrating the occurrence of the
larval representative of this species (Oysticercus tenuicollis) in
man, rests upon the two cases recorded in Schleissner's ' Noso-
graphy' of Iceland. One of the alleged instances, however, has
been proved by Kiichenmeister and Krabbe to be that of an
echinococcus ; so that, after all, there only remains the solitary
case observed by Schleissner himself, in which the parasite can
fairly be considered as the " slender-necked hydatid."

To the above, however, may probably be added a specimen
preserved in the Anatomical Collection at King's College, London.
It was found connected with an ovarian cyst.

Tcenia elliptica, Batsch. — This parasite is readily recognised
not merely by its delicate form and small size, but also by the
circumstance of its supporting two sets of reproductive organs
in each mature joint. Their outlets are situated at the centre
of the margin of each segment, one on either side. ■ Ordinarily
infesting the cat, this worm is a mere variety of the common
Tcenia cucumerina of the dog. At all events, from the evidence
put forth by Eschricht, seconded by Leuckart, there is every
reason for believing that one or other of these closely-allied
varieties is liable to infest the human body. It was originally
stated by Eschricht that he had received a Tcenia canina which
had been passed by a negro slave at St Thomas, Antilles. This
is a synonym of T. elliptica, which must therefore be very rare
in the human body, possibly only occurring in the negro race.

In regard to the source of this parasite, it has been shown
by Melnikow that the scolex of Tcenia cucumerina resides in
the louse of the dog {Trichodectes latus), and thus it is
exceedingly probable that the scolex of Tcenia ellijptica resides
in the louse of the cat [Trich. subrostratus) . How man becomes
infested is not so clear. Melnikow's paper on the juvenile state
of this cestode is contained in the ' Archiv fur Naturgeschichte'
for 1 869, and is illustrated by a figure of the measle.

Tcenia flavopuncta, Weinland. — Eegarded as a new species,
the discovery of this little tapeworm is due to the investigations
of Weinland. In Dr Jackson's ' Catalogue of the Boston
Medical Improvement Society' an account of the contents of a
phial is recorded as follows : — " Specimen of Bothriocephalus
three feet in length, and from half a line to one line and a
quarter in width, from an infant. The joints are very regular
except at one extremity, where they approach the triangular
form, are very delicate, and but slightly connected, as shown in

102

PARASITES OF MAN

a drawing by Dr Wyman." It is further stated that the infant
was nineteen months old, and that the worm was discharged
without medicine, its presence having never been suspected. It
was presented by Dr Ezra Palmer in the year 1842. On
examining the fragments, Dr Weinland found, instead of a
solitary specimen, at least six different tapeworms, all of them
being referable to a hitherto undescribed species. There were
no heads ; nevertheless, it was ascertained that the worms varied
from eight to twelve inches in length, the joints or segments
being very broad, and at the same time narrowed from above
downwards. The parasite was named " the spotted tapeworm,"
in consequence of the presence of yellow spots near the middle
of the joint. They represent the male organs of reproduction,
the outlets of which, as in my T. lophosoma, occur all along one
side of the body or strobile. In Weinland's estimation this
parasite forms the type of a new genus which he calls Hymeno-
lejpis. A full account of the worm is given in his well-known
essay (1. c, Bibl. No. 2).

Tcenia abietina and other varieties. I can only notice very
briefly certain cestodes which either present malformations or
which may be regarded as mere varieties. First in this series is
~Wem\a,nd'sT. abietina. Noonewhohas studied his 'Beschreibung
zweier neuer Teenioiden aus dem Menschen/ Jena, 1861, can
doubt that it is a mere variety of T. mediocanellata. The
monstrosity described by him as referable to T. solium must also
be referred to the beef tapeworm. The variations in the cha-
racter of cestode proglottides is practically infinite. A museum
might be filled with them. Most common with T. medio-
canellata, these varieties more or less prevail with other species.
Thus I have seen them in Tsenise and Bothriocephali alike. I
have obtained segments of T. mediocanellata having sexual
outlets on both sides of the proglottis, so regularly disposed in
a few segments as to suggest the notion of a new species. The
coalescence of several segments into one compound segment is
frequent, but the most remarkable specimen that I have seen is
one contained in the museum of the Royal College of Surgeons.
In the old Hunterian catalogue the specimen is described as
" two joints of the Tcenia solium, with a number of orifices in
unequal series on either side." As stated in the new catalogue
of the series, prepared by myself, the "lower segment is furnished
with twenty-two sexual orifices, one of which is situated in the
central line" on the ventral surface (as in Bothriocephali).

CESTODA

103

Keferences to this and other specimens in the Hunterian
Collection will be found below (see Pittard). In regard to
Weinland's conjectural Tcenia acanthotrias, based on the
circumstance of his having found a Cysticercus that presented
three rows of hooks on its rostellum, I need only say that if
such a Tarda were found it would only turn out to be a
malformed T. solium. The specimens, however, are none the
less interesting. Very remarkable and altogether exceptional
characters are presented by the strobile of the cestode described
by Mr Cullingworth, of Manchester, and of which I possess
specimens. Here, apparently, at least two tapeworms are joined
together throughout the entire chain of proglottides without
intermission. The three margins of each compound segment
project at equi-distant angles. Could we have secured the head
we should certainly have found six or eight suckers present,
since the finest neck-segments showed that the malformation
pervaded the entire colony of zooids, sexually mature and
otherwise. Mr Cullingworth' s specimen is so remarkable that
I subscribe full particulars of the case in his own words. He
says : — " A respectable married woman, named Ann H — , forty
years of age, residing in Salford, brought to my out-patient
room at St Mary's Hospital, Manchester, on September 3rd,
1873, a few segments of tapeworm as a sample of what she had
been passing per anum for about two years. Although never
in the habit of taking meat absolutely raw, she told me, on
inquiry, that she was particularly fond of tasting it when only
partially cooked. The segments were unlike anything I had
seen before, and I took them home for examination, ordering the
patient meanwhile a draught containing a drachm of the oil of
male fern, and giving her strict injunctions to bring to me every
fragment that passed away as a result.

" On September 1 7th she brought me portions of a tapeworm
corresponding throughout to the segments I had already seen,
and measuring altogether nine feet in length. Unfortunately,
the head was not to be found. Along the middle line of every
segment in the body a crest or ridge runs longitudinally, and in
the centre of the margin of this crest the genital pore is situ-
ated. [In 304 segments examined, only four had the genital
opening placed laterally. One segment had two openings,
viz. one at the lateral margin and the other in the crest.]
Underneath the segment there is a longitudinal groove, and
the lateral portions are folded together by the apposition of their

104

PARASITES OF MAN

under surfaces. When hardened in spirit the section of a
segment presents a three-branched appearance, the branches
being of unequal length, but placed at equal angles. The
uterus sends vessels into the crest as well as into the sides of the
segment ; and the contained ova are exactly like the ova of an
ordinary Taenia mediocanellata. Wedged in between, or
attached to, the segments here and there, is a stunted and
ill-shaped joint, with irregular and unequal sides. A mature
joint measures from five eighths of an inch to three quarters of
an inch in length, and about half an inch in breadth, and
the breadth or depth of the crest is usually one eighth of
an inch.

" There are only two specimens that I can find on record at
all similar to the one here described, and both of these differ
from it in several important particulars. Kiichenmeister men-
tions, as a variety of Taenia mediocanellata, a tapeworm sent
to him from the Cape of Good Hope by Dr Rose. This worm
possessed a longitudinal ridge, but he describes its mature
segments as 'extremely massive' — more than an inch in length
and §" in breadth. The genital pores, too, were irregularly
alternate, and not situated on the crest. On March 20th, 1866,
Dr Cobbold exhibited to the Pathological Society of London a
specimen of crested tapeworm which was discovered in the
museum of Middlesex Hospital, and to which he proposed to
give the name Taenia lophosoma (Xotyoq, crest ; awjua, body). The
reproductive papillee were all on one side of the chain of seg-
ments, a peculiarity which entirely distinguished it from the
Cape of Good Hope variety of Kiichenmeister. The head of
the creature was wanting. It will thus be seen that my
specimen does not correspond with either of these in the situa-
tion of the genital aperture. Here it is placed in the crest
itself, and not unilaterally, as in Dr Cobbold's specimen, or
alternately, as in Kuchenmeister's. It further differs from the
Cape variety in the more moderate dimensions of its proglot-
tides. I have adopted, however, the name suggested by
Dr Cobbold in the communication referred to, inasmuch as it
sufficiently indicates the principal distinguishing feature of the
specimen. I may mention that Dr Cobbold saw the specimen
during his visit to Manchester, and that he regarded it as a
most remarkable and unique abnormality."

Further, in connection with abnormal cestodes, I may observe
that Weinland's case of a triple-crowned Cysticercus does not

OESTODA

105

stand alone, since a similar specimen is, I believe, in the
possession of the Rev. W. Dallinger. This was removed from
the human brain. Curious as this subject is, I cannot dwell
upon it. Not only are the mature tapeworms and their Cysti-
cerci liable to present monstrosities, but even also their prosco-
lices or six-hooked embryos. Thus, twelve hooks were observed
by Salzmann in the embryo of T. elliptica,, and Heller also
figures two embryos of T. medio canellata (T. saginata, Grceze)
with numerous hooklets. Dujardin saw seven in a Bothrioce-
phalic embryo. Occasionally there have been errors of inter-
pretation made by observers. Thus, Diesing has given beautiful
figures of Dibothrium hians in such a way as to suggest different
degrees of monstrosity affecting the tail end of the strobile ; but
this splitting has clearly resulted from injury. Thus also, when
I removed five specimens of a new cestode (Diphyllobothrium
stemmacephalum) from the intestines of a porpoise, one of them
was cleft nearly half way up the strobile. This had been done
by the scissors employed in slitting up the gut ; but owing to
perfect contraction of the incised edges, it was some time
before I discovered that the apparent monstrosity had been
artificially produced. Lastly, I may add that many of the
older writers were well acquainted with larval and other
anomalies. Thus Rudolphi described a two-headed Cysticercus
from a Lemur, and also a double-headed Taenia crassicollis. This
worm had a tripartite body j as had likewise a Taenia crassi-
collis of which he did not possess the head {corpore prismatico).
Other monstrosities were described and figured by Bremser
and Creplin. Pallas mentions a two-headed Tricuspidaria
(Tricenophori nodulosi bicipites), and, as already stated at p. 97,
a double-headed Cysticercus has been obtained from the liver
of a sheep.

Before quitting the Taeniae proper, I may observe that several
other species have been indicated, based on ovular and other
insufficient characters. To these belong Ransom's supposed
tapeworm, and also Weinland's Taenia meg aid on.

Bibliography (No. 18).— Bonnet, C, ' CEuv. Compl./ torn vi,
p. 191, 1791. — Bremser, Atlas, by Leblond, PI. iv. — Chaussat,
' Comptes Rendus/ p. 20, 1850. — Cobbold, ' Catalogue of the
specimens of Entozoa in the Museum of the Royal College of
Surgeons of England/ Nos. 118—121, London, 1866.— Idem,
'Worms/ 1. c, p. 78.— Idem, "On a Cysticercus from the
Human Brain/' ' Brit. Assoc. Rep./ 1870— Creplin, ( Ttenia

106

PARASITES OP MAN

Monstrum, &c./ Berlin, 1839.— Culling worth, 0. J., " Notes on
a remarkable specimen of Tapeworm {Taenia lophosoma, Cob-
bold)," 'Med. Times and Gaz./ Dec, 1873.— Davaine, 'Les
Cesto'ides/ 1. c, p. 570. — Biesing, ' Zwanzig Arten von
Cephalocotyleen/ figs. 1 and 2, taf. ii (aus dem xii, Bd. d.
denkschr. d. Math.-nat. 01. d. k. Akad.), Wien, 1856. — Dujardin,
1. c, p. 619. — Heller, 1. c, s. 600. — Euchenmeister, 1. a, Eng.
edit., p. 139. — Leuclcart, 1. a, s. 303 and 465. — Levacher, ' Journ.
Tlnstitut/ p. 329, 1841. — Pittard, S. R., Eemarks in his article
" Symmetry," Todd's ' Cyclop./ vol. iv, p. 848, 1849-52, in
which he refers to a monstrous Bothriocephalus (T. lata) in the
Hunterian Museum, old ' Catalogue of Nat. Hist./ pi. iv, p. 50,
No. 205; see also my ' Catalogue/ 1. c, supra, No. 167. —
Hansom, in Reynolds' ' System of Medicine/ — Budolphi,
* Synops./ p. 545 and 598-9, with fig. showing the heads of
Cystic. Simice (biceps), widely apart, 1819. — Weinland (T.
megalbon) , in Zoolog. Garten, Frankf., 1861, s. 118. — Idem,
' Essay/ 1. c, p. 11.

Bothriocephalus latus, Bremser. — This species, though seldom
seen in England, is sometimes brought hither by persons who
have been residing for a time in foreign countries. It is indi-
genous in Ireland, and, though by no means common there, has
been called the Irish Tapeworm. As regards its distribution
in Europe it is much more prevalent in some districts than in
others. On this point Leuckart remarks that "foremost amongst
these are the cantons of West Switzerland, with the adjacent
French districts. In Geneva, according to Odier, almost a
fourth part of all the inhabitants suffer from Bothriocephalus.
It is also common in the north-western and northern provinces
of Russia, in Sweden, and in Poland. In Holland and Belgium it
is likewise found, but, on the whole, not so frequently as in the
first-named countries. Our German fatherland also harbours
them in some districts, especially in eastern Prussia and Pome-
rania, and there have appeared cases in other places, as in
Rhenish Hesse, Hamburg, and even in Berlin ; these being
apparently spontaneous instances."

Unlike the ordinary tapeworms, the segments of the broad
tapeworm do not individually separate so as to become indepen-
dent organisms, a circumstance which is highly favorable to the
bearer. Its remarkable breadth, and the extremely numerous
and closely-packed proglottides, impart a sufficiently distinctive
character ; but this parasite may be more fully characterised as

CESTODA

107

the largest human cestode at present known, attaining a length
of more than twenty -five feet, and sometimes measuring nearly
an inch in breadth ; the so-called head £" in width, bluntly
pointed at the tip, much elongated or club-
shaped, slightly flattened from behind for-
wards, and furnished with two laterally disposed
slit-like fossae or grooves, but destitute of any
armature : anterior or sexually-immature seg-
ments of the body extremely narrow, enlarging
in a very gradual manner from above down-
wards ; joints of the lower half of the body
gradually decreasing in width, but enlarging
in depth j sexually-mature segments usually
about | of an inch in depth, but those near
the caudal extremity frequently \" , and quad-
rate in form ; body flattened, but not so uni-
formly as obtains in the ordinary tapeworms,
being rather thicker near the central line ;
total number of joints estimated at nearly
4000, the first sexually-mature ones being
somewhere about the six hundredth from the
head ; reproductive orifices at the central line,
towards the upper part of the segment at the
ventral aspect, the vaginal aperture being
immediately below the male outlet, and both
openings surrounded bypapilleeform eminences;
uterus consisting of a single tube, often seen
regularly folded upon itself, forming an opaque,
conspicuous, centrally- situated rosette ; eggs
oval, measuring in length by -L" in breadth,
having three shell-coverings, and a
operculum at one end, as occurs in the fluke-
worms. Owing to the dark color of the egg
shells, the uterine rosette is readily seen by
the naked eye as a conspicuous deep brown spot at the centre
of each successive segment.

The source and development of this parasite are points of
considerable interest. The eggs are of comparatively large size,
and after expulsion and immersion in water they give passage
to beautifully ciliated embryos, which latter produce larvse
furnished with a boring apparatus. These larvse resemble the
six-hooked embryos of other tapeworms. In what animals the

lid-like Fig. 25.— Head and nerk
of Bothriocephalus latits.
a, Front view. 'Ihe
smaller figure represents
tlie head as seen from the
side. After Knoch.

108

rAUASITES OF MAN

larvae subsequently develop themselves is not ascertained with
certainty, but it is probable that persons become infested by
eating imperfectly cooked fresh-water fish. Leuckart has
suggested that the intermediary bearers are species of the
salmon and trout family. Dr Knoch, of Petersburg, thought

that there was no need of the inter-
mediate host. He believed that he had
succeeded in rearing young broad tape-
worms in the intestines of dogs. It
was Leuckart who first explained the
source of Knoch's errors of interpre-
tation. Although Knoch administered
eggs of Bothriocejphalus latus to dogs,
and afterwards found young tapeworms
of the species in question in the intes-
tines of the dogs, it did not logically

Fig. 26. — Proscolex, or six-hooked em- „ n. _ . n,. , ■>

bryoofBothriocephaius.escapmgfrom follow that any genetic relation (as be-

its ciliated covering. After Leuckart. , . , . , -, . . -. . .

tween the egg -contents and the adult
worms) had been thereby established. The circumstance that ripe
ova of the Bothriocephalus always contain six-hooked embryos,
must alone imply that an intermediate host is necessary for the
formation of Oysticerci or measles. If the broad tapeworm could
be reared in a direct manner by the administration of Bothrio-
cephalus eggs, there would be no need for the presence of
boring hooklets in the proscolex. These are necessary for
invading the flesh of some intermediate host.

Dr Fock, of Utrecht, has sent me particulars of an interesting
case, and he suggests that infection comes from the little river
bleak (Leuciscus alburnus). Writing from Utrecht in December,
1877, Dr Fock, after referring to a former case, goes on to
say :—

" Permettez moi, cher confrere, que je rappelle a votre
souvenir que vous avez eu l'obligeance de communiquer au
public une observation, de ma main, sur un cas tres rare de ver
rubanaire, d'un Bothriocephale, chez une petite fille juive.
Malheureusement je n'ai pu en donner, de plus amples details,
parce que cette enfant n'a plus, depuis ce temps-la, rendu la plus
petite parcelle de ver. II y a maintenant quinze mois, et voila
que de nouveau un cas pareil se presente. Une femme mariee,
frisonne, et, cette fois-ci encore, juive, s'est adressee a moi pour
la debaraaser de son ver. Elle me disait avoir rendu, il y a
quelque temps, des fragments, ou plutdt un fragment de la

OESTODA

109

longueur (Tun metre, (Tun ver solitaire, pour lequel elle avait ete
traitee, sans succes, par son medecin ordinaire. A cause de cela
clle s'adressa a moi, et je lui ai repondu quelle devrait revenir
la premiere fois qu'elle rendrait de nouveau, spontanement, un
nouveau fragment. Apres unmois d'intervalle elle est revenue
en me montrant un fragment de la longueur d'un demi-metre
qu'elle venait de rendre spontanement, apres avoir jeuni par
precepte religieuse, et deux jours apres cela, traitee par Tecorce
de grenadier, elle a rendu un Bothriocephale parfaitement con-
ditional en entier.

" Ce cas me semble assez interessant pour etre communique
de nouveau, d'abord parce que jusqu'ici personne n'a pu dire par
quel chemin a pu sJintroduire un tel helminthe, et ensuite parce
que ce chemin doit se presenter bien rarement dans nos
contrees (ou en Angleterre) puisque dans le courant d'une
trentaine d'annees ayant rencontre des centaines de tsenias, ce
cas-ci est seulement le second dont je suis gratifie. II me
semble digne de reflexion que ce Cas-ci se presente cette fois-ci
de nouveau chez une juive. Est ce cas-ci fortuit, ou bien y-a-
t'il un lien de causalite entre ce ver rare et le genre de nourri-
ture ou de boisson de ces bonnes gens ? La dame me recontait
que, en Frise, il y a un poisson tres recherche qui s'appelle en
Hollandais bleck, en Anglais blay ou bleak, et dont ils sont tres
friands, dans lequel, ils rencontrent tres souvent un tres grand
ver rubanaire. Une autre personne me disait avoir ete a
table chez un ami, qui ne sachant probablement ce qu'il
mangeait, savoura avec beaucoup de delice cette friandise de-
goutante."

After describing the specimen, Dr Fock concludes his remarks
with a suggestion as to the possibility of introducing tapeworm
into the human body by potable water, into which Cysticerci
have accidentally found their way. Dr Fock remarks : — " J'ajoute
une reflexion par rapport a la provenance des autres tsenias, qui
jusqu'ici sont introduits par Tusage de la viandenon assez cuite
ou rotie, ou saignante ; mais, ne se pourrait-il pas que des debris
de la chair d'un animal ladre fussent introduits fortuitement
dans l'eau, par example, d'un fosse, et que celle-ci employee
comme boisson contint des Cysticerques et par ainsi aussi uno
cause de Taenia ? Ce n'est qu'une conjecture que je propose en
terminant cet article."

Although I cannot at all agree with Dr Fock in regarding
water as a source of infection in the manner he indicates, yet

110

rARASlTES OF MAN

the still more recently expressed opinions of MM. Bertolus and
Duchamp, based on experimental researches, render it tolerably-
certain that Leuckart' s original surmise was correct, and that

we must look to freshwater fishes for
the larvae of the broad tapeworm. In
the section of this work devoted to the
parasites of fishes I shall make par-
ticular allusion to the experiences of
Dr Bertolus; but as confirming the view
of Leuckart I may here observe, that
Bertolus has almost proved that the
so-called Ligula nodosa infesting the
common trout is merely a sexually in-
complete example of Bothriocephalus
latus. The bleak (Leuciscus alburnus)
shares with other freshwater fishes the
privilege of harbouring a species of
Ligula (L. digramma) ; but whether
this form bears any genetic relation to
our human Bothriocephalus latus can
only be determined by actual experi-
ment. If, as Duchamp and others have
either indicated or implied, Ligula aU
bumi is a synonym of the bleak's ces-
tode in question, then it is evident that
the sexually mature form of the Ligula
of the bleak is the well-known L. sim-
jplicissima of many water birds and of a
few other avian species. Probably the
bleak-eaters of Holland consume many
kinds of freshwater fishes, including
various species of the salmon and trout
family.

The symptoms occasioned by Bothrio-
cephalus latus do not differ materially
from those produced by other tapeworms.
According to Odier, as quoted by Davaine, there is not unfre-
quently a tumid condition of the abdomen, with sickness,
giddiness, and various hysterical phenomena occurring at night.
Pain in the region of the heart, palpitations, and faintness are
also mentioned.

As already hinted, this cestode is very liable to present

Fig 27. — Stroliile of Bothriocephalus
cordatus. After Leuckart.

CESTODA

111

abnormalities of structure, the proglottides frequently displaying
double sexual orifices, witli corresponding duplication of the
reproductive organs internally. For details respecting the
anatomy of Bothriocephalus I must refer to the works of
Kiichenmeister and Leuckart ; and more particularly to the
memoir of Drs F. Sommer and L. Landois, who have supple-
mented the previous researches of von Siebold, Leuckart,
Bottcher, Stieda and others by beautiful investigations of their
In the pages of f Nature/ for 1872, I gave a resume of

own.

Sommer's memoir, which will be found quoted below.

Bothriocephalus cordatus, Leuckart. — This species is identical
with a worm long ago described by Pallas and Linneus. At
present it is only known to infest the residents of North Green-
land, but it is probably distributed throughout the north gene-
rally. It attains the length of about one foot, and has a small
heart-shaped head, whose apex is directed
forwards. The neck is so obscure that it
may be said to be altogether wanting, the
segmentation of the body being well marked
immediately below the head. Though so small
a species, Leuckart, who first described it,
counted between six and seven hundred joints.
As in the broad tapeworm, the reproductive
orifices are serially disposed along the centre
of the ventral line, but a close inspection
shows that the folds of the egg-bearing organ
are comparatively more numerous. This
worm does not appear to be a frequent
resident in the human body, though it is by
no means uncommon in the dog. Possibly
it may yet be found in the inhabitants of
some of our northern and western isles.

Bothriocephalus cristatus, Davaine. — This
cestode measures between nine and ten feet
in length, and is characterised by the
presence of two remarkable prominences,
together forming a sort of rostellum or
crest which is covered by numerous minute
papillae. The full-grown segments are less
than half an inch in breadth ; the body
of the parasite being narrower than that of the broad
species. The original description of the parasite by Davaine

WW
m. v

W-r IP
f If .

Fig. 28.— Head of Bothrio-
cephalut erutatut, viewed
from the front. After
Duvninc.

112

1'AR.ASITES OF MAN

is based on two specimens, one of which, quite perfect, was
obtained from a child five years old, under Dr FereoPs care
at Paris. The other was passed spontaneously by an adult
residing at Haute- Saone. I have here copied one of Davaine's
original figures of the head of the worm.

Bibliography (No. 19). — Bertolus, " Mem. sur le development
du Dibothrium latum " (in Appendix to Duchamp's work, see
Bibliog. No. 59). — Blanchard, " Recherches, &c./' 'Ann. des
Sci. Nat./ ser. 3, Zool., PI. 11, 12, 1848.— Bottcher, " Studien
ueber den Bau des Both, latus," ' Virchow's Archiv/ s. 97 et sea,
1864.— Bremser, 1. c, Bibl. No. 1, s. 88, 1824.— Chiaje, ' Com-
pendio, &c./ Tab. iii,'figs. 1—5, 1833.— Gobi old, ' Entoz./ p.
289, 1864. Idem, " Remarks oh the Broad Tapeworm" (with
a letter from Dr Pock), the 'Veterinarian/ July, 1878. — Greplin,
in Ersch and Gruber's ' Encyclop./ 1839, p. 296. — Bavaine,
' Traite/ 1. c, 1860 ; 2nd edit, (passim), . 1877. — Idem, art.
' Les Cestoides/ 1. c, Bibl. No. 2, p. 580— 591, 1876.— Bujar din,
1. c, Bibl. No. 1, p. 612, 1845.— Eschricht, B. F.,< Anat-physiol.
Untersuchungen ueber die Bothriocephalen/ Breslau, 1840.
— Bock (see Cobbold). — Heller, 'Darmschmarotzer/ 1. c, s. 606,
1876. — Knoch, ' Petersburger Med. Zeitschrift/ 1861. — Idem,
' Die Naturgeschichte des breiten Bandwurms (B. latus, auct.)/
St Petersburg, 1862. — Kiichenmeister, ' Ueber cestoden/ 1. c,
1853. — Beuckart, 'Die Blasen Bandwiirmer/ 1856. — Idem, 'Die
mensch. Par./ Bd. i, s. 414—448, und 757, 1863 ; and Bd.
ii, s. 866, 1876.— Owen, Todd's ' Cyclop./ 1837— Sommer und
Bandois, aus Sieb. und Koll. Zeitschr., ' Beitrage zur Anato-
mie der Plattwiirmer/ Leipsig, 1872 ; see also the resume in
' Nature ' for Aug., 1872, p. 278. — Wawruch, ' Pract. Mono-
graph, d. Bandwiirm-Krankheit/ 1844, s. 33.

Echinococcus hominis (the common hydatid). — This larval
entozoon has acquired various names according to the kind of
bearer in which it happens to have been found ; but all the true
hydatids or acephalocysts, whether infesting man or animals, are
referable to one and the same species of parasite. They have
been termed Echinococcus hominis, E. veterinorum, E. polymor-
phus, E. exogena, E. endogena, E. multilocularis, according to
circumstances. All of them represent a juvenile stage of the
Taenia echmococcus or hydatid-forming tapeworm which infests
the dog and wolf. Experimental proof of this fact has been
furnished by Yon Siebold (1852), Haubner, Leuckart, Kiichen-
meister, Van Beneden, Naunyn, Nettleship, Krabbe, and others.

CESTODA

The first successful rearing of Taeniae
with human hydatids was accomplished by
Naunyn (1864), his results being subse-
quently verified by Krabbe and Finsen
(1865). Zenker, Ercolani, and several
others, including myself, also conducted
feeding experiments with human hydatids
which were attended with negative re-
sults. In the case of one of my experi-
mental dogs the animal was liberated by
an ill-disposed person before I had oppor-
tunity to destroy it. As the experiment
was carefully conducted, the animal may
have proved a source of fresh echino-
coccus-infection. Mr B. Nettleship's emi-
nently successful experiment was made
with hydatids obtained from a sheep.
The converse experiment, namely, that of
rearing hydatids with the mature pro-
glottides of Taenia echinococcus adminis-
tered to animals, has been performed most
successfully by Leuckart, and by Krabbe
and Finsen ; by the former in the pig, by
the latter in a lamb, with tapeworms
that had also been reared by experiment.
Zenker, later on, reared the Taenia from
hydatids obtained from an ox.

The sexually mature Taenia echinococcus
may, for the purposes of diagnosis, be
characterised as a remarkably small ces-
tode, seldom reaching the fourth of an
inch in length and developing only four
segments, including that of the head •
cephalic extremity capped by a pointed
rostellum, armed with a double crown of
comparatively large-rooted hooks, from
thirty to forty in number; the four
suckers prominent, and succeeded by an
elongation of the segment forming the
so-called neck j final segment, when sexu-
ally mature, equalling in length the three
anterior ones ; reproductive papilla at the

114

PARASITES OP MAN

margin of the proglottis rather below the central line j proscolex
or embryo giving rise to the formation of large proliferous
vesicles, within which the scolices or echinococcus-heads are
developed by gemmation.

When an animal is fed with the mature proglottides of Tania
echinococcus the earliest changes that take place are the same
as obtain in other cestodes. The segments are digested ; the
shells of the ova are dissolved ; the six-hooked embryos
escape. The embryos bore their way into the organs of
circulation, and thence they transfer themselves to the different
organs of the host ; being especially liable to take up their
abode in the lungs and liver. Having arrived at this, their
resting stage, the embryos are next metamorphosed into
hydatids. According to Leuckart's investigations the juvenile
hydatid is spherical at the earliest stages ; being surrounded by
a capsule of connective tissue formed from the organs of the
host. After removal from its capsular covering, the vesicle
consists of a thick laminated membrane, forming the so-called
cuticular layer, and a central granular mass, which subsequently
becomes enveloped by a delicate granular membrane. At the
fourth week the echinococcus capsule measures about in
diameter, its contained hydatid being little more than half this
size. Its future growth is by no means rapid, seeing that at
the eighth week the hydatid has attained only the ^' in diameter.
At this period the central granular mass develops a number of
nucleated cells on the inner surface of the so-called cuticle.
These cells, which at first are rounded or oval, become angular
or elongated in various directions, and even distinctly stellate ;
and in this way a new membrane is formed, constituting the
so-called inner membrane or granular layer. The intermediate
stages between this condition and that of the fully-formed
echinococcus hydatid have not been satisfactorily traced in
detail ; nevertheless, Krabbe and Finsen's experiment on a lamb
showed that within a period of little more than three months
well-developed echinococcus-heads may be formed in the interior
of the vesicles. It is thus clear that the production of scolices
immediately follows the formation of the granular layer, and
this is succeeded, though not invariably, by the formation of
daughter- and grand-daughter- vesicles, which are sometimes
termed "nurses." These latter maybe developed exogenously
or endogenously.

The appearance of hydatids varies very much according to

CESTODA

115

their mode of formation, to the kind of host in which they are
present, and to the character of the organs in which they happen
to take up their residence. The so-called exogenous type
occurs sparingly in man, whilst the endogenous type is very
abundant. The peculiar form known as the multilocular echino-
coccus is probably a mere variety of the exogenous type. The
exogenous and endogenous hydatids may coexist in the same
bearer. In the lower animals we commonly find the organs of
the body occupied by numerous lobulated cysts, varying in size
from a walnut to a goose's egg, but sometimes rather larger.
They are rarely solitary, being particularly liable to occupy both
the liver and lungs in the same animal. The viscera are some-
times crowded with cysts. The hydatids do not usually protrude
much beyond the surface of the infested organ, but lie imbedded
within its parenchymatous substance.

The multilocular variety was first described by Yirchow. In
reference to it Leuckart writes as follows :

" Hitherto we know this growth only from the liver, in which
it forms a firm, solid, and tolerably rounded mass of the size of
the fist or even of a child's head. At first sight it looks more
like a pseudoplasm than a living animal parasite. If you cut
through the tumour, you recognise in its interior numerous
small caverns, mostly of irregular shape, and separated from
one another by bundles of connective tissue, more or less thick,
and including a tolerably transparent jelly-like substance. In
the intervening stroma a blood-vessel or a collapsed bile-duct
runs here and there ; but there is nowhere any trace of true liver
substance. The outer boundaries of the tumour are in most
cases pretty well defined, so that the attempt to cut these
growths out is not difficult. In particular spots, especially at
the surface, one sometimes sees white, moniliform, jointed lines
passing off from the tumour, and even thicker terminations
which, perhaps, expand in the neighbouring liver-parenchyme
into new (multilocular) groups of different size. In one case,
recorded by Virchow, the growth extended, together with
Glisson's capsule, a long way towards the intestine." To this
description it may be added, that the growth on section pre-
sents an appeai'ance not altogether unlike alveolar colloid,
having, in point of fact, been confounded with that pathological
product, with which, however, as stated by Yirchow, it has
nothing in common. This is proved not only by the occurrence
of the pathological features above mentioned, but also, more

116

PARASITES OP MAN

particularly, by the well-ascertained presence of echinococcus-
heads in most of the so-called alveoli. Several hypotheses
have been broached with the view of explaining the mode in
which these multilocular hydatid growths are formed. Virchow
thought that the echinococcus vesicles were primarily formed in
the lymphatic vessels, whilst Schroder van der Kolk supposed
that they originally took up their abode in the biliary ducts.
Although, thanks to the courtesy of Professor Arnold Heller in
giving me a specimen, I have been enabled to confirm much that
has been written in respect of the morbid appearances, I can
add nothing towards the solution of the difficulty in question.
Until lately it was supposed that the multilocular variety of
hydatids only existed in man, but Professor Bollinger has
encountered it in the liver of a calf.

Selecting any ordinary fresh example of the exogenous kind,
and laying the tumour open with a scalpel, we notice in the

first instance an escape
of a clear transparent,
amber -coloured fluid.
This previously caused
the distension of the
sac. If the tumour is
large, this escape will
probably be followed
by a falling in, as it
were, of the gelatinif orm
hydatid membrane, in
which case the inner wall
of the external adventi-
tious investment or true
fibrous cvst will be laid
bare. If the hydatid be
next withdrawn from the
cyst, it will be seen to
display a peculiar tremu-
lous motion, at the same
time coiling upon itself
wherever there is a free-
cut margin. Further
examination of portions
of the hydatid will show
that we have two distinct

Fig. 30.— Ectocyst, endocyat, and brood capsule of Echinococcus.
' From a Zebra. After Huxley.

CESTODA

117

membranes j an outer, thick, laminated, homogeneous elastic
layer (the ectocyst of Huxley), and an internal, thin, soft, granu-
lated, comparatively inelastic layer — the endocyst of the same
author. The terms are convenient. The ectocyst is structure-
less, consisting of a substance closely allied to chitine. For this
and other reasons it has been called the cuticular layer, but
the endocyst is the essential vital part of the animal, repre-
senting a huge compound caudal vesicle. In an hydatid from
the zebra, Huxley found that the endocyst was " not more than
^th of an inch in thickness, being composed of very delicate
cells of 5^" to in diameter, without obvious nuclei ; but
often containing clear, strongly refracting corpuscles, generally
a single one only in a cell/' Prof. Huxley adds : " These
corpuscles appear to be solid, but by the action of dilute acetic
acid the interior generally clears up very rapidly, and a hollow
vesicle is left of the same size as the original corpuscle. No
gas is developed during this process, and sometimes the
corpuscles are not acted upon at all by the acid, appearing then
to be of a fatty nature. A strong solution of caustic ammonia
produces a concentrically laminated or fissured appearance in
them. Under pressure and with commencing putrefaction a
number of them sometimes flow together into an irregular or
rounded mass."

The precise mode of development of the echinococcus-heads
or scolices has been a subject of lengthened discussion between
Leuckart and Naunyn. According to Leuckart the earliest
indication of the scolex consists of a slight papillary eminence on
the inner surface of the granular endocyst. After a short
period this prominence displays in its interior a vacuole-like
cavity, the latter being occupied, however, with a clear limpid
fluid. Its margins become more and more clearly defined, until
the cavity is by and by seen to be lined with a distinct cuticular
membrane. The papilla increasing in size, becomes at first
elongated or oval, eventually scoleciform, or even, perhaps, a
true echinococcus-head. Thus far the description bears out, in
a measure, the theoretical notions entertained by the older
authors ; but the developmental process does not stop here. The
scolex-development has now to sacrifice itself by developing in
its interior a brood of scolices or echinococcus-heads. In other
words, it becomes transformed into the so-called brood-capsules
of Leuckart and other authors. These structures were pre-
viously well known to Professors Erasmus Wilson and George

1 [° PARASITES OF MAN

Busk. Mr Wilson spoke of the capsule as " a delicately thin
■proper membrane, by which the Echinococci are connected with
the internal membrane of the acephalocyst " (' Med.-Chir.
Trans./ 1845, vol. xxviii, p. 21). Mr Busk described the
echinococcus-heads as " attached to a common central mass by

Fig. 31. — Group of Ecliinococcus-heads, from an hydatid found in the liver of a sheeg.
Magnified about 25 diameters. From a drawing by Professor Busk.

Fig. 32. — Three brood-capsules, containing Echinococcus-hends. Magnified 76 diameters.
After Professor Erasmus Wilson.

short pedicles, which appear to be composed of a substance
more coarsely granular, by far, than that of which the laminas
of the cyst are formed. This granular matter is prolonged
beyond the mass of Echinococci into a short pedicle common
to the whole, and by which the granulation is attached to the

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119

interior of the hydatid cyst." What Mr Busk here describes
as a granulation can only be equivalent to the brood-capsule
and its entire contents, but he elsewhere speaks of the capsule
itself as a " delicate membranous envelope." It should be
borne in mind that Busk's paper was communicated to the
Microscopical Society so early as the 13th Nov., 1844; being
published in the ' Transactions ' for that year.

In the completely developed state the echinococcus-heads
exhibit somewhat variable characters as to size and form, the
latter differences being, for the most part, dependent upon their
degree of contraction and vitality. In the perfect condition
they vary from the ^ to the
■reo" in diameter, being usu-
ally about the They are
solid, and when stretched out
exhibit an hour-glass-like
constriction at the centre of
the body, which divides the
scolex into an anterior part
supporting the rostellum and
suckers, and a posterior
part which has been com-
pared to the caudal vesi-
cle of ordinary Cysticerci.
The rostellum supports a
double crown of hooks, but
the disparity of the two series
is scarcely sufficiently marked
to render their distinction ob-
vious. The hooks of the
smaller row vary in size from

ToV t° 53o" °f an inch* whilst
those of the larger series are
from ^g" to jfc". In all in-
stances the root-processes are incompletely developed, and con-
sequently vary in thickness. They are, as Leuckart also has
stated, apt to exhibit abnormalities.

In regard to the development of the echinococcus-heads it
further remains for me to observe that a distinct water-
vascular system is recognisable in the scolices. By the
intervention of the pedicle of the scolex this system is
connected with the brood-capsule, and also with the vessels of

Fig 33.— Sep irate scolex, or ecliinococcus-head.
Magnified 500 diameters. After Huxley.

120

PARASITES OF MAN

the maternal endocyst. In the scolex there exists a circular
channel immediately below the rostellum, and this ring, on
either side, gives off two vessels which pass downwards in a
tortuous manner, internally, until they arrive at the pedicle
where they unite to form two channels, which latter are
continued into the vascular system of the maternal endocyst.
In the retracted condition their position, of course, becomes
very much altered, and they form loops on either side of the
central line which marks the space leading down to the
inverted head. Neither Prof. Huxley nor myself have seen
these vessels, which Leuckart observed in the scolex itself, but
Huxley discerned some apparently loose cilia in the granular
parenchyma of the body ; their longitudinal measurement being
about the ~ of an inch.

Fig. 34.— An Echinococcus brood-cnpsule (flattened by pressure). Magnified about 120 diameters.

From a drawing by Prolessor Busk.

As regards the production of " nurses " by the phenomenon
of proliferation, I can only remark that the endocyst is
primarily concerned. The secondary and tertiary vesicles must
be regarded as so many special bud-developments which, instead
of becoming brood-capsules, become daughter- vesicles and

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121

grand-daughter vesicles, constantly developing in their interior
secondary and tertiary brood-capsules and scolices, but some-
times, it would appear, developing neither the one nor the other.
This is the view of Naunyn, which is somewhat opposed by
Leuckart, who holds that the vesicles ordinarily arise from within
the layers of the ectocyst. Speaking of these daughter-hydatids
Leuckart remarks that " Naunyn denies that they take their
origin between the lamellae of the mother bladder — a fact, how-
ever, which, in agreement with Kuhl and Davaine, I have seen
more than once and have followed out step by step." For my
own part I incline to the belief that the process as observed by
Leuckart is exceptional, and that under ordinary circumstances
it occurs as Naunyn has described it. Thus the long and short
of the whole matter appears to be that the endocyst is capable
of forming solitary scolices. Some of the scolices become
differentiated to form brood-capsules, a portion of whose indi-
vidual echinococcus-heads may, in their turn, become secondary
brood-capsules, whilst others fail to become either scolices or
brood-capsules. It accords with our knowledge of the general
plan of development to believe that the daughter and grand-
daughter hydatids are likewise peculiarly modified scolices.
They are, in short, buds of the endocyst.

The distribution of hydatids throughout the organs of the
bearer, and their prevalence in particular countries, has espe-
cially engaged my attention. I have personally examined
upwards of a thousand preparations of entozoa in our public
collections ; and of these, 788 are preserved in the anatomical and
pathological museums of the metropolis. By this inspection I
have obtained a tolerably accurate knowledge of the pathology,
localisation and effects produced by the presence of bladder-
worms in at least 200 unpublished cases of hydatid disease.
Most of our museums exhibit one or more specimens that are
unique. After making certain necessary deductions, I find that
I have 192 new cases to add to the 135 cases of hydatid
disease that I had previously recorded, affording a total of 327
cases available for statistical purposes. If an analysis of these
cases be made and compared with the statistics furnished by
Davaine, and if the whole be reduced to the lowest number of
practically available terms, we at length obtain a result which,
although it may be only approximatively correct, is nevertheless
of much practical, value and significance. The statistics in
question stand as follows :

122

rAKASITES OF MAN

Organs affected.

Davaine.

Cobbold.

Total.

165

161

326

Lungs

o o
DO

373

327

700

In the main Davaine' s table and my own show a remarkable
correspondency, as is seen in the numbers referring to hydatids
of the liver, heart, and bones respectively. Where our results
do not correspond the explanation of the discrepancy is suffi-
ciently simple. The abdominal cases here credited as such in
Davaine's table are placed by him under pelvis, whilst the
abdominal cases in my own table not only include the pelvic
hydatids, but also two spleen cases, and nineteen others from
the peritoneum and intestines.

As the facts here stand, the liver cases comprise nearly
46^ per cent. In a large number of cases the entozoon has
taken up its abode in organs of vital importance. If statisti-
cians and officers of health would obtain an adequate concep-
tion of the fatal capabilities of parasites, they should consider
these data. In 6 per cent, of all these cases the bladder worm
has found its way into the brain, and of course proved fatal to
the bearers ; in about 3^ per cent, more they took up their
residence in the heart, also proving fatal ; whilst of all the
other cases put together I reckon that not less than 15 per cent,
were concerned in bringing about the death of their hosts. I
probably underrate the fatal capabilities of echinococcus disease
when I express the conviction that hydatids prove fatal to 25
per cent, of all their human victims.

The recently published analysis of 983 cases by Dr Albert
Neisser affords similar results. Of these, 451 were referable
to the liver, or 45*765 per cent. The other cases, reduced as
above, show in the main a similar correspondency.

It may be asked if these facts afford us any assistance in
determining the amount of injury that we, as a people, sustain
either directly or indirectly from hydatids. On carefully
reviewing all the data before me, I may say that it is difficult

CESTODA

123

to draw very precise conclusions ; albeit it is not mere guess-
work when I assert that in the United Kingdom several hun-
dred human deaths occur annually from this cause. In some
other countries the proportion is far greater ; the offc-quoted
case of Iceland, where the disorder is fatally endemic, still
standing at the head of the afflicted territories.

Our Australian colonies are probably entitled to the next
place of distinction in this respect. We have strong and
recent evidence of the truth of this statement. Thus a writer
in the ' Australian Med. and Surg. Eeview ' says : " This dis-
ease is becoming unpleasantly frequent, and at present we have
no reliable mode of treatment, either theoretical or empirical. "
Another writer observes (f Melbourne Argus/ May 18th, 1874),
" Hydatid disease is endemic in this colony ; and, though not
so constantly met with as in Iceland, we may probably claim
the doubtful honor of holding the second place in the list of
countries so affected/-' In the ' Argus' for June 20th of the
same year, another writer refers to the frequent notices of
cases of hydatids published in the various local newspapers. A
retired medical man, the late Mr J. P. Eowe, writing in the ' Mel-
bourne Leader ' (Sept. 7th, 1872), incidentally remarked on the
" notable increase of hydatid disease in the human subject/'
Again, still more satisfactory evidence is afforded by a reviewer
in the ' Leader ' of the 31st January, 1874. Commenting on
my manual, he not only takes occasion, to speak of the preva-
lence of hydatids generally, but also supplies that kind of
accurate statistical evidence of which we so much stand in need.
He gives the following table, showing the number of deaths
from hydatids in Victoria for eleven years. It is instructive in
many ways.

Years.

Males.

Females.

• Total.

1862

1863

1864

1865

1866

• 25

1867

1868

1869

1870

1871

1872

Total deaths in eleven years

125

200

124

PARASITES OF MAN

To employ the writer's own words, "this mortality gives only
a faint notion of the extreme prevalence of hydatids in Victoria,
since numbers of cases are cured by tapping, and otherwise by
medical treatment, or by spontaneous bursting of the cysts."
Hydatids are often found post mortem where their presence has
never been suspected during life. " To meet with hydatids as a
cause of deranged health is now a matter of daily expectation
with every medical practitioner." Lastly, Dr Dougan Bird, in
his able brochure on ' Hydatids of the Lung/ fully confirms
these statements, remarking that the rich and poor of the Aus-
tralian metropolis suffer just as much from hydatids as do either
the shepherds of the western plains, or the miners of Ballarat
and Sandhurst.

Such are the facts from Australia. As regards home evi-
dence, so far as T am aware, little or nothing has been done
towards securing an accurate estimate of the mortality in
England from echinococcus disease. The reports of the Regis-
trar General give no sufficient sign. The explanation is not far
to seek, since for the most part hydatids are either classed
with diseases of the liver, or with those of the other organs in
which they happen to have been present.

One of the most valuable contributions to our knowledge of
the prevalence of hydatid disease affecting animals is that
supplied by Dr Cleghorn, from a statistical table constructed
by the executive commissariat officers stationed at Mooltan.
The record in question shows that out of 2109 slaughtered
animals, no fewer than 899 were affected with hydatid
disease. This is equal to more than forty-two per cent. In
the majority of cases, both the lungs and liver were affected,
cysts were found 829 times in the liver and 726 times in the
lungs. In a few instances they were present in the kidneys,
and also occasionally in the spleen. The inference from all
this is that in India, if not elsewhere, the echinococcus disease
is much less common in man than it is in animals. The
explanation is simple enough, since cattle have more ready access
to, and less scruple in partaking of filthy water and food in
or upon which the eggs of the Taenia echinococcus abound.

Into purely professional questions connected with the treat-
ment of the echinococcus malady I do not here enter ; never-
theless, in connection with hygiene I may observe that the
prevalence of hydatids in any country is strictly dependent
upon the habits of the people. The close intimacy subsisting

CESTODA

125

between the peasantry and their canine companions is the
primary source of the endemic j and where dogs are not kept,
it is well nigh impossible that the disease should be contracted.
The fact that every Icelandic peasant possesses, on an average,
six dogs, and that these dogs share the same dwelling (eating
off the same plates and enjoying many other privileges of
intimate relationship) sufficiently explains the frequency of
hydatids in that country. According to Krabbe, the sexually
mature Taniae occur in 28 p. c. of Icelandic dogs, whereas in
Copenhagen he found it twice only in 500 dogs examined. In
his work (quoted below, p. 58, or Fr. Edit., p. 60) Krabbe
comments on a sensational passage which, in my introductory
treatise (p. 283), I had quoted from a popular memoir by
Leuckart (' TJnsere Zeit/ s. 654, 1862). The practitioners
whom we had spoken of as " quacks " are mostly homoeopaths ;
and it appears that even those who are not in any legal sense
professional men " treat their patients much in the same way
as ordinary medical men." It simply comes to this, that,
instead of dog's excrement forming with the aforesaid " quacks "
a conspicuous or common remedy (as Leuckart's description had
led me to infer), this nasty drug is now rarely administered,
and by the grossly ignorant only.

Up to the present time no person has seen the Tcenia echino-
coccus in any English dog which has not been previously made
the subject of experiment, but considering the prevalence of
hydatid disease amongst us, there can be no doubt that
English dogs are quite as much if not more infested than conti-
nental ones. Probably, at least one per cent, of our dogs harbour
the mature tapeworm. Certainly a great deal of good might
accrue from the acquisition of more extended evidence respect-
ing the prevalence of this and other forms of entozoa infesting
man and animals in this country.

From Schleisner's table it appears that hydatids are more
frequent in women than in men. Apparently, it is not so in
Australia. As regards Iceland the explanation must be sought
for in the different habits of life. No doubt, water used as drink
by women is constantly obtained from supplies in the immediate
neighbourhood of dwellings, and in localities to which dogs
have continual access. The comparative rarity of the echino-
coccus disease amongst sailors is not so much dependent upon
the circumstance that seamen's diet usually consists of salted
provisions, as upon the fact that these men can seldom have

126

PARASITES OF MAN

opportunities of procuring water from localities where dogs
abound. In regard to water drinking, there is ground for
believing that the addition of a very little alcohol' is sufficient
to destroy the six-hooked embryos of Taenia echinococcus whilst
still in ovo ; and there is no doubt that water raised to a tem-
perature of 212° Fahr. will always ensure the destruction of the
larvae. Boiled water by itself is by no means palatable. The
reason why the upper classes comparatively seldom suffer from
hydatids may be attributed to the circumstance that those few
who drink water take the very proper precaution to see that it
is either " pump " or fresh spring water in which no living
six-hooked embryos are likely to exist. So far as hydatids are
concerned, wine and beer drinking is preferable to water-
drinking ; yet if water is carefully filtered no evil of the para-
sitic kind can possibly result from its imbibition. An ordinary
charcoal filter will effectually prevent the passage of the ova,
since their diameter is nearly ~ of an inch.

From what has been stated it follows that personal and
general cleanliness are eminently serviceable as preventions
against infection, but to ensure perfect success other precautions
must be exercised, especially in relation to our contact with
and management of dogs. Leuckart puts this very clearly when
he says : — " In order to escape the dangers of infection, the
dog must be watched, not only within the house, but whilst he
is outside of it. He must not be allowed to visit either
slaughter-houses or knackeries, and care must be taken that
neither the offals nor hydatids found in such places are acces-
sible to him. In this matter the sanitary inspector has many
important duties to perform. The carelessness with which
these offals have hitherto been disposed of, or even purposely
given to the dog, must no longer be permitted if the welfare
of: the digestive organs of mankind is to be considered.
What blessed results may follow from these precautions may
be readily gathered from the consideration of the fact that,
at the present time, almost the sixth part of all the inhabitants
annually dying in Iceland fall victims to the echinococcus
epidemic" (1. c, s. 654). Similar measures had previously
been recommended in less explicit terms by Kiichenmeister,
who in effect remarked that the principal thing was to ensure
the destruction of the echinococcus vesicles. He also recom-
mended the expulsion and annihilation of the Ta-nia echinococcus.
In order to carry out this idea, it was suggested by Dr Leared

CESTODA

127

that every dog should be periodically physicked, and that all
the excreta, tapeworms included, should be buried at a con-
siderable depth in the soil. I advised, however, that in place
of burying the excreta, they should, in all cases, be burnt. I
had, indeed, long previously urged this measure (in a paper
"on the Sclerostoma causing the gape-disease of fowls/'
published in 1861), with the view of lessening the prevalence of
entozoa in general, whether of man or animals. The rule I
suggested stood as follows : — All entozoa which are not preserved
for scientific investigation or experiment should be thoroughly
destroyed by fire, when practicable, and under no circumstances
whatever should they be thrown aside as harmless refuse. As
an additional security I recommended that boiling hot water
be occasionally thrown over the floor of all kennels where dogs
are kept. In this way not only would the escaped tapeworms
be effectually destroyed, but also their eggs and egg-contents,
including the six-hooked embryos. These measures were again
advocated at the Cambridge Meeting of the British Association
in 1862, and also more fully in a paper communicated to the
Zoological Society, during the autumn of the same year (' Pro-
ceedings/ vol. xxx, pt. 3, pp. 288, 315).

As the scope and tendency of this work preclude the textual
admission of clinical details, I must limit my remaining obser-
vations to the pathology of hydatid disease. At very great
labor, pursued at distant intervals during a period of ten years,
I sought to ascertain the probable extent and fatality of this
form of parasitism in England, by going over such evidence as
our pathological museums might supply. Although, from a sta-
tistical point of view, the investigation could hardly be expected
to yield any very striking results; yet clinically viewed the
study was most instructive. The evidence which I thus pro-
cured of numerous slow and painful deaths from echinococcus
disease, further stimulated me to place a summary of the facts
on record. Physicians, surgeons, scientific pathologists, and
veterinary practitioners are alike interested in the study of
hydatid disease ; and I had not proceeded far in my careful
investigation before it became evident to me that very great
practical results would ensue if, in this kind of effort, the prin-
ciple of division of labor had full play. At all events, within
these museums lie concealed a mass of pathological data which,
although well within reach, have not been utilised to the extent
they ought to have been.

128

PARASITES OF MAN

As a student of parasites for some thirty years, I must
without offence be permitted to protest against the too
frequent omission of parasites in statistical evidence as a
cause of mortality. From facts within my own knowledge 1
can confidently assert that parasites in general, and hydatids in
particular, play a far more important part in the production of
disease and death than is commonly supposed. In saying thus
much, however, I am not insensible to the fact that, in recent
times, new methods of treatment combined with higher surgical
skill, have greatly tended to lessen the fatality of this affection.
In this connection I would especially refer to the recorded
experiences of an able colonial surgeon, Dr MacGillivray, as
made known in the pages of the ' Australian Medical Journal/
The able surgeon to the Bendigo Hospital, treated as in-patients,
from 1862 to 1872, inclusive, no fewer than seventy-four cases
of hydatid disease. He operated on fifty-eight of them. Two
patients were tapped for temporary relief (as they were dying
of other diseases) ; and of the remaining fifty-six only eleven
died. No fewer than forty-five were discharged cured — a fact
redounding largely, I should think, to the credit of Australian
surgery.

In reference to museum evidence I have no hesitation in
saying that the pathological collections in the metropolis
abound in rare and remarkable illustrations of hydatid disease ;
most of the preparations being practically known only to such
few members of the medical profession as have been at some
time or other officially connected with the museums. Not
without justice, curators often complain that their work and
catalogues are turned to little account. As a former conservator
of the Edinburgh University Anatomical Museum (1851-56), and
subsequently as museum- curator at the Middlesex Hospital
Medical College, I am in a position to sympathise with them.
Valuable, however, as the catalogues are, it is often necessary
to make a close inspection of the preparations in order to arrive
at a correct interpretation of the facts presented.

Although the entozoal preparations in the museum attached
to St Bartholomew's Hospital are, comparatively speaking,
few in number, there are some choice specimens of hydatid
disease. There is a remarkable case in which hydatids invaded
the right half of the bones of the pelvis ; death resulting from
suppurative inflammation of the cysts. This patient, a woman,
had also another hydatid cyst which was connected with the

CESTODA

129

ovary. Amongst the series contributed by Dr Farre, there is
a case represented where a large cyst containing numerous
hydatids " occupied the pelvis of an infant and produced reten-
tion of urine/' which ultimately proved fatal. There are also
several fine examples of hydatids from the omentum (Dr Farce's
case), besides a good specimen of acephalocysts connected with
the vesiculaa seminales. There are two other cases in which
these larval entozoa were passed with the urine. At the time
when I made my inspection, the entire series represented
twenty-five separate cases, of which only one appears to have
been published in detail (Mr Evans's case, ' Medico- Chirurgical
Transactions,' 1832). In addition to the above, I must not
omit to particularise two instructive preparations illustrative
of a case in which an hydatid was lodged in the right
half of the cerebrum. This was from a girl in whom head
symptoms showed themselves a year before death, and in whom
there was partial hemiplegia of the left side. I may add that
there is also in the series a doubtfully genuine example of
hydatids of the breast.

The collection in connection with the Westminster Hospital
contains several highly interesting specimens of entozoa (one
of which I believe to be altogether unique), but it is by no
means rich in the matter of hydatids. Out of a score of
preparations of parasites of various kinds, only four (apparently
representing the same number of cases) are hydatids, all of
which appear to have been connected with the liver. Two are
certainly so, one of the latter (Mr Holthouse's case) showing
calcareous degeneration.

The museum connected with St Mary's Hospital Medical
School, in addition to several liver cases, contains one interest-
ing example of hydatids of the lung (Dr Chambers's case), and
also three valuable preparations illustrating Mr Coulson's
remarkable case of hydatids affecting the tibia. One of the
preparations shows the bone itself, which was eventually removed
at the joint, the operation having been performed by Mr
Spencer Wells.

Here, perhaps, it will not be out of place to mention as a
fact of special clinical interest that I have encountered records
of no fewer than nine other similar cases where hydatids
have taken up their abode in the tibia, generally selecting
the head or upper part of the bone. Some of my notes have
been mislaid, but, speaking from recollection, one of the

130

PARASITES OF MAN

choicest specimens which I have examined is that contained
in the pathological museum of the Nottingham Hospital.

When I first went over the collection of the Middlesex
Hospital Museum, I found it to contain fifty-four preparations
of entozoa, of which some fourteen only were true hydatids,
representing as many separate cases. There are now upwards
of a score of preparations of hydatids, several of the cases
having already had ample justice done to them by Dr Murchison
in his well-known memoir (fEdinb. Med. Journ./ Dec, 1865).
Amongst the most interesting preparations I would especially
call attention to two fine and genuine specimens from the
kidney, another very large example of an hydatid situated
between the bladder and rectum, a simple acephalocyst removed
from the orbit (Mr Hulke's case), and the hydatid removed
from the axilla by the late Mr Charles Moore. There is a jar
containing hundreds of hydatids that were taken from the
thoracic cavity of a dissecting-room subject, who was reported
to have died of phthisis ; and there is another preparation of an
hydatid of the heart, which also proved fatal, without there
having been the slightest suspicion entertained as to the true
nature of the disease. For this fine preparation the museum
stands indebted to Dr Moxon, of Guy's Hospital. Several of
the liver cases are particularly instructive ; but amongst the
specimens presented by Mr Mitchell Henry is a small bottle full
of minute hydatid vesicles, all of which were removed from the
interior of the tibia. The history of this case has been lost ;
and, unfortunately, the bone from which the parasites were
taken does not appear to have been preserved.

The museum connected with King's College contains at
least a dozen good specimens of liver hydatids, several of the
cases being of special interest from a pathological point of
view. There are two remarkably fine examples of hydatids
contributed by Dr Hooper, the parasites in one case affecting
the spleen, and in the other involving the ovary and uterus.
The spleen contained numerous encysted hydatids, whilst the
uterine organs exhibited "an immense collection" of the same
growths. In this place, also, I may refer to an hydatid-like
entozoon, taken from a cyst in the ovary of a female who had
been under the care of Dr Johnson (I860). It is, apparently, a
genuine example of the slender-necked hydatid (Cysticercus
tenuicollis) ; and if so (as might be determined by dissection),
is, so far as I aware, the only specimen of the kind in existence

CESTODA

from the human bearer. There is a renal hydatid (presented
by Dr Pass, of Warwick) which was obtained from a lunatic,
its presence being "quite unsuspected during life." Amongst
the liver cases (the majority of which are from Dr Hooper's
collection), there is one enormous hydatid that was obtained
from a young woman who had died during a fit of laughter.
The tumour had pushed the diaphragm up to a level with the
fourth rib ; and it is stated that, on puncturing the cyst, the
fluid contents were ejected "in a jet nearly two feet high."
There is one case represented where numerous hydatids were
expectorated after hepatitis, whence it was concluded that they
were originally connected with the liver. There is a large
solitary hydatid that was removed from a young female who
died of phthisis, and in whom the consequent swelling had
formed in the neighbourhood of the navel. Especially instruc-
tive, also, from a clinical point of view, is a case of peritoneal
hydatids where the tumours had been diagnosed to represent a
case of extra-uterine fostation. It appears that there were two
cysts, one of them being connected with the uterus. Two of
the enormous hydatids taken from these cysts are preserved in
the collection of the Anatomy School of Oxford. Several of
the preparations show to perfection the stages of natural cure
produced by calcareous degeneration ; and there is one liver
showing three of these so-called ossified cysts. The disease in
this case proved fatal.

Most of the entozoa displayed in the Charing Cross Hospital
Museum have been contributed by Dr Wiltshire, the series
being particularly strong in tapeworms. There are four cha-
racteristic examples of hydatids of the liver, representing as
many separate cases. Two were from abscesses of this organ.
In one of these, Mr Canton's case, the hydatid was, I believe,
expelled after operation ; but in the other example (presented
by Mr Rose, of Swaffham) the parasite was evacuated from an
abscess, which burst of itself, externally.

In the museum at University College, I examined sixteen
preparations of hydatid disease, representing almost as many
distinct cases. One is a wax model. Bight of the specimens
were from the liver, five from the abdomen (including those of
the omentum and mesentery), two from the lungs, and one
from the heart. The model displayed ordinary hydatids of the
liver bursting into the lungs. The mesenteric example is par-
ticularly fine, whilst that from the omentum is undergoing cal-

132

PARASITES OF MAN

careous degeneration. Probably the most interesting of all is
the example showing an hydatid lodged in the septum of the
heart. This was from a middle-aged female, who died sud-
denly whilst pursuing her ordinary domestic avocations.

The museum of the Royal College of Surgeons contains a
fine collection of parasites, its chief strength in this respect
being due to the special series of entozoa. Were visitors to
judge by the contents of the catalogue of this series (which I
prepared some years ago at the instance of the Council of the
College), they might be led to suppose that the hydatids were
only feebly represented. Out of nine preparations of hydatids
in this section, only six have come from the human body.
However, scattered throughout the collection, I found that
there were no fewer than thirty-five preparations of hydatids
belonging, apparently, to as many as thirty separate cases.
Omitting, for the present, all mention of those derived from
animals, I ascertained that, of the thirty human cases, thirteen
were referable to the liver, four to the abdomen, three to the
lungs (one of which was originally connected with the liver),
and two to the brain. Five were of uncertain seat. With the
abdominal cases we may also include one case of hydatids of
the spleen, and another where these organisms were found in
the region of the bladder. There is a characteristic breast
case. One of the original Hunterian cases (in which ' ' a pro-
digious number of hydatids were found in the sac of the liver
and dispersed throughout the cavity of the abdomen ") appears,
though it is not expressly so stated in the catalogue, to have
been regarded as an ordinary example of abdominal dropsy.
In one of the three lung cases two small hydatids were sepa-
rately expectorated at an interval of about a month. This
occurred in a female.

I may here incidentally remark that many cases are on record
where abdominal hydatids have been overlooked, the patient
being supposed to be suffering from ascites. One such in-
stance took place a few years ago at the Middlesex Hospital.
I well remember a similar case of supposed hydrothorax, where
the post-mortem examination revealed the presence of immense
numbers of these formations occupying the right side of the
chest. This case occurred at the Norfolk and Norwich
Hospital, at the time when I was a student there, some
thirty-five years ago.

The pathological collection connected with St George's

OESTODA.

133

Hospital displays several good hydatid preparations, the entire
series representing at least twenty-two separate cases. Of
these, fifteen are referable to the liver, that is, if we include Dr
Dickinson's case, already published, where hydatids were found
within the hepatic duct. There are two renal cases; also one
from the brain (Dr Dickinson's case), and another where an
hydatid was expectorated. Besides these, there are three other
highly characteristic examples of echinococcus disease affecting
the region of the neck, breast, and axilla respectively.

The museum of the London Hospital Medical School contains
a large collection of parasites. Out of fifty-seven preparations
of entozoa, I found twenty-two referable to hydatids; and, so
far as I could gather, all of them belonged to different cases.
Only one case seems to have been published in detail. This,
though a very old preparation, is a fine example of an hydatid,
nearly three inches in length, occupying one of the cerebral
hemispheres (' Edinb. Med. Journ./ vol. xv). There is a second
brain case, where the vesicles were of small size, but very
numerous. Of the other twenty cases, fourteen belong to the
liver, two to the spleen, one to the lung, one to the uterus ;
one being a very large hydatid of doubtful seat, and another
being referable to the lumbar region, where it formed a tumour
containing "a, large number of small hydatids." Amongst the
more remarkable specimens is that described in the MS. cata-
logue as " a true hydatid cyst developed in connection with the
broad ligament." This preparation, unique of its kind, shows
no trace of the ovary, which, indeed, seems to have disappeared
altogether. One of the liver cases should rather be classed as
abdominal, since the large cyst is situated between the dia-
phragm and liver, pressing upon the latter organ below and also
upon the lung above, but apparently not involving either of
these viscera structurally. Another very striking case is that
in which there is an external opening communicating with the
cyst in the liver, and an internal opening through the dia-
phragm communicating with the lungs and bronchial tubes.
The patient had actually coughed up liver hydatids by the
mouth, and had passed others through the right wall of his
abdomen. There is another liver case in which the hydatids,
in place of escaping externally, had gained access to the inferior
cava; and if I understand the MS. record rightly, in the same
patient a second hydatid communicated with the portal vein,
and a third with the hepatic vein. Lastly, I must add that

134

PARASITES OF MAN

there is yet another fine preparation of liver hydatids, occurring
in a lad, nineteen years of age. He had, it seems, met with
<fa slight accident, and died with obscure head symptoms;" but
the odd part of the case is that at the post-mortem examina-
tion there was positively nothing found that could explain the
patient's death. He was under the care of Mr Luke (1834).

Comparatively recently I inspected the collection at St
Thomas's Hospital, which I found to be particularly rich
in entozoa of various kinds, especially tapeworms and hydatids.
I encountered seventy- six preparations of internal parasites ;
and of these, forty-two were of the hydatid kind, representing
at least thirty-three different cases. I say "at least/' because
it is often impossible to decide in instances where no history of
the specimens can be obtained. Thus, there are three similar
preparations of hydatids passed by the urethra, and, from their
appearance, I judge them to have come from one and the same
patient ; yet there is no statement in the catalogue to that
effect.

Of the thirty-three cases of hydatids represented in this
museum, I reckoned eighteen as referable to the liver, two to
the brain, two to the bones, two to the urinary organs, and one
to the lung, spleen, uterus, and soft parts of the thigh respec-
tively. There are also three that may be classed as peritoneal.
There is another choice example in which the disease cannot be
referred to any particular organ. I allude to Dr Peacock's case,
already published (' Pathological Transactions,' vol. xv), where
the lungs, liver, heart, spleen, and some other organs, were all
occupied by hydatid formations. As an instance of extensive
visceral infection by Echinococci in the human subject, I believe
this case to be unique. The brain hydatids are particularly
fine. In the specimen presented by Mr Boot, of Lincolu, the
hydatid, two inches in diameter, is lodged in the anterior horn
of the left ventricle. One of the peritoneal cases is remarkable
for the amount of forward displacement of the pelvic viscera,
caused by four or more hydatids, each of them nearly as large
as a cricket-ball. Amongst the abdominal cases I have included
a recent preparation, to which Mr Stewart has called my
attention. The hydatid in question, of the size of a large
lemon, existed near the fundus of the bladder, its walls being
one third of an inch in thickness, and forming an unusually
firm tumour. Of all the fine specimens of hydatids in the col-
lection, however, none have struck me so much as those affect-

OESTODA

135

ing the bones. There is a humerus, taken from a man thirty-
four years of age, in which the shaft is occupied throughout by
small hydatids that have destroyed almost all the cancellous
structure ; in some places, also, the absorption of the cortical
layer has gone on to such an extent as to have left little more
than the periosteum. Of course, the bone was at last fractured
easily. It is a beautiful specimen ; and the existence of Echi-
nococcus-heads was proved by microscopic evidence. Scarcely
less interesting are two preparations illustrative of Mr Travels
case of a man, thirty-eight years of age, in whom numerous
small hydatids occupied both the head of the tibia and the lower
end of the femur. Each set of parasites freely communicated
with the knee-joint, necessitating amputation of the limb.

The very large museum connected with Gny's Hospital is
rich in hydatids. When, some time ago, I spent several days
in going over the collection, I examined seventy-six prepara-
tions, representing apparently seventy separate cases of this
affection. Amongst the noteworthy specimens one lung
hydatid was intimately associated with a thoracic aneurism,
two others being connected with the pleura ; and of seven
abdominal cases, five were connected with the peritoneum, one
with the mesocolon, and one with the aorta. This last-men-
tioned instance occurred in a woman of sixty years, who, until
her death, was treated for dropsy. She complained of incessant
pain, which was only relieved when she rested on her hands
and knees. Of the three cases affecting the heart one has
been published (Mr Henderson's), where the patient, a girl of
nineteen years, died suddenly whilst in the apparent enjoyment
of perfect health. In one of the other two cases (Mr May's, of
Tottenham), the left lung was also involved. One case of
hydatid disease affecting the spinal column appears to have been
originally an ordinary liver case. In Mr Cock's example of
genuine mammary hydatids, the hooklets and echinoccocus
heads were detected ; but I am not sure that a similar
result of microscopic examination was obtained in the equally
interesting example of hydatids of the thyroid gland (also
removed by Mr Cock). There are five bladder cases, all
apparently genuine (of which one has been published) ; and
there are also five other cases referred in the catalogue to the
kidneys, of which I regard two as doubtfully parasitic in
character. Of three cases of hydatid growths occupying the
soft parts of the thigh, two were under Mr Bryant's care. The

136

PARASITES OF MAN

museum likewise contains an old preparation of hydatids of the
tibia, but its history has been lost. There are also two brain
cases, besides upwards of a score of more or less characteristic
and instructive cases of hydatids affecting the liver.

Scattered amongst the museums connected with the larger
provincial schools and recognised hospitals there must be a
great many valuable preparations of hydatid disease ; at all
events, I judge so from the inspection I have incidentally made
of a few of the collections.

Of eleven preparations of human hydatids which I observed
in the Cambridge Anatomical Museum, apparently representing
the same number of cases, seven were connected with the
liver and one with the lungs. Those hydatids displayed in the
" special series " of entozoa were of uncertain seat. From the
recently published and valuable e Notes ' by Dr Bradbury, I
have no doubt that considerable additions have been made to
the Cambridge Collection since my last visit.

The museum at Oxford contains some choice specimens of
hydatids, but I have only personally inspected a few of them.
In the absence of original notes, however, I am indebted to
the kindness of Mr W. Hatchett Jackson for supplying me with
several interesting particulars. The anatomical department of
the Oxford Collection shows from one particular case two
hydatids that were found " under the dura mater." In the
pathological department we find one hydatid from the liver of
a male subject, and also a preparation showing a number of
small hydatids that were " coughed up from the lungs of a
female." There are also in this department (Dr. Acland's)
two examples of hydatids from the diaphragm, apparently
belonging to two separate cases. One is described as a large
" hydatid in the diaphragm covered by the pleura," whilst the
other is spoken of as " springing from the diaphragm and
projecting into the sac of the pericardium." There is likewise
a preparation showing a number of small specimens of hydatids
that were passed per anum by a female. It is conjectured
that they came from the liver.

The small pathological museum attached to the Brighton and
Sussex Hospital is particularly rich in hydatids. Amongst
others, it contains preparations illustrative of the remarkable
case of hydatids in the region of the prostate, communicated
by Mr Lowdell, in the ( Lancet/ in 1846.

The comparatively large museum adjoining the Norfolk and

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137

Norwich Hospital displays a choice series of hydatids, chiefly
from the collection of the late Mr Crosse. That eminent
surgeon prepared a special set of specimens to illustrate the
process of natural cure by calcareous degeneration ; and I may
here, perhaps, be pardoned for mentioning that it was the study
of these and other entozoa in Mr Crosse's Collection, some
thirty or more years ago, that first drew my attention to the
phenomena of parasitic life. Illustrations of the helminths in
question are still in my possession. In one case (which is
instructive as indicating the possibility of death from the
simplest form and commonest habitat of an hydatid) a lad,
twelve years old, received a slight blow from a playmate. Some-
thing gave way, and death speedily followed. It was found by
post-mortem examination that a solitary liver hydatid, rather
larger than a cricket-ball, had been ruptured. Although the
case is almost unique, it is nevertheless by no means pleasant
to reflect upon the fact that under similar circumstances a slight
blow might prove fatal to any one, no matter in what internal
organ the bladder worm happened to be situated.

Before concluding my summary notice of the human
hydatids contained in the metropolitan and certain other
museums, there is an interesting literary contribution that I
cannot pass unnoticed. In the November number of the
' Indian Medical Gazette' for 1870 an article occurs in which it
is stated that the Calcutta Medical College Museum contains
eighteen specimens of hydatid cysts of liver. This fact was, it
seems, originally adduced to show, not the frequency, but rather
the rarity, of the occurrence of hydatids in India. However,
from a valuable communication by Dr James Cleghorn, which
was published in the same periodical for the following March, it
appears that hydatids of the liver are much more common in
India than is generally supposed. This, he says, is owing to
the circumstance that many of the so-called cases of tropical
abscess are neither more nor less than examples of hydatid cysts
that have suppurated. Besides Cleghorn's evidence, we have
the previous testimony of the Inspector General I. M. D.,
whose Report for 1868-69 I have already referred to in con-
nection with Cysticercus in beef. He says : u Daring some
three months' regular observation of the animals killed at the
Commissariat slaughter-house here, at least 70 per cent, of the
beef livers may be calculated as thus affected. Cobbold,
writing of the Taenia echinococcus, says that ' this little tapeworm

]38

PARASITES OF MAN

infests only the dog and the wolf/ Therefore, considering the
immense number of pariah dogs fed on the refuse of animals
infected with hydatids, it seems more than probable that the
parasite must attain its strobila condition in their intestines, and
through them be eventually disseminated over the pastures on
which the cattle graze."

I now turn to a neglected phase of the subject from which
much practical instruction may be gathered. The consideration
of the pathological phenomena of hydatid disease as it affects the
lower animals is of high interest, and no prejudice should induce
any medical man from accepting such useful data as may be
gathered from this source. The facts of hydatid parasitism in
animals, though often peculiar, are, for the most part, of an
order similar to those presented in the human subject. If any
medical practitioner thinks it beneath his dignity to study the
pathology of the lower animals, the conduct of John Hunter in
this respect is a standing protest against such narrowness.

The museum of the Royal College of Surgeons of England
contains some of the finest specimens of hydatids from the lower
animals that are to be seen anywhere, the very choicest of them
having been selected by Hunter himself. That distinguished
man sought information from every available source, and
hydatids were for him of almost equal interest, whether found
in the body of a human being or in the carcase of an ox or an
ass. Now, at all events, neither pathologists nor sanitarians can
well afford to neglect comparative pathology ; and, for myself, I
am free to say that the yearly exposition to the students of the
Royal Veterinary College of the phenomena of parasitic life
amongst animals has brought with it an ever-increasing know-
ledge of the most curious and often unlooked-for information.
Some of the data thus supplied are quite remarkable. Let me
also add that my studies of the entozoa of wild animals have put
me in possession of particulars of high value in regard to
the larger question of the origin of epidemics. Beasts, birds,
reptiles and fishes, of every description, are liable to succumb
to internal parasites, and there is practically no end to the
variety of useful information to be obtained from this source.
I have collected materials almost sufficient for a separate treatise
on this department of the subject, but I fear I shall never have
either the time or opportunity to give the facts due publicity.
Here, for obvious reasons, I must for the most part restrict
myself to the hydatids properly so called.

CESTODA

139

Referring, in the first instance, to the hydatids of animals
that have the same mode of origin and exhibit the same general
characteristics as those found in man, I notice that four of
the metropolitan museums exhibit nine examples of liver Echi-
nococci. The Hunterian Collection shows specimens of this
kind from the pig, monkey, zebra, and lion. The museum at
St Bartholomew's Hospital contains two examples from the pig
and one from a cow ; whilst the animal liver-hydatids preserved
in the King's College and Guy's Museums, respectively, are from
the pig and sheep. That from the latter is partly calcified.
Respecting animal hydatids affecting the lungs, the Cambridge
Museum exhibits a simple acephalocyst from a monkey, and
the Guy's Hospital Museum shows a pulmonary hydatid from
the kangaroo. In the museum at Oxford, Dr Acland's
(pathological) department shows a preparation of " one large
echinococcus cyst from the abdomen of a baboon," whilst Dr
Rolleston's department (anatomical) displays the echinococcus
itself from the " cavity of the abdomen of the same
animal." The collection also contains a variety of other
bladder worms from different animals. The Hunterian Museum,
Lincoln's Inn, exhibits four or five alleged examples of hydatids
from the kidney of the sheep, besides another from the spleen.
Some of these are of very doubtful character. A cystic kidney
from the sheep, preserved in the London Hospital Museum, and
originally supposed to have been due to hydatids, is (as hinted
in the MS. catalogue) certainly not of parasitic origin. In
regard to the occurrence of hydatids in the heart of animals the
Hunterian series shows two good examples from cattle, whilst
the collection at University College exhibits one taken from the
wall of the left ventricle of a sow. This was presented by Dr
Elliotson.

In the museum of the Royal Veterinary College there are a
number of excellent preparations of true hydatids taken from
various animals, especially from cattle, swine, and sheep ; and
there are also many kinds of bladder worms which, though
often called " hydatid " by veterinarians, have a totally different
origin from that of the true Bchinococci. The so-called gid-
hydatids (Ccenuri) and slender-necked hydatids (Gysticercus
tenuicollis) are of this description. Specimens of the poly-
cephalous brain hydatid, or Ccenurus, also exist in the museums
connected with St Bartholomew's, Guy's, and St Thomas's
Hospital Medical Colleges, as well as in both the anatomical

140

PARASITES OF MAN

and pathological departments of the Oxford Museum. Speci-
mens of large Ccenuri occurring in the soft parts of rabbits may
be seen in the Guy's Museum (presented by Mr Carpenter).
Similar characteristic specimens exist in the Oxford Collection,
labelled 0. cuniculi, obtained from the " masseter and infra-
spinatus 13 muscles of a rabbit. My private collection also
contains a recent addition of this remarkable hydatid, sent to
me by Mr Alston from Ayrshire. It is the only one I have
seen from Scotland. In the second half of this work these
Coenuri will again come under notice. Three examples of the
slender-necked hydatid (from a monkey and two sheep respec-
tively) may be seen in the Guy's and University College Collec-
tions, and there are several in the museum of the Royal
Veterinary College.

I cannot go out of my way to speak of other bladder worms,
except so far as to call attention to the heart of a bear preserved
in the museum at Guy's, the walls of which are crowded with
Cysticerci. That unique preparation ought to be carefully
examined and described. The Hunterian Museum contains two
magnificent specimens of hydatids affecting the bones of cattle.
In the one case a solitary vesicle occupies the shaft of the
humerus ; whilst in the other several " acephalocysts 33 have
taken up their residence within the cancellous structure of the
ilium.

In the matter of human mortality from hydatids I have
already supplied statistical evidence of the unenviable distinction
which our Australian colonies exhibit, and in addition to the
facts brought forward I may add that Dr Lewellin has
mentioned to me a fatal case in which an hydatid occupied the
whole length of the vertebral canal. The patient was under
Dr Annand's care. There could be no doubt as to the genuine-
ness of the case, as the spinal cyst was tapped during life,
when echinococcus hooklets were found.

Through Dr Lewellin I am also indebted to Dr H. B.
Allen, pathologist at the Melbourne Hospital, for the particulars
of a case of hydatids of the cerebrum, which are given as
follows :

" J. Q — , aged 15, was admitted into the Melbourne Hos-
pital on the 13th November, 1877, suffering from partial left
hemiplegia. He rapidly became insensible and died next day.
His mother furnished the following history.

"He had been woodcarting in the bush for a considerable

CESTODA

141

time, and while thus engaged eight weeks before admission
began to lose power in his left arm and leg j gradually the
paralysis increased, and he was taken home, where he remained
for six weeks. During this time he had every week an attack
of severe headache, and once he lost all sight for over half an
hour. Gradually the symptoms increased, and he was taken to
the hospital, but even then was able to walk with assistance
part of the way.

" At the autopsy, when the calvarium was removed, a large
cyst about four inches in diameter was found on the mid-
convexity of the right hemisphere of the cerebrum, slightly
towards its anterior part. It formed a marked prominence on
the anterior surface of the brain, and was bounded superficially
by the pia mater and arachnoid, which were neither noticeably
thickened nor adherent to the dura mater. On opening the
cyst it was seen to extend inwards and abut on the wall of the
lateral ventricle, and consisted of the ordinary gelatinous
membrane, studded internally with little granular eminences,
some pellucid, some opaque white. The contents were thin
limpid fluid. The brain tissues around presented scarcely any
induration. All other organs structurally healthy, congestion
being the only morbid condition present.

" The specimen is preserved in the hospital museum, which
contains two other preparations of hydatids in the brain, and
also an hydatid cyst of large size growing from the interior of
the frontal bone."

In concluding this account of hydatids I may remark that,
by the employment of sanitary measures, the disorder might,
in course of time, be thoroughly stamped out. What these
measures are I have already stated.

I need hardly say that the following bibliography by no
means exhausts the records of echinococcus disease. In Dr
Albert Neisser's recent monograph nearly a thousand separate
cases are quoted and classified. The monograph of Dr Hearn,
which is not mentioned in Neisseria work, also contains a
valuable bibliography.

English literature. Hydatids in general (Bibliography No.
20 a).— Ballard, E. (review of Henoch), 'Med.-Chir. Rev./
1854. — Bird, 8. D., ' On Hydatids of the Lung • their diagnosis,
prognosis, and treatment/ 2nd edit., Melbourne, 1877. — Bush,
" On the Nat. Hist, of the Echinococcus," ' Micr. Soc. Trans./
orig. series, vol. ii, 1849.— Budd, 0., ' Diseases of the Liver/

142

PARASITES OF MAN

Lond., 1845.— Oarmichael, E. (lecture), ' Dub. Med. Press/ 1840,
p. 9.1. — Gobbold, T. 8., ' Entozoa/ chap, vii and viii, 1864. —
— Idem, " On Hydatid Disease " (lecture), ' Lancet/ June, 1875,
p. 850. — Idem, " On Hydatid Diseases of Man and Animals"
(museum specimens), in a series of articles contributed to
' Brit. Med. Journ./ Oct., 1875, to Jan., 1876 ; fully reprinted
in the ' Veterinarian/ Feb., 1876. — Copland, J. (Art. " Hyda-
tids ") in his ' Dictionary/1 1848. — Davies, T., 'Lond. Med.
Gaz./ 1835. — Gairdner, J., and Lee, ' Edinb. Med. and Surg.
Journ./ 1844. — Goodsir, H. 8. D. (same ref., Lee's case), 1844.
— Goodsir, J. (same ref.), 1844. — Gross, 8. D., ' Elements of
Path. Anat./ chap, xv, Boston, TT.S., 1839. — Hawkins, 0.,
'Lancet/ 1833.— Hjaltelin, 'Edinb. Med. Journ./ 1867; see
also Dobell's ' Report on the Progress of Practical and Scien-
tific Medicine/ London, 1870. — Hodgkin, T., in his 'Lectures on
the Serous and Mucous Membrane/ 1838. — Kerr, W. (art.
"Hydatids") 'Cyclop, of Pract. Med./ 1833.— Leared, A.
(prevention), ' Med. Times and Gaz./ 1863. — MacGillivray, P.
H. (see below, miscell. cases). — Murchison G., in his 'Clinical
Lectures/ Lond., 1868, p. 54 ; 2nd edit., 1877. — Idem,
" Hydatid Tumours of the Liver ; their danger, their diagnosis,
and their treatment," ' Edinb. Med. Journ./ 1865. — Nettleship,
E., " Notes on the Rearing of Taenia echino coccus in the Dog
from Hydatids, &c," ' Proc. Royal Soc./ 1866, p. 224.— Rose,
0. B., " On the Vesicular Entozoa, and particularly Hydatids,"
'Lond. Med. Gaz./ 1833-4, p. 204.— Stephens, 'Lancet/ 1833;
the 'Veterinarian/ 1831, p. 284. — Thompson, T. (remarks),
'Lancet/ 1851. — Wilson E., "On the Structure, Classification,
and Development of the Echinococcus hominis," ' Med.-Chir.
Trans./ 1845. — Yates, G., " On Hydatid Disease," ' Assoc.
Med. Journ./ vol. iii, 1855.

Hydatids of the liver (Bibliography No. 20&). — Abercrombie,
T., 'Lond. Med. Journ./ vol. ii, p. 276, 1829.— Alison, 8. 8.,
'Lond .Med. Gaz./ 1844. — Barclay, 'Brit. Med. Journ./ Nov.,
1868, p. 494. — Barker, T. A. 'Lancet/ and ' Path. Soc. Trans/
1855.— Barlow, 'Lond. Med. Gaz./ 1857.— Beith, 'Path. Soc.
Trans./ 1852. — Bradbury, J. B. (six cases), 'Brit. Med. Journ.,'
Oct., 1874, pp. 526-558.— Idem, 'Brit. Med. Journ./ 1876,
vol. ii, p. 646. — Brinton, ' Lancet/ 1854. — Idem, ' Lancet/
1 858— Bristowe, T. 8., 'Path. Soc. Trans./ 1851.— Idem,
' Path. Soc. Trans./ 1858.— Broadbent, W. H., " Hydatids of
the Liver ; Paracentesis followed by free Incision ;" ' British

CESTODA

1 13

Med. Journ./ Nov. 30th, 1878— Brodie, B. G. (supposed),
'Lond. Med. Gaz./ 1828.— BrooJc, C, 'Lancet./ Feb., 1868,
p. 162— Buchanan, 'Surg. Med. Gaz./ 1861.— Budci, W.,
'Brit. Med. Journ./ 1859.— Chambers, T. K., ' Lond. Med.
Gaz./ 1846.— Cox, T., ' Lancet/ and ' Med-Chir. Trans./ 1838.—
Crosse, J. G., 'Lancet/ 1837.— Curling T. B., ' Med.-Chir.
Trans./ 1840.— Daly, 0. (supposed), ' Brit. Med. Journ./ 1859.
— Davies E., ' Path. Soc. Trans./ 1848. — Dickenson, ' Lond. Med.
Gaz./ 1861; 'Path. Soc. Trans./ 1862. — Duncan, A. (near the
portal vein), ' Edin. Med. and Surg. Journ./ 1808. — Duncan,
P. M. (several cases), ' Prov. Med. and Surg. Journ./ 1850-52
Elliotson, J., ' Lancet/ 1832. — Feam, 8. W. (immense cyst),
' Brit. Med. Journ./ Nov., 1868, p. 496.— Fletcher, T. B. E.,
' Prov. Med. and Surg. Journ./ 1846. — Freer, W. G., ' Lancet/
and 'Prov. Med. and Surg. Journ./ 184b.—Gaitskell, W. (1000
discharged) ' Lond. Med. Expository/ 1815. — Gulland, 'Edin.
Med. Journ./ I860.— Barley, J., ' Lancet/ May, 1866, p. 538,
and 'Med. Chir. Trans./ 1866. — Idem, in 'St Thomas's Hos-
pital Eeports/ 1877, p. 291. — Hastings, C, 'Brit. Med. Journ./
1858.— Beaton, ' Brit. Med. Journ./ Oct. 31st, 1874, p. 557.—
Eeckford, N., ' Brit. Med. Journ./ Sept., 1868, p. 332.— Eillier,
' Lancet/ and ' Path. Soc. Trans./ 1855. — Hutchinson, J.,
' Lancet/ Oct., 1862.— Inglis, A., ' Brit. Med. Journ./ 1 859.—
Beared A. (Hjaltelm's case), ' Path. Soc. Trans./ 1863. —
Logan (thousands present), 'Path. Soc. Trans./ and' Med. Times
and Gaz./ March, 1865, p. 243. — Lyon, E. (several cases),
' Prov. Med. and Surg. Journ.,' 1850. — Murchison, C. (rupture
through pleura), 'Path. Soc. Trans./ 1861. — Idem, ' Lancet,''
July, p. 75,1868. — Page, 'Brit. Med. Journ./ and 'Lancet/ Nov.,
1864. — Pavy, F. W. (expectorated), ' Med. Gaz./ 1851. —
Idem, 'Med.-Chir. Trans./ and ' Lancet/ Sept., 1866, p. 234.—
Peacock (two cases, expectorated), ' Lond. Med. Gaz./ and
' Lancet/ 1850. — Pemberton, O. A. (rupturing diaphragm),
' Prov. Med. Journ./ 1848. — Philipson, ' Brit. Med. Journ./
Oct. 31st, 1874, p. 557 '.—Pollock, J. (fatal), ' Path. Soc. Trans./
1854. — Idem (opening into lung), ' Lancet/ Jan., 1865, p. 63. —
Bees, G. O., ' Guy's Hosp. Eep./ 1848. — Idem (lecture), ' Lond.
Med. Gaz./ 1849— Richards, C. C, 'Lancet/ Jan., 1865,
p. 261. — Roberts, 'Lancet/ 1833. — Russell, J., ' Prov. Med. and
Surg. Journ./ 1851. — Sadler, M. T. (Cesarean section), ' Med.
Times and Gaz./ Aug., 1864, p. 141.— Salter, E, ' Path. Soc.
Trans./ I860— Savory, W. S. (letter), 'Lancet/ May, 1866,

144

PARASITES OF MAN

p. 410. — Shemuin, H. C. (fatal);' Edin. Med. and Surg. Journ./
1823.— Sibson, F., ' Lancet/ July, 1868, p. IQ.—Sloane, J.
(puncture), ' Brit. Med. Journ./ 1858. — Thompson H., ' Path.
Soc. Trans./ and ' Lancet/ 1858. — Trimnell, G. 0., ' Lond.
Med. Eepos./ 1821.— Ward, 8. H., 'Lancet/ 1868, vol. ii,
pp. 141, 305, and 474. — Wearne, V. (perforating diaphragm),
' Brit. Med. Journ./ July, 1864, p. 31. — Willcs (escaping by
gall-ducts), ' Path. Soc. Trans./ 1860. — Young, J., ' Edin.
Med. and Surg. Journ./ 1829.

Hydatids of the liver and other organs together (Bibliography
No. 20 c). — Beale, L. (kidney), 'Arch, of Med./ vol. i, p. 31,
1857 ; see also same case by Bristowe, 'Path. Soc. Trans./
1853.— Billing (lungs), 'Lond. M. and S. Journ./ 1831, p. 58.
— Griffith, J. W. (abdomen), 'Lond. Med. Gaz./ 1844. — Heslop,
T. P. (kidney), 'Month. Journ. of Med. Sci./ 1850.— Richard-
son (kidney, Dr Mackinder's case), 'Lancet,' 1855.

Liver cases occurring in America (Bibliography No. 20 d). —
Alexander, E. (200 present), ' Boston Med. and Surg. Journ./
1838.— Finnell, 'New York Med. Journ./ 1856, p. 216.— Minot,
T. (expectorated), Bost. Soc. for Med. Improv., 1859, and
'Brit. Med. and Surg. Journ./ 1860, p. 297.— Webber, J. E.,
'New York Med. Times/ 1853, and 'Bost. Med. and Surg.
Journ./ 1853, p. 126.

Hydatids of the lungs and pleura (Bibliography No. 20 e). —
Oholmeley, ' Guy's Hosp. Rep./ 1837. — Bowling, F., 'Australian
Med. Journ./ 1864.— Buffin, A. B., ' Beale' s Archives/ 1857,
vol. i, p. 253.— Hare, 'Path. Soc. Trans./ 1857-8.— Hill, J.,
'Med. and Philos. Comm./ 1784, vol. ii, p. 303. — Hutchinson,
J., 'Path. Soc. Trans./ 1854.— Kirhes, W. S., ' Med. Times
and Gaz./ 1851. — Beared, A., 'Path. Soc. Trans./ 1857. —
Peacock, 'Lancet/ 1850. — Ridge, J., 'Guy's Hosp. Rep./ 1836,
p. 507. — Rigden, G., ' Prov. Med. and Surg. Journ./ 1852. —
Smith, F. G., 'North Amer. Med.-Ohir. Rev./ 1858, p. 333.—
Todd, B. B., ' Med. Times and Gaz./ 1852.

Hydatids of the Kidney (Bibliography No. 20/.) — Adam*,
A. L., ' Lancet/ 1 864, p. 375. — Barker, T. H, 'Glasg.Med. Journ./
1855-6, p. 439. — Buncan, 'Liverpool Med. Journ./ 1834. — Dunn,
J., 'Lond. Med. Repos./ 1817.— Fussell, E. F., ' Lancet/ 1851.—
Lettsom (two cases), ' Trans. Med. Soc. of Lond./ 1789, p. 33.—
Ward, W., 'Lancet/ 1846.— Wilson, J. (lecture) 'Lond. Med.
Repos./ 1822.

Hydatids of the spleen, omentum, and abdominal cavity

CESTODA

145

(Bibliography No. 20 g.). — (Anonymous), 'Edin. Med. and Surg.
Journ./ 1819, p. 50.— Bailey, F., ' Lond. Med. Repos./ 1826.—
Bright, B. (remarks on cases) ' Guy's Hosp. Rep./ 1838. — Bryant,
T. (simulating ovarian disease), 'Guy's Hosp. Rep./ 1868,
p. 235.— Budd, G. (omentum), ' Med. Times/ 1838.— Idem (rep.
by Parsons), 'Brit. Med. Journ./ 1859. — Burman, ' Prov. Med.
Journ./ 1847. — Oroivther, C, ' Edin. Med. and Surg. Journ./
1826, p. 49.— Greenhow, K H, 'Lancet/ 1862.— Little, W. I.
(simulating ovarian disease), 'Brit. Med. Journ./ 1857. — Macleay,
K., 'Edin. Med. and Surg. Journ./ 1806.— Morley, J. (partly
pelvic), ' Lancet/ 1845. — Newman, W. (simulating ovarian
disease), ' Obstetr. Soc. Trans./ vol. iv, 1862. — Obre (peri-
toneal), ' Path. Soc. Trans./ 1854. — Ogle, J. (omentum), ' Path.
Soc. Trans./ 1860. — Simpson, A. B. (peritoneal), ' Edin. Med.
Journ./ 1861-62. — Simpson, J. Y., ' Assoc. Med. Journ./ 1854,
p. 137. — Thompson, T., 'Lancet, 1843. — Thompson, A. T.
(simulating ovarian disease), ' Lancet/ 1833.

Hydatids within the pelvic cavity (Biblioqeaphy No. 20 h). —
Birkett, /.(voided), ' Guy's Hosp. Rep./ 1851, p. 300. — Bryant,
T., ' Lancet/ 1865, pp. 566 and 589. — Gorrigan (ovarian), ' Dub.
Quart. Journ./ vol. i, 1846. — Crampton (ovarian), ' Dub. Quart.
Journ./ vol. ii, 1846. — Curling, T. B. (bladder), 'Med. Times
and Gaz./ 1863. — Farre, A., ' Lancet/ 1862. — Habershon,
' Path. Soc. Trans./ I860.— Hughes, ' Lond. Med. Gaz./ 1861.—
Hunter, T., ' Trans, of Soc. for Improv. of Med. and Chir.
Knowledge/ 1793, p. 34. — Jennings (simulating pregnancy),
'Dublin Quart. Journ./ 1855. — Lotvdell, 'Lancet/ 1846. —
Maunder, 'Lancet/ Sept., 1864, p. 351. — Sadler, M. T. (voided),
' Med. Times and Gaz.' 1865. — Simon, J. (voided), ' Lancet/
1853.— Wakley, 'Lancet/ 1863.— White, 'Med. Gaz./ 1842.

Hydatids of the heart and blood-vessels (Biblliogeaphy No.
20 i).— Bigger, 'Dub. Path. Soc' Rep. in ' Lancet/ 1830.—
Budd, G., ' Path. Soc. Trans./ 1839.— Goote, H, ' Lond. Med.
Gaz./ 1854.— Goodhart, ' Brit. Med. Journ., Nov. 27, 1875.—
Price, D., 'Lond. Med. Repos./ 1822. — Smith, B., 'Lancet/
1838.— Trotter, ' Chem. and Med. Essays/ 1736.— Wilks (Hen-
derson's case), 'Path. Soc. Trans./ 1860.

Hydatids of the brain and cranial cavity (Bibliogeaphy, No. 20
7c). — (Anonymous) 'Lancet/ April, 1864, p. 444. — Bailey, F.}
'Lancet/ 1825 ; 'Lond. Med. Repos./ 1826. — Barker, T. A.,
'Path. Soc. Trans./ 1858.— Bennett, J. R., 'Med. Times and Gaz.,'
Jan.,1862.— Berncastle, J., ' Lancet/ 1846. — Bree, 0. R., ' Lan-

146

PARASITES OF MAN

cet/1837.— Brittan,F., 'Brit. Med Journ./ 1859.— Burton, 'Med.
Times and Gaz./ 1862. — Dagleish, G., ' Lancet/ 1832.— Fletcher,
T. B. E., 'Assoc. Med. Journ./ vol. iii, p. 161, 1855. — Heading-
ton, 'Edin. Med. and Surg. Journ./ vol. xv, 1819, p. 504. —
Hclsham, 'Med. Comment./ vol. xiii, 1788, p. 289. — Macnamara,
W. H., ' Brit. Med. Journ./ vol. ii, p. 616, 1876.— Big den, G.,
' Prov. Med. and Surg. Journ.,' 1852. — Stewart, J., ' Lancet/
1848.— Sturton, 'Lancet/ 1840.— Wilson, E., 'Lancet/ 1848.

Hydatids of the bones (Bibliography No. 20 I). — Cobbold, T. S.,
" Notice of Specimens of Tibial Hydatids in Nottingham," ' Brit.
Med. Journ./ 1865, and in the 'Veterinarian/ Feb., 1866. —
Idem, " Notice of Specimens from the Tibia in the Mid. Hosp.
Museum/'' ibid. — Cooper, A., " Foster and Lucas's case affecting
the Tibia/' f Surg. Essays/ Lond., 1818. — Ooulson, W. (tibia),
' Med.-Chir. Trans./ 1858 ; see also Daubeny, H., ' Path. Soc.
Trans.,' 1858. — Erichsen, J. E., in his ' Surgery/ 4th edit., pp.
728, 823, and 948, Lond., 1864.— Hunter, W. (tibial, Mus. Spec,
at Glasgow), quoted in ' L'Experience/ 1838, p. 531. — Keate,
B. (os frontis), 'Med.-Chir. Trans./ 1819. — Lambert, J. (tibia),
'Lancet/ 1826. — Thompson, H. (Hearne's tibial case), ' Path.
Soc. Trans./ 1859.— Webster, F. W. (tibia), ' New Eng. Med.
Journ. of Med. and Surg./ 1819. — Wichham, W. J. (tibia), 'Lond.
Med. and Phys. Journ./ 1827.

Hydatids of the breast, muscles, and soft parts (Bibliography
No. 20 m). — Adams, J. (abdominal parietes), 'Lancet/ 1851. —
(Anonymous), " Hyd. in the Eye of a Girl/' ' Boston Med. and
Surg. Journ./ 1849, p. 28. — Baird, J. (muscles), ' Edin. Med. and
Surg. Journ./ 1821. — Birkett, J. (mammary), 'Lancet/ March,
1867, p. 263. — Brodie, B. G. (near scapula), 'Lancet/ 1818. —
Bryant, T. (thigh), ' Path. Soc. Trans./ 1859.— Idem (thigh),
' Lancet/ 1862. — Idem (breast), ' Path. Soc. Trans./ and
'Lancet/ Nov., 1865, p. 565. — Gholmeley (from right side),
'Lancet/ 1826. — Cooper, B. B. (neck and breast, two cases),
'Guy's Hosp. Rep.' 1851. — Idem, in Birkett's work on the
'Breast/ p. 183; the 'Institute/ vol. i. p. 119, 1850. — Dixon,
J. (neck), 'Lancet/ 1851.— Henry, M. (breast), 'Lancet/ Nov.,
1861, p. 497— Hewndon, A. (neck), by Tyson, in ' Phil. Trans./
1706-7, vol. xxv, p. 2344. — Jones, S. (subperitoneal), 'Path.
Soc. Trans./ 1854.— BanMne, J., " Supposed Hyd. in Synovial
Sheaths," 'Edin. M. and S. Journ./ 1830.— Sands (neck),
'Amer. Med. Times/ 1861, vol. ii, p. 376.— White (breast and
arm), 'Lancet,' 1839.

OESTODA

14-7

Hydatids of uncertain seat, or miscellaneous cases and obser-
vations (Bibliography No. 20 n).— Barrett, ' Lond. Med. Gaz./
1838.— Durrani, 0. M. (Ipswich Hosp.), ' Prov. Med. and Surg.
Journ./ IQbl.—Fagge, H., ' Lancet/ July, 1868, p. 76.—
Greenhow, J. M. (intestinal), 'Lancet/ 1823.— Howship, J.
(case, with speculative remarks), 'Edin. M. and S. Journ./
1835.— MacGillivray, P. H. (orbit, &c), 'Austral. Med. Journ./
Aug., 1865.— Idem, ibid., March, 1867.— Idem (3rd series of
cases), ibid., July, 1872. — Idem (treatment with kamala), ibid.,
July, 1872.— Marl-ham, W. 0., " On the ' son hydatique/ "
'Assoc. Med. Journ./ 1856, p. 1072. — Musgrave (letter to Sir H.
Sloane), 'Phil. Trans./ vol. xxiv, 17 '04-5.— Phillips, 'Lancet/
July, 1868, p. 77.— Russell, J J., 'Dub. Journ./ 1838.—
Salter, H, 'Path. Soc. Trans./ 1854.— Ward, T. 0., 'Lond.
Med. Gaz./ 1837.

Hydatids of animals (acephalocysts) (Bibliography No. 20 o). —
Bollinger (see Bibl. No. 49).— Oobbold, 'Manual/ 1. c. (Bibl.
No. 2), 1874. — Crisp, E. (in a turkey and in hogs), 'Path.
Soc. Trans./ 1863. — East, J. (see Steel). — Findeisen, " Echin.
in der Lunge," ' Repertorium fiir Thierheilkund./ 1875,
s. 48. — Gross, S. B. (in swine), ' Elements of Path. Anat./
1845, p. 118. — Hunter, J., "A Cyst (hydatid) which was
filled with water, formed in and filling up the Bone (humerus)
of an Ox (from Hunterian MS.)," more fully described in the
' Catalogue of the Mus. Lond. Coll. Surg./ " Path.," vol. ii,
prep. No. 864, p. 201, 1847.— Idem, " On Hydatids in Sheep "
(supp. to Trans, of a Soc, 1. c, supra), 1793. — Hutchinson, J.,
" Hydatid in the Eye of a Horse," ' Path. Soc. Trans./ and
'Lancet/ 1857. — Huxley, T. H., " On the Anatomy and Deve-
lopment of Echinoc. veterinorum (from a Zebra)," ' Proc. Zool.
Soc./ 1852. — Kirhman, J., " Chronic Disease of the Bones of
the Cranium of a Horse, associated with the existence of Hyda-
tids within a Cyst at the inferior part of the Orbit," the ' Vete-
rinarian/ vol. xxxvi, p. 77, 1863. — Lepper, " Hydatids in the
Kidney of a Lamb," the ' Veterinarian/ 1863, p. 524. — Martin,
J. (in the liver of a sow), ' Vet. Assoc. Trans./ 1842-3, pp.
330 and 364. — Moorcroft, W. (in the brain of a cow), ' Med.
Facts and Observ./ vol. iii, 1792. — Morgan, A. (in the brain of
a mare), the ' Veterinarian/ 1855, p. 396. — Peech, 8. (in the
muscles of a horse), the 'Veterinarian/ 1854, pp. 80 and 209.
— Siedamagrotzlcy (see Bibliog. No. 49). — Simonds, J. B., "Re-
marks on Mr. Scruby's case of Hydatids in the Liver of a

148

PARASITES OF MAN

Sheep," ' Trans, of Vet. Assoc./ 1842-3, p. 331.— Steel, J. E.
(in liver of a cow; Mr East's case), the 'Veterinarian, 1878,
p. 441. — Stoddart (in liver of a cow), the ' Veterinarian/ 1838,
p. 637. — Thudichum, J. L. W. (in sheep), ' Assoc. Med. Journ./
1856, p. 195. — Vincent, J. P. (in horse, causing lameness), the
' Veterinarian/ 1848, p. 674. — Walker, A. (in the liver of a sow),
' Vet. Becord/ 1846, p. 185. — Woodger (in the brain of a horse),
the ' Veterinarian/ 1863, p. 75.

Foreign Literature. Human Hydatids (Bibliography No. 20
p). — Bocher, 'Zur statistik der Echinoc./ Berlin, 1868. — Davaine,
C, ' Traite > (1. c, Bibl. No. 1), p. 350, 1860 ; 2nd edit. p. 369,
1877. — Idem., " Becherches sur les hydatides, les echinoc, &c.,"
' Graz. Med. de Paris/ 1855. — Idem, " Becherch. sur le fremisse-
ment hydatique," fGaz. Med.,' 1862. — JEschricht, ' Danske vi-
denskab. selsk. forhandl./ 1853. — Finsen, ' Ugeskrif t f or Lager/
Bd. iii, 1867; see also c Brit, and For. Med.-Chir. Bev./ 1868,
p. 324; also ' Schmidt's Jahrb. fur Med./ 1867, s. 181.— Gue-
rault, " Sur lamaladie hydatique, &c," ' Gaz. des H6p./ 1857. —
Hearn, A. W., ' Kystes hydatiques du poumon et de la plevre/
Paris, 1875. — Heller, A., " Die Schmarotzer der Leber," von
' Ziemssen's ' Handbuch d. Spec. Pathol, und Therapie,' Bd. viii,
s. 559. — Krabbe, H., " Maladie causee en Island par les Echino-
coques," in his ' Becherches Helminthologiques,' p. 41, Paris,
1866; c Helm. TJnders^gelser,' Copenhagen, 1865, p. 40. — Idem,
' Die echinoc. der Islander," ' Archiv fur Naturg./ 1865, and in
fDen med. Skole i Beykjavik/ 1868. — Idem (see T. B. Jones,
Bibl. No. 2).—Leuchart, R. (1. c, Bibl. No. 1), Bd. i, s. 335,
1863 ; Bd. ii, s. 859, 1876. — Linder, ' Echinococcen der Leber/
Leipsic, 1869. — Naunyn, ' Archiv fur Anat., Physiol./ &c,
1862-3. — Neisser, A., ' Die echinococcen Krankheit./ Berlin,
1877. — Bassmussen, 'Bidrag til Kundskab om Echinoc./ &c,
1865; see also ' Brit, and For. Med.-Chir. Bev./ 1866, p. 285,
and 1867, p. 424. — Schmalfuss, 'Ueber Leberechinococcus/
Breslau, 1868. — Tommasi, T., e Storia di un caso di Echinoc./
&c, in an appendix (Nota) to his edition of my 'Lectures'
(Vermi, &c), Milan, 1873, p. 153.

N0TE> — As Leuckart, Davaine, and especially Neisser offer
exhaustive analyses of the French and German literature of
human hydatids, I will only give the authors' names attached
to such additional foreign memoirs and cases as have been
published in England. These are quoted in my e Introductory
Treatise on the Entozoa.' Full references will be found in the

NEMATODA

149

"Bibliography" of that work under the following heads:— Andral
(pulmonary veins), Angeli, Auglagnier (bladder), Baillarger
(brain), Boinet (liver), Chaubasse (abdominal), Gruveilhier (liver
and spleen), Dupuy (hydatids in animals), Demarquay (liver),
Dupuytren (muscles and viscera, &c), Fouquier (lungs), Gayet
(liver), Goyrand (liver), Guerard, Guillot, Hedinger (brain),
Heintz (liver), Held (thigh), Heller (lip), Klenclce (blood, &c),
KuJm, Lafforgue (liver), Legroux, Livois, Luschha (liver), Martinet
(brain, liver), Maug (hand), Meissner, Miched (brain), Moissenet
(liver), Montault (brain), Morrisseau, Nicolai (liver), Oerstelen
(kidney), Pohl (abdominal), Quinquirez (bladder), Recamier
(abdominal), Richard (liver), Roget (lungs), Roux (pelvic),
Riittel (brain), Schleissner, Siehel, Sommering (eye), Skoda,
Tomowitz (bladder), Zeder (brain).

Additional references to the echinococcus disease as it occurs
in animals will be found at the close of the section devoted to
the parasites of Kuminants (Bibliography No. 49), and I shall
recur to the subject of mortality from " worms " further on.

SECTION III.— Nematoda (Round Worms).

Trichina spiralis, Owen. — The progressive triumphs of
biological science are well epitomised in the history of the
discovery, and in the record of the gradual manner in which we
have obtained our present complete knowledge of the structure
and development of this small entozoon.

Although the facts connected with the original discovery are
clear and indisputable, much error still pervades foreign litera-
ture on this head. Without a doubt Mr Hilton was the first to
suggest the parasitic nature of the capsules first spoken of as
" gritty particles." With Sir James Paget, however, rests the
true discovery and determination of the nematoid character of
the worm itself. With Professor Owen remains the honor of
having first scientifically verified, described, and named the
entozoon. Some have sought, without good reason, to alter
Owen's nomenclature ; yet not only the generic title, but nearly
all else that he wrote concerning the parasite, must be allowed
to stand.

In relation to the capsules, it is true that prior claims of
discovery have been put forward ; but whilst Peacock's prepara-
tion of the "little bodies" testifies to the fact of his having

150

PARASITES OF MAN

seen the capsules before other English observers, including
Wormald, it was Hilton who first surmised their parasitic
character. As for the claims of Klencke and Tiedemann, they
are practically of no value, even if it be admitted that the
former may have at an early period seen something resembling
this nematode, and that the " stony concretions " encountered
by the latter were degenerated capsules.

On no subject have I desired to write with more accuracy
and precision than on this, and lest the above remarks should
appear to be somewhat partial, I now purposely re-state the
facts as they have presented themselves to me during a full and
prolonged study of the entire literature of the subject. If it be
asked with whom rests the discovery of Trichina, the reply
must be framed with a due regard to precise issue at stake.
The first recognition of the capsules as parasitic products is
fairly claimed by Hilton ; the worm by Paget ; the zoological
allocation and nomenclature by Owen ; the adult worm by
Yirchow ; the developmental phenomena by Leuckart ; the
rearing of the larvae by Herbst ; and to crown all, the clinical
importance of the parasite by Zenker. Due regard being had
to these relative claims, I think the following more extended
statement will be found to be true and just in all its bearings.

In the year 1834 Sir James Paget, then a student, first
actually determined the existence of the nematode entozoon,
which was subsequently more completely described by Professor
Owen. The discoverer was assisted by the celebrated
botanist, Robert Brown, who lent his microscope for the
purposes of examination. In the following year Professor
Owen first scientifically described and named the flesh-worm
(Trichina spiralis) in the published transactions of a learned
society. He first fully interpreted the true zoological position
of the parasite. Sir J. Paget's colleague, Mr. Wormald, had
"more than once" previously noticed the characteristic specks
" in subjects dissected at St Bartholomew's Hospital." He
transmitted the individual specimens which enabled Owen to
draw up his valuable paper. It is clear, however, that Mr
Hilton was the first to suggest the parasitic and animal nature
of the specks observed in human muscle. As the " find " was
made in 1832, he anticipated "Wormald in his observation of
the " gritty " particles in dissecting-room subjects, describing
the bodies as " probably depending upon the formation of very
small Cysticerci." Nevertheless, according to Dr Hodgkin,

N EM A TO DA

]51

" the first observation of these little bodies was made in 1828 "
by Mr H. Peacock. The latter made a dry preparation of the
sterno-hyoideus muscle to display the specks. That preparation
is the oldest in existence, and may be seen in Guy's Museum.
It may further be remarked that Henle, Kiichenmeister,
Davaine, myself, and others, have pointed to a notice by
Tiedemann as probably, or possibly, indicating a prior observa-
tion of the specks. Leuckart rejects the evidence. Dr Pagen-
stecher appears to be in doubt as to the nature of the bodies in
question. As the passage in question possibly gave a rough
and imperfect description of the now familiarly known calcified
Trichina capsules, I give a translation of it (Froriep's ' Notizen,'
1822, Bd. i, s. 64) : — "At a post-mortem examination of a man
who had been a great brandy- drinker, and who died from
thoracic dropsy after several severe attacks of gout, Tiede-
mann found white stony concretions in most of the muscles,
especially at the extremities. They lay in the cellular tissue
between the fibre-bundles, frequently also attached to (or near)
the walls of the arteries, being from two to four lines long,
and roundish. The chemical examination conducted by Gmelin
yielded seventy-three parts phosphate of lime, seven parts car-
bonate of lime and twenty parts animal matter, resembling albu-
men or fibrin." In regard to this notice Dr Pagenstecher (' Die
Trichinen/ s. 4) has remarked that Tiedemann' s " communica-
tion was also referred by Henle to such a parasitic development
when he subsequently found Trichina ; and in this sense it was
afterwards received by Diesing, Kiichenmeister, and Davaine.
But it has been rejected by Leuckart on account of the size (from
two to four lines) and seat of the concretions. True, it has
never yet been observed that the capsuled Trichina (not measur-
ing a tenth part of that diameter) subsequently constituted centres
of gouty deposit exceeding their own bulk, nor is it likely that
they should. Seeing, however, as we often do, that errors
respecting size have crept into works on Trichina, we shall not
need to lay much stress upon these statements; still less so
since the notice is very superficial, and its character is essen-
tially of a physiologico- chemical nature. But this, at least,
seems to us decisive, that when Bischoff, at Heidelberg, wrote
on a case which occurred in Heidelberg, not one single word
was mentioned respecting a former case, if such should have
happened, although Tiedemann and himself were on terms of
close intimacy." So much for Tiedemann. In regard to

152

PARASITES OF MAN

Klencke' s claims, the same observer writes : — u Klencke has
asserted that he had already drawn Trichinae in the year 1829,
and that he had seen them again in 1831. This subsequent
statement has no kind of confirmation. The unreliableness,
mistakes, and self-deceptions in the helminthological writings of
Klencke have been repeatedly exposed some twenty years ago."
Prior to this criticism by Pagenstecher, Professor von Siebold
and several other well-known helminthologists had already com-
mented on Klencke' s assertions in the same destructive manner.
In regard to the experimentation and the valuable instruction
thus acquired, it appears that Herbst was the first
to rear muscle-flesh- worms, or encapsuled Trichinae,
in animals (1850) ; whilst Virchow was probably the
first to rear and recognise sexually-mature intestinal
Trichinae in a dog ('Deutsche Klinik/ 1859, s. 430) j
yet, without doing injustice to others, it must be
added that it remained for Prof. Leuckart to offer a
fall, complete, and correct solution of the principal
questions relating to the source and mode of genesis
of the flesh-worm (1860). Leuckart likewise did
good service by disproving the erroneous views
that had been put forth by Kiichenmeister. Lastly,
all these brilliant results culminated in the clinical
observations of Zenker, who opened out a new
epoch in the history of trichinal discovery. Pro-
fessor Zenker was the first to detect the young in
the act of migration, and he likewise primarily
demonstrated the fact that the larval parasites
were capable of producing a violent disease in the
human body.

Never in the history of biological science have
more valuable issues followed the method of expe-
riment upon animals. Not only has human life
been thus saved, but animal life also. State-
medicine and sanitation have received an immense
impulse. The good that has already resulted is
simply incalculable ; nevertheless, in the eyes of a
set of ignorant fanatics who infest this country,
all experiments " involving cruelty to animals "
oufifht to be prevented at any cost. The further

Fig. 35.— Sexually ° r _ J

mature Trichina proerress of biological science in England has

spiralis ; mule. r ° °

Afier Leuckart. hereby sustained a severe check.

NEMATODA

153

The Trichina spiralis in its sexually-mature state is an ex-
tremely minute nematode helminth, the adult male measuring
only the ^th of an inch, whilst the perfectly developed female
reaches a length of about ±". The body is rounded and filiform,
usually slightly bent upon it-
self, and rather thicker behind
than in front, especially in the
males. The head is narrow,
finely pointed, unarmed, with a
simple, central, minute oral aper-
ture. The posterior extremity
of the male is furnished with
a bilobed caudal appendage, its
cloacal or anal aperture being
situated between these divergent
appendages. The penis consists
of a single spicule, cleft above,
so as to assume a V-shaped
outline. The female is stouter
than the male, bluntly rounded
posteriorly, having the genital
outlet placed far forward, at
about the end of the first fifth
of the long diameter of the
body. The eggs measure 7— {'
from pole to pole. The mode
of reproduction is viviparous.

As commonly observed in the
human body our young Tri chinas
appear as spirally-coiled worms
in the interior of small, globular,
oval, or lemon-shaped cysts,
which latter appear as minute
specks scarcely visible to the
naked eye. These specks resem-
ble little particles of lime, being
more or less calcareous accord-
ing to the degree of degenera-
tion which their walls have undergone. In shape and general
aspect they are not altogether unlike the eggs of certain nema-
toid worms, but their size alone sufficiently distinguishes them.
They measure on an average ^" in length by in breadth.

Fig. 36. — Larval Trichina coiled within its
capsule. Alter Bristowe and Ruiney.

154

PARASITES OF MAN

The organised capsules are not essential to the further develop-
ment of the parasite, and are rather to be regarded as abnormal
formations, or rather, perhaps, as products resulting from an
effort of nature to protect and thus prolong the life of the
occupant. They are frequently altogether wantiug. The cap-
suled Trichinae measure §3" in length by about in breadth.
When fully formed they not only exhibit a well-marked digestive
apparatus, but also reproductive organs, which are often, indeed,
sufficiently developed to determine the sex.

Notwithstanding the large number of experiments that have
been more or less recently made by investigators, little or
nothing has been discovered calculated to disturb the conclu-
sions set forth by Leuckart, who writes as follows : — " (1) Tri-
china spiralis is the juvenile state of a little round worm, pre-
viously unknown, to which the generic title of Trichina must
remain attached. (2) The sexually mature Trichina inhabits
the intestinal canal of numerous warm-blooded animals, espe-
cially mammalia (also of man), and constantly in great num-
bers. The duration of its life extends from four to five
weeks. (3) At the second day after their introduction the
intestinal Trichinae attain their full sexual maturity. (4) The
eggs of the female Trichinae are developed within the uterus of
the mother, into minute filaria-like embryos, which, from the
sixth day, are born without their egg-shells. The number of
young in each mother-worm is at least from ten to fifteen
thousand. (5) The new-born young soon after commence
their wandering. They penetrate the walls of the intestine and
pass directly through the abdominal cavity into the muscles of
their bearers, where, if the conditions are otherwise favorable,
they are developed into the form hitherto known. (6) The
directions in which they proceed are in the course of the inter-
muscular connective tissues. (7) Only the striped muscle
(that of the heart excepted) contains Trichinae. The majority
of the wandering embryos remain in those sheathed muscular
groups which are nearest to the cavity of the body, especially
in those which are smaller and most supplied with connective
tissue. Speaking generally, their number decreases with the
distance from the abdomen, being, however, more numerous in
the anterior half of the body. (8) The embryos penetrate
into the interior of the separate muscular bundles, and here
already, after fourteen days, acquire the size and organisation
of the well-known Trichina spiralis. (9) Soon after the in-

NEMATODA iOO

trusion of the parasite the infested muscular fibre loses its
original structure, the fibrillar collapse into a finely granular
substance, whilst the muscular corpuscles change into oval
nucleated cells. (10) The infected muscular bundle retains
its original sheathing up to the time of the complete develop -
meut of the young Trichinae, but afterwards its sarcolemma
thickens, and begins to shrivel at the extremities. 11. The
spot inhabited by the rolled-up parasites is converted into a
spindle-shaped widening, and within this space, under the
thickened sarcolemma, the formation of the well-known lemon-
shaped or globular cysts commences by a peripheric hardening
and calcification. This degeneration commences several months
after the wandering. Immature muscle - Trichinae are not
capable of producing
infection. (12) The
migration and deve-
lopment of the em-
bryos also take place
after the transporta-
tion of impregnated
Trichinae into the in-
testines of a new
host. (13) The fur-
ther development of
the muscle-Trichinae
into adult animals is
altogether indepen-
dent of the forma-
tion of the calcareous
shell, and occurs as soon as the former have reached their
completion. (14) Males and females are already recog-
nisable in their larval state. (15) The immigration of the
Trichina-brood in masses produces very grave or even fatal
consequences, such as peritonitis (from the embryos perforat-
ing the intestinal walls), pain, and paralysis (resulting from the
destructionof the infected muscular fibres). (16) The infection
of man occurs especially through swine. (17) The muscle-Tri-
chinae are so capable of resistance that they are by no means
in all cases destroyed by the ordinary methods of roasting,
cooking, pickling and smoking. (18) As a rule, swine obtain
Trichinae from rats, to which latter we also as the natural
bearers have to convey them. Microscopic examination of

!56 PARASITES OF MAN

flesh is, therefore, urgently recommended as a public preventive
against all danger from Trichinae."

As a summary the above conclusions are well nigh exhaustive;
but whilst I purposely avoid entering into mere clinical details,
there are points of hygienic interest to which I must allude.
Thus, as regards the number of larval Trichinae in any one
" bearer " at a time, this, of course, must be extremely variable,
but it may amount to many millions. In one of the cats on
which Leuckart experimented, he estimated a single ounce of
its muscle-flesh to harbour no less than 325,000 Trichinae. I
find that a relatively similar degree of infection in an ordinary
human " bearer" would yield thirty millions. In the case of one
of my own experimental animals, a pig, I reckoned that there
were at least sixteen millions of Trichinae. The larvae were about
ten months old and enclosed within perfectly formed capsules ;
nevertheless, the animal had never displayed any symptom of
irritation. In a trichinised human subject, examined by Dr
Thudichum, it was estimated that 40,000,000 parasites were
present. My own estimate, calculated from specimens of
muscle obtained from the same case, gave 100,000,000 as the
approximate number of worms present. In the only outbreak
of Trichinosis occurring in England, details of which will be
given further on, I found that the flesh of the hog that had
caused the local endemic contained upwards of 80,000 Trichinae
to the ounce. The consumption of a pound of such flesh
would be capable of producing a collective progeny of some-
thing like 400,000,000 within the human " bearer."

In the year 1865 I conducted a series of experiments upon
upwards of a score of animals, including seven birds, the latter
all yielding only negative results. So far as muscle-Trichinae
were concerned my experiences accorded with those of Professors
H. A. Pagenstecher and C. J. Fuchs, at the Zoological Institute
in Heidelberg. These experimenters found that the ingested
muscle-Trichinae acquired sexual maturity within the intestinal
canal of their avian "hosts;" but they never found young
Trichinae in the muscles of the birds, nor did they perceive
any evidences of an attempt on the part of the escaped embryos
to effect a wandering or active migration on their own account.
Clearly, if the bird's intestinal canal were a proper territory
for the residence of sexually mature Trichinae, we should have
found abundance of wandering non-encapsuled flesh-worms and
also sexually-immature muscle-Trichinae enclosed in well-formed

NEMATODA.

157

capsules. Not a few persons still entertain the notion that
Trichina) are liable to infest all kinds of warm-blooded, and
even also many kinds of cold-blooded animals, such as reptiles
and fishes. Certain nematodes found in earth-worms have been
described as Trichinae ; and consquently, pigs and hedgehogs
were said to become trichinous through eating these annelids.
The minute flesh- worms described by Bowman from the muscle
of the eel are not true Trichinae, any more than the somewhat
similar parasites which Eberth found to infest the muscles of
the frog. The same may also be said of Dr Salisbury's
urinary Trichinae, which are the larva) of Filaria Bancrofti.

Deducting the seven birds, and also six other animals where
no examination after death was possible, I ascertained the
result of my worm-feedings in sixteen instances. Nine of the
experiments were entirely successful, the infected animals
comprising four dogs, two cats, one pig, one guinea-pig, and a
hedgehog.

Carnivorous mammals, especially those subsisting on a
mixed diet, are the most liable to entertain Trichina), but it is
quite possible to rear them in herbivora. Thus, Pagenstecher
and Fuchs succeeded in rearing muscle-Trichina) in a calf, and
they found three female intestinal Trichinae in a goat, but
apparently no muscle-flesh- worms, althougth twenty-seven days
had elapsed since the first feeding with trichinised rabbit's flesh.
In three sheep on which I experimented no trace of Trichina)
could be found. There is no practical need for any further experi-
ments on herbivora, for it is quite clear that, in their natural
state, herbivorous mammals can seldom have an opportunity of
infesting themselves,, whilst the reverse is the case with
swine, carnivorous mammals, and ourselves. Because many
quadrupeds become trichinous, it does not follow that all
mammals are liable to be infested. In the case of most
parasites we find the species limited to a larger or smaller
number of hosts. On the other hand, in not a few cases, the
range of the entozoon is limited to a single territory or host.

In conducting the experiments above mentioned I was
assisted by Professors Simonds and Pritchard, of the Royal
Veterinary College. As they were the only researches con-
ducted on any considerable scale in England, I subjoin a few
details of them. Dr Thudichum's experiments were, I believe,
confined to rabbits.

Exps. 1 and 2. — On the 15th of March, 1865, an ounce of

158

PARASITES OF MAN

flesh containing Trichinae was administered by myself to a black
bitch. The dog being destroyed five days subsequently,
neither intestinal nor muscle-Trichinae were discovered. It was
thought that the dog had thrown up the bolus, which was
strongly saturated with chloride-of-zinc solution. The bolus
consisted of a portion of the pectoralis major of a subject
brought to the dissecting-room at the Middlesex Hospital. The
cysts were highly calcified, but the majority contained living
embryos, which were quite unaffected by the zinc solution
injected into the body to prevent decomposition. At the same
date a small white puppy was experimented on and examined
with precisely the same results. In either case it was too early
to expect muscle-flesh-worms to have become developed.

Exp. 3. — Half an ounce of the same trichinous human flesh
was given (at the same date) to a black-and-tan puppy reared
at the Royal Veterinary College, a second " feeding" being
administered on the 21st of March, or six days after the first.
In this case Mr Pritchard, who fed the animal, took the pre-
caution to chop the muscle into small pieces, and to mix it with
other food, in order that the flesh might be the more readily
retained in the stomach. The puppy was not destroyed until
the 15th of the following June, when, on examination, numerous
encysted but non-calcified muscle-Trichinae were found in all the
voluntary muscles subjected to microscopic scrutiny.

Exp. 4. — An ounce of the same flesh was given to a dark-
colored pig on the 15th of March, and again on the 20th,
several other " feedings" being also administered during the
month of April, 1865. It was destroyed on the 16th of May, but
no Trichinae were detected.

Exp. 5. — An ounce of the same human muscle-flesh admi-
nistered to a small sheep (which was subsequently killed on the
29th of June) also produced negative results.

Exps. 6 and 7. — " Feedings" were at the same time adminis-
tered to a rat and mouse. The mouse died on the 2nd April,
when I examined its muscles without success. On the follow-
ing day the rat unfortunately made its escape, but whether
trichinised or not cannot be said.

Exp. 8. — An ounce of trichinous human flesh was given to a
donkey, in the form of " balls," on the 20th of March ; and
during the month of June four other separate " feedings" with
trichinous dog's flesh were also administered. The animal was
removed from the College without the result being ascertained.

NEMATODA

159

]j;Xp, 9. — From the 15th to the 20th March, 1865, inclusive,
three small Trichina " feedings" were likewise administered to
a guinea-pig. This little animal was not destroyed until the
15th of the following June, when a positive result was obtained.
The pectoralis transversus and other muscles were found to
harbour a considerable number of encysted Trichinae.

Ewp. 10. — On the 20th March, and again on the 21st (1865),
" feedings" from the same human subject were administered to
a hedgehog. On the 26th of April the animal seemed to be
attacked with symptoms of Trichinosis. It refused food, kept
its head extended, and the eyelids closed. On the 27th it
appeared much worse, and on the morning of the 28th it was
found dead. On the 29th I examined the flesh, and found abun-
dance of living Trichinae in the muscles. The capsules were
very thin and transparent. A few days later Mr Simonds also
examined the flesh, and confirmed this result.

Exps. 11 and 12. — Two chickens were fed, on the 21st of
March, with the same material. One of the birds died on the
24th, when I examined the intestines and detected one or two
very minute nematodes, which, at the time, I believed to be
imperfectly developed Trichinae, but subsequently saw reason
to alter my opinion. The other bird died on the 3rd of April,
and certainly contained no muscle-Trichinae.

Exp. 13.— On the 22nd and 23rd of March " feedings,"
amounting to an ounce of flesh in all, were given to a mole.
This animal was returned to the care of Mr Charles Land, who
had previously sent it to the Veterinary College. He subse-
quently reported that, after observing the mole to be "working"
for two or three days, he lost all trace of it, and concluded that
it had either escaped or was dead.

Exp. 14. — On the 1st and 2nd of May portions of the left
fore extremity of the hedgehog (in which we had successfully
reared Trichina from the Middlesex-Hospital subject) were
offered by Mr Simonds to a cat. It ate the flesh very readily,
consuming the entire limb. On the 15th of the following June
the cat was killed, when living Trichinae were found within all
the muscles which we examined.

Exp. 15. — At the same dates a young terrier dog was simi-
larly treated, but did not take the " feeding" so readily. In
this case the left hind extremity of the hedgehog was employed,
and what was not eaten voluntarily was forcibly introduced. On
the 1st of June the dog was attacked with "distemper," and

160

PARASITES OF MAN

died on the 8th of the same month. On examination we found
several living Trichinae in the sterno-maxillaris and other
muscles. Some of the parasites were encysted.

Exp. 16. — From the 9th to the 12th of June inclusive four
separate worm-feedings with the flesh of the trichinised
terrier- dog were administered to a crow. The bird was killed
some months afterwards and sent to me for examination. Its
muscles were entirely free from Trichinae.

Exp. 17. — From the 9th to the 17th of June inclusive seven
separate worm-feedings were administered to a pig. One of
the " feedings " was with the trichinised guinea-pig's flesh, the
others from the dog. This animal was not destroyed until the
4th of April, 1866, when all the muscles which I examined were
found extensively infested with Trichinae. There were probably
not less than 16,000,000 present, all being alive and enclosed
within perfectly-formed capsules, none of which latter exhibited
any traces of calcareous deposition.

Exp. 18. — Four separate feedings with trichinous dog's flesh
were likewise, at the same dates as the foregoing, administered
to a rat. This experimental animal, however, like the one
previously mentioned, contrived to make its escape. I fear it
was well trichinised.

Exp. 19. — About the same date trichinous "feedings " were
given to a black puppy (bred at the Veterinary College). The
dog was killed on the 18th of August, 1866, having also been
made the subject of an echinococcus-feeding, when I found
abundance of encysted Trichinae within the voluntary muscles.

Exp. 20. — Four separate worm-feedings with the flesh of the
trichinised guinea-pig were given to a sheep on the 15th, 16th,
17th, and 19th days of June, 1865. The experimental animal
was destroyed on the 29th of the same month, but the result
was negative.

Exps. 21 and 22. — "Feedings" with the guinea-pig's flesh —
four in the one case and three in the other — were also ad-
ministered by Mr Simonds (from the 15th to the 19th of June,
inclusive) to a chicken and goose respectively. These birds were
destroyed some months afterwards and sent to me for examina-
tion, but the most careful scrutiny failed to detect any
Trichinae within their muscles. The goose was cooked and
eaten without the slightest hesitation. The chicken I found
too tough for consumption.

Exp. 23. — On the 28th of March, 1866, I obtained a small

NEMATODA

161

quantity of muscle from a highly trichinised German subject,
who died from the effects of an accident at the London
Hospital the day previous. The case was fully reported by
Dr Thudichum in a new journal, called ' Scientific Opinion '
(No. 4, April 25th 1866, p. 55). During the same day (at
2.30 p.m.) I fed a dog with part of this human flesh. On the
morning of the 31st I killed the dog, and examined the
intestinal canal (at 11.30 a.m.), which revealed the presence of
sexually -mature living Trichinae. The males (of one of which
I retain an accurate figure) displayed the characteristic bilobed
caudal appendage, leaving no doubt as to their source and
nature. I have mentioned the precise time of the experiment,
in order to show that a period of sixty -nine hours proved amply
sufficient for the development of the young muscle-flesh- worms
of the human subject into the sexually-mature adult Trichinae
of the dog.

Uxp. 24. — With another portion of this human flesh (taken
from the muscles of the tongue) in which the Trichinae were
extraordinarily abundant, I fed a cat. In about ten days the
animal showed the most marked symptoms of trichinosis. It
refused to eat ; the eye lost its lustre ; the body became very
thin, and I thought the animal would die. By very great care,
keeping it warm before the fire, and subsequently inducing it
to take a little milk, the creature improved, gained flesh, and
eventually recovered. About three months afterwards I de-
stroyed this cat, when on examining the jpanniculus camosus,
latissimus dorsi, and other superficial muscles, I found great
quantities of well-developed, capsuled Trichinae. Although the
animal had swallowed scarcely a quarter of an ounce by weight
of the infested flesh, yet thousands of parasites had been pro-
pagated and dispersed throughout its muscular system. In
this way the helminthiasis nearly proved fatal to my cat. As
has been already stated, Dr Thudichum, who I believe had
an opportunity of examining the corpse of this trichinised
German, estimated the number of parasites in his body at
40,000,000. I do not think this estimate likely to be exag-
gerated, for if all the flesh had been infested to the extent I
found to obtain in respect of the muscles of the tongue, I
believe 100,000,000 would have been nearer the mark. In
places the point of a needle could scarcely be thrust between
the capsules, so closely were they agglomerated.

JExp. 25.— From the 19th to the 25th of April, 1866, inclusive,

162

PARASITES OF MAN

daily administrations of trichinous pork, in the form of bolus,
were made to a sheep by Mr Pritchard. The Trichinoe were
obtained from one of our experimental animals at the Veterinary
College, about two ounces of the flesh being given at each
feeding. The flesh of this sheep (destroyed in the following
November) failed to give any indication of the presence of
parasites.

Exps. 26 and 27. — About the same time, and occasionally at
intervals extending over a period of five weeks, Mr Pritchard
also fed two young fowls with the same trichinous pork.
Towards the close of October, 1866, both birds died, when Mr
Pritchard carefully examined the flesh of them, but failed to find
any trace of Trichinae.

Exps. 28 and 29. — From April 2nd to the 9th of the same
month, 1866, inclusive, feedings with trichinous pork were
likewise given to two dogs. These animals were destroyed and
examined by Mr Pritchard in November, 1866, but the result
appears to have been negative.

It is perfectly certain that the infection of man by Tri-
china is invariably due to the ingestion of verminiferously
diseased meat, and as remarked in my ' Lectures/ whenever the
parasites are taken in large numbers unpleasant symptoms soon
show themselves in the infested person. There is, first of all,
restlessness, loss of appetite, and more or less prostration.
This is succeeded by rheumatoid pains in the limbs, with the
frequent accompaniment of considerable swelling. The pain is
not situated in the joints, but in the intermediate soft parts.
In severe cases the limbs are drawn up and half bent, as in
instances of severe and continued cramp. Sometimes the
suffering is excruciating and unbearable, patients having been
known to request the surgeon to put an end to their lives. In
the worst forms of the malady death rapidly ensues from
diarrhoea and exhaustion. If the parasites have gained admis-
sion to the muscles all hope of destroying them is at an end ;
but if a person suspects himself to have eaten diseased or
trichinised meat he should lose no time in seeking professional
assistance, seeing that the administration of suitable anthel-
mintics might be the means of saving his life, whereas a few
days' delay would probably prove fatal. So long as the worms
remain in the stomach or intestinal canal they can be got rid
of, but when once the trichinal brood have invaded the flesh
then they cannot be expelled. As remarked in my ' Entozoa/

NEMATODA

163

it is easy to perceive that although, in the majority of instances,
Trichiniasis does not cause death, yet the percentage of fatal
cases is by no means insignificant.

The notion that particular breeds of swine are more liable to
be infested than others is absurd, since infection must be due
to the facilities offered for swallowing garbage, especially dead
rats. According to Drs Belfield and Atwood 8 per cent, of
slaughtered American swine contain Trichinae. In infested
hogs they found from 35 to 13,000 parasites in a cubic inch of
muscle, and by repeated feedings they succeeded in rearing
about 100,000 Trichinae in the body of a rat.

In regard to the disease in man let us glance at the
phenomena that presented themselves in Plauen, a town of
Central Saxony. Drs Bohler and Konigsdorffer, who first
saw this disease and treated it, state, according to Leuc-
kart, that "the affection began with a sense of prostration,
attended with extreme painfulness of the limbs, and, after
these symptoms had lasted several days, an enormous swelling
of the face very suddenly supervened. The pain occasioned by
this swelling and the fever troubled the patients night and day.
In serious cases the patients could not voluntarily extend their
limbs, nor at any time without pain. They lay mostly with
their arms and legs half bent — heavily, as it were, and almost
motionless, like a log. Afterwards, in the more serious cases,
during the second and third week, an extremely painful and
general swelling of the body took place ; yet, although the fifth
part of all the patients were numbered amongst the serious
cases, only one died."

Satisfactory as it may be to note the numerous recoveries
which take place, this circumstance is very much marred by
the fact that a large proportion of the patients suffer the most
excruciating agony. In the main it will be observed that
Bohler's and Konigsdorffer's experience, as recorded by
Leuckart, corresponds very closely with that given by other
observers. The symptoms, moreover, are very similar to those
produced in the original case published by Zenker. In this case,
which occurred in the Dresden Hospital (1860), the patient was
a servant girl, aged twenty, and the principal symptoms were
loss of appetite, prostration, violent pains, contraction of the
limbs, and finally oedema, which, in association, perhaps, with a
certain amount of pneumonia, terminated her career within a
period of thirty days. The post-mortem appearances showed

164

PARASITES OF MAN

that the larval Trichinae were the cause of death. The intes-
tinal canal contained numerous sexually-mature worms.

The effects produced by Trichinae on animals are similar to
those occasioned in man. The phenomena were summarised
by Davaine (in the journals quoted below) in 1863 as follows :

" The first phase is characterised by intestinal disorder, pro-
duced by the development of the larvae in large numbers, and
their adhesion to the mucous membrane of the intestine. In
thiB stage M. Davaine has seen rabbits die with intense diar-
rhoea ; one of two cats which he fed with trichinised meat had
diarrhoea for at least a fortnight, but survived. Of five or six
rats fed on a similar diet, one only, which was pregnant, died
of diarrhoea, after abortion, on the eighth day. According to
M. Leuckart, the passage of the embryos of Trichinae through
the intestinal walls sometimes produces peritonitis. This intes-
tinal phase often becomes blended with the next ; it may be
relieved by the expulsion of the worms by means of the diar-
rhoea, or may cease with the natural death of the worms.

" The second stage presents general symptoms — muscular
pains, &c. These phenomena are dependent on the introduc-
tion of the Trichinae into the muscles ; they rapidly acquire
their maximum intensity, and have not a long duration. The
appearance and duration of this stage are in complete relation
with the development and length of sojourn of the Trichinae in
the intestines ; in fact, in this entozoon, oviposition is not slow
and of long duration, as in many nematoid worms ; the genital
tube is rapidly formed, and the ova, in its whole length, are
developed almost simultaneously, so that the embryos, arriving
soon at maturity, are at once thrown out in large numbers into
the intestine, and the mother Trichina dies exhausted. If it
be remembered that the embryos do not escape before the
eighth day, that a certain number of days are required for their
arrival in the muscles, and that new ones are not produced
after six or seven weeks, it will be understood that the first
symptoms of this stage can scarcely appear until the end of a
fortnight after ingestion of the diseased food, that they must
continue four or five weeks, and that after this they may dis-
appear. This course of events is observed in animals j and in
man the symptoms of this stage have shown themselves and
become aggravated from the third to the sixth week after
infection. Most animals die during this stage ; rabbits rarely
survive ; rats, on the contrary, generally resist it.

NEMATODA

165

« If the animals do not die of the general symptoms or local
disturbances proper to these two stages, the inflammatory sym-
ptoms cease, respiration becomes natural, and order is re-esta-
blished. But, in some cases, the number of cysts formed in
the muscles are sufficiently great to impede them in the proper
exercise of their functions, and hence arises general debility, a
kind of consumption which persists or becomes aggravated, and
the animal dies of marasmus. M. Davaine has noticed this in
rabbits, but especially in a rat.

" Eecovery from these phases of trichinal infection may be
apparently perfect. A rabbit which M. Davaine kept during
five months became large and fat, although it had a large
number of Trichinae in its muscles ; a rat which had had these
entozoa in considerable numbers during six months was, to all
appearance, in good health. Hence he concludes that the
Trichinae produce symptoms only when they are in the intes-
tinal canal, and when they are entering the muscles. Having
become lodged in their cysts among the muscular fibres, they
may remain harmless for an indefinite time. In every case
except one, down to 1859, Trichinae have been found in the
bodies of persons who have died of disease (generally chronic)
or by accident ; or in the dissecting-room, in bodies regarding
which the previous history could not be obtained. In most cases
the cysts contained a cretaceous or fatty deposit, showing that
they had probably existed several years.

" The observations which have been made on the human
subject, in regard to the symptoms caused by Trichinae, show
that they belong, as in animals, to the initial period of infec-
tion. They consist in intestinal and in muscular lesions ; the
latter coincide with the entrance of the parasite into the
muscles, and are truly traumatic. In Zenker's case the intestinal
symptoms were swelling and pain ; in a case described by
Friedreich diarrhoea was present. In all cases the most
remarkable symptoms were violent rheumatoid pains in the
muscles, not in the joints, which were considerably aggravated
by attempts to extend the half-bent limbs. The other sym-
ptoms have been variable, but have had a strong resemblance
to those of typhoid fever. In several cases there has been
abundant sweating ; and in one there was a very remarkable
miliary and furuncular eruption. The animal heat was dimin-
ished in Friedreich's case ; and in those observed in Voigtland
by Freytag the temperature never exceeded 1 02° Fahr.

166

PARASITES OP MAX

" The progress, duration, and severity of the disease in man
are in relation to the number of Trichinae taken into the
digestive canal. Of sixteen patients observed at Plauen by
Drs Bohler and Konigsdorffer, eight, who were moderately
affected, recovered in a month ; four, more severely diseased,
were ill two months ; of four others, one died with ascites and
colliquative diarrhoea at the end of two months, and three
recovered slowly at the end of three or four months. Recovery
does not imply the death of the Trichinae, it follows their
enclosure in cysts.

" The diagnosis of trichinal infection has several times been
made in the living human subject by removing a portion of
muscle. M. Davaine thinks it probable that, during the first
six or eight weeks of the disease, the diagnosis may be con-
firmed by searching for adult Trichinae in the alvine evacuations,
produced naturally or by means of a purgative.

" The prophylactic treatment consists simply in the avoidance
of uncooked meat. The medicinal treatment must vary with
the stage of the disease. At first, attempts must be made to
expel the parasites from the intestines by purgatives and
anthelmintics. Which amongst the latter is the most energetic
is not yet determined. Calomel is, perhaps, M. Davaine
thinks, the best. After six or eight weeks all treatment
directed towards the intestines is superfluous. It is scarcely
probable that any substance will act on the larvae disseminated
through the muscles. Friedreich has recommended picronitrate
of potash ; but, in the case in which he used it, live Trichinae
were found in the muscular tissue after the patient was con-
sidered to be cured."

In regard to the possibility of curing trichiniasis by the
administration of drugs which should act as trichinacides upon
the parasites in the condition of flesh-worms, the absurdity of
the proposal only equals that which was made in reference to
the destruction of hydatids by the administration of kamala.
As has been shown in the record of my first experiment the
flesh of a trichinised corpse may be thoroughly saturated with
a strong solution of chloride of zinc, and yet the worms will
remain quite unaffected.

In reference to the dangers arising from the consumption of
diseased meat, Professor Gamgee has very cogently put two
questions : — " Did Moses know more about pigs than we do ? "
" Was it a knowledge of the parasitic diseases of swine and mau

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167

that led Moses to condemn pork as human food?" Mr
Gamgee answered both questions negatively, thus : — " The
wisdom of the Mosaic law can only be justly estimated with a
knowledge of the accidents arising in warm countries from
eating pork throughout long and hot periods of the year ; and
there is no doubt that the direct evil results, as manifested by
human sickness, led to the exclusion of pork from the list of
Israelitish viands. The masses of measly pork which may be
seen hanging from the butchers' stalls in Southern Europe
prove that the long-legged swine which hunt the forests for
acorns, and rove about to pick up all kinds of offal, are often
unfit for human food, and that they were so to no less extent
in the land of Israel is probable." As supplementing Pro-
fessor Gaingee's argument, I may remark that, if Moses had
been furnished with special knowledge beyond that of his con-
temporaries, he would not, in the matter of meat-parasitism,
have confined his restrictions to pork. Had he possessed any
knowledge of measly beef, he would not have spared the ox on
the ground that although " it divideth the hoof, yet it cheweth
the cud/'' As regards home-reared animals, Professor Gamgee
cogently remarked : " It is interesting to observe that parasitic
maladies in the pig specially abound in that section of the
United Kingdom where swine live most amongst human beings.
The Yorkshire and Berkshire pigs, in their native counties
enclosed in the farmyards of their breeders, are free from worms
which are likely to live in the body of man. The Irish pig
is the one most commonly injured by entozoa, and the reason
for this is evident when we know how much the cottager relies
on rearing a porker which is permitted the free range of house
and road, where every description of filth is devoured, charged
with the ova of parasites expelled by man or some of the lower
animals/' He also adds : " The conditions under which we live
in the British isles are certainly much less favorable to the
propagation of worms ; but we disregard, in our ignorance, the
most common precautions to protect ourselves from loathsome
diseases, and not only permit dogs to eat any kind of offal
in and around slaughterhouses, but sanction the existence of
piggeries where all kinds of garbage, charged with worms or
their eggs, are daily devoured by swine. The majority of
germs calculated to engender parasites are to be found in
abundance in the contents of the alimentary canal of human
beings and domestic quadrupeds. If pigs are permitted to eat

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PARASITES OF MAN

these, as in Ireland or in many British piggeries, we must
expect hams, bacon, and pork sausages to be charged with the
embryonic forms of human entozoa. Whereas in Iceland the
dog is the victim of human negligence, and en revanche the
cause of human disease, in the British isles the pig holds this
unenviable position. We have good reason to believe, with
Moses, that the pig is an unclean beast ; but without discarding
him from the scanty list of animals to be eaten, it is evident
that we can purify the race of swine, and thus prevent human
as well as porcine maladies."

On the authority of Kupprecht, as quoted by Davaine, I
append a list of the principal epidemics observed in Germany
during the first six years immediately following the discovery
of trichinosis :

1. Two slight epidemics in 1860 in the Island of Kiigen ;
10 to 20 patients (Dr Landois).

2. An epidemic at Stolberg, 1860 (Dr Fricinus). The
number of trichinised persons was not stated with certainty.

3. Five epidemics during five summers, 1858 to 1862, at
Magdebourg. The number of patients was 300, two only died
(Dr Sendler).

4. An epidemic at Plauen in 1862, 20 patients (Bohler).

5. Gusten, 1861, 40 cases, all cured (Frankel).

6. Epidemic in the Province of Armsted (Mansfeld), 1861,
8 patients.

7. Hettstadt, January and March, 1862, 8 to 10 patients.

8. Blankenburg, 1862, 278 cases, 2 deaths.

9. Calbe (Prussia), 1862, 38 cases (9 men, 25 women, 4
children), 8 deaths (Dr Simon and Dr Herbst).

10. Burg, in Magdebourg, 1863, 50 patients, 10 deaths
(Dr Klusemann).

11. Quedlinburg, 1863, 9 patients, 1 death (Dr Behrens).

12. Plauen, 1863, 21 patients (Konigsdorffer) .

13. Falkenstein, 1863, 4 patients (Drs Bascher and Pinter).

14. Posen, August and September, 1863, 37 patients (Dr
Samter).

15. Hamburg, 1863, 2 patients (Dr Tiingel).

16. Blankenburg, 1863, 32 patients, 2 deaths; new cases in

1864 (Dr Schok).

17. Hettstadt (Prussian- Saxony), October, 1863, 158 patients,

27 deaths (Ruppreeht).

18. Eisleben, December, 1863, and January, 1864, 18 cases,

NEMATODA

169

no deaths. This result was attributed to the employment of
phosphoric acid (Eupprecht).

19. Hettstadt, February and March, 1864, 8 patients, no
deaths. Two cats were also attacked, one of which died.
Nearly 50,000 Trichina were counted in an infected leg of pork
(Eupprecht) .

20. Quedlinburg, 1864, 120 patients, 2 deaths; benzine was
employed (Dr. Wolf).

21. Hettstadt, January, 1865, 15 patients (Eupprecht).

22. Berlin, 1864, 3 cases (Dr Cronfeld). Several butcher
boys (Frischer).

23. Leipzig, 1864, 14 patients, 2 deaths ; 4 Were infested
after having eaten raw beef hashed on a block which had
previously received the flesh of a trichinised hog (Dr B. Wagner) .

24. Potsdam, 1864, 5 slight cases (Dr Mollendorf ) .

25. Celle (Hanover), 1864, 8 patients (Dr Scheller and Dr
Baring) ; Trichina proven in the pork by Grerlach. In 1855, 12
Trichina (?) patients were treated by Schuchart.

26. Hedersleben, 25th October, 1865, a pig was killed and
sold ; on the 28th the malady appeared amongst the workmen ;
350 patients, 100 deaths. Of 100 children infected, none died.
Trichina found in the autopsies (Dr F. Kratz).

Dr Davaine also adds the following outbreaks :

In Massachusetts, 1867, 6 patients from having eaten raw
ham, 1 death (' Medical Times/ 20th April, 1867, p. 431).

Eavecchia (Bellinzona), 1868, 5 patients, 4 deaths (Dr Zang-
ger (in Landbote of Winterthur).

Up to a comparatively recent date no case of trichinosis had
been recognised in England during the life of the victim. As
regards diagnosis, what was happening every day on the Con-
tinent was utterly unknown here. Whilst, however, not a
single instance of Trichina- disease had been observed by British
physicians in actual practice, as many as thirty or forty instances
had occurred where the parasites in question had been found
post mortem. I had myself examined the trichinised flesh
taken from a dozen of these corpses, but in no instance had
the faintest suspicion of trichinosis been entertained during
life. The circumstances attending the only outbreak of tri-
chiniasis that has been witnessed in England are very inter-
esting. In the month of April, 1871, I received from Dr
W. L. Dickinson, of Workington, Cumberland, specimens of
pork which he requested me to examine ; and in complying with

170

PARASITES OF MAN

his request I confirmed his opinion that the diseased meat was
infested with Trichinae. A few days afterwards I announced
the discovery in the pages of the ' British Medical Journal'
for April 22, p. 435. It happened, also, that at the time
I was delivering a course of lectures before the Society
of Arts ; consequently, in my second discourse (which was
devoted to the parasites of cattle) I gave full details of the facts
that were obtained. Taking a small portion of the flesh which
I judged to be affected to an average extent I addressed the
audience as follows :

" If you calculate from one portion only, you might, if
you had stumbled upon a part which was more infested with
parasites than another, be led to over-estimate the degree
of trichinisation. Taking proper precautions I have calcu-
lated that one scruple of this trichinous flesh would give us
4320 parasites, and two scruples would therefore yield 8640.
Without speaking so precisely to numbers, I do not hesi-
tate to aver mv belief that there are at least 5000 of these

parasites inside this small piece of ham. The number is pro-
bably close upon 8640. In one drachm that would give us 12,000,
and in an ounce 103,000, according to the old apothecaries'
weight. If, however, we calculate according to the ordinary
weight used by butchers, we should say that one ounce contains
437^ grains of meat, and therefore the number of parasites in one
ounce would be 85,000. Thus, in one ounce of meat from this
particular pig you have 85,000 Trichinae, calculated at the rate
of 200 in the grain, for I have purposely cut off the odd numbers.
You may say, if a person can survive 18, 20, 30, or 40 millions,
he would not take much harm from eating a piece of flesh
containing only 8640 parasites. Such a portion, however,
would be quite sufficient to make any one of us extremely
uncomfortable were we to eat it, for supposing its contained
parasites to be alive, it might prove dangerous to life. Why ?
The explanation is this : — Half of those 85,000 parasites, at the
very least, will, in forty-eight hours after ingestion, have become
fully-developed females; and from each of these 42,000 there will
proceed at least 1000 as a brood, so that the entire progeny
(and it is they that do the mischief by their independent
migrations through our tissues) will eventually yield about
42,000,000 entozoa. If we should be so voracious as to eat a
pound of such trichinised flesh, then there would be 400,000,000
as the result of a single meal.

NEMATODA

171

u Having detailed these facts and inferences, I now wish to
bring to your notice some other particulars connected with the
Cumberland outbreak.

" Dr Dickinson, of Workington, tells me that he was at first
suspicious that his patients were suffering from fever, but was
not quite able to make out what the disorder was. At length
certain symptoms occurred, which suggested that it might
possibly be the German flesh-worm epidemic making its appear-
ance in this country for the first time, and, therefore, in view of
verifying the facts of the case, he sent me portions of the flesh of
the pig. He describes the symptoms, which in their character
corresponded with those previously recorded as experienced by
persons similarly attacked. Dr Dickinson remarks, towards the
close of his communication, that the victims form a small family
who have carefully reared their own swine. The British farmer
is thus here introduced to us at his own table playing the part
of ' host' — at her own table, I should say, for, to be more pre-
cise, it is a widow, her daughter, and a man-servant who are
suffering. Dr Dickinson informs me that for two or three weeks
before he was called to see them they had been eating sausages
and boiled pork from one of their own home-fed pigs, which
pig, by the way, turns out to have been an old sow. He
brought away some sections of the leaner portions of the flesh
for microscopic examination. You will observe that there can
be no mistake about the source of the food on this occasion.
Hitherto, Trichina has not been observed in our British-fed
swine in more than one or two, or possibly three instances.
Therefore it would be very interesting to ascertain how it hap-
pened that this poor pig became trichinised. In my communi-
cation addressed to the ' British Medical Journal/ I wrote as
follows : — " Dr Dickinson has at the present time under his care
a family suffering from the so-called flesh-worm disease,
resulting from the consumption of ham prepared from pigs
reared by the family themselves. A portion of ham sent to me
swarmed with recently encapsuled Trichinae. Dr Dickinson being
thus the first person who has diagnosed trichiniasis in the living
subject in England, I hope he may be induced to give us further
particulars/ The editor, in commenting upon this letter,
added a practical point, which I wish especially to bring to your
notice. He says : — ' The subject of parasitic diseases of
domestic animals is one of widespread and increasing interest.

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PARASITES OF MAN

It is immediately related to the irrigation of fields with
sewage/ The editor, of course, made this statement on
independent grounds, and on his own responsibility. If
he had said the subject bears an indirect relation to the
sewage question, he would have said no more than is abso-
lutely true, for, as I shall take occasion to explain, there is
every reason to suppose that certain forms of parasitic disease
may be propagated by means of sewage. In this connection
some of you may be disposed to ask the question : — ' Are there
any sources of comfort to be gathered from the facts V Or
you may say, supposing that in future our British swine are not
as free from Trichinae as they have been hitherto, can we possibly
avoid the contingency of playing the part of host to those crea-
tures ? Certainly, I reply, it is simply a question of properly cook-
ing the food. If these farmers have not cooked their food at all,
or scarcely at all, that will at once account for their being laid up.
I should tell you that the lady and the daughter are recovering,
and that they are convalescent, but the man-servant is very ill.
If, during cooking, the flesh consumed by these persons had
been raised to a persistent temperature of 170° Fahr., then,
doubtless, the ingestion of trichinised pork would have done
no harm. You observe that Dr Dickinson says in his letter
that they partook of it roasted and boiled. Now, few of us are
in the habit of eating underdone pork, although there are other
meats that we devour very readily in an imperfectly cooked
state. It must be remembered, also, that although the exterior
may have been subjected to a temperature of 212 degrees, it by
no means follows that the whole of the joint throughout must
have been submitted to that temperature. Under rapid cooking,
the centre of a large joint may remain much below even 140
degrees. If the man-servant ate only one ounce of the flesh
with living Trichinae in it, he will probably have at this present
moment at least 42,000,000 of these guests in his muscles.
You will ask, ' Will he recover V 1 Yes ; if he ate no more than
that/ If he has eaten 2 oz. thoroughly underdone, depend
upon it he has 80,000,000, and if he has eaten 3 oz. he will have
over 100,000,000 of Trichinae in his muscles. Could he survive
if he had eaten over 3 oz., and thus have 100,000,000 and
upwards of these inhabitants ? I think he could. Wc have
evidence on this point from the case in which I estimated that
there were upwards of 100,000,000 of Trichinae present, and yet
the man survived the attack.

NEMATODA

173

" Incidentally I may remark that in the course of the last
twenty years, although millions of parasites and their eggs have
passed through my hands, I have almost entirely escaped infec-
tion. It is something to know what you are either handling
or looking at, because there are many parasites besides Tri-
china which are dangerous. There are gregariniform entozoa
residing in meat which we eat every day without any bad
consequences. They are as harmless as cheese-mites. There
is no need to be in the slightest degree nervous about flesh-
food, provided it is properly cooked. I believe there will be
no fatal issue in the case of any of the three individuals just
alluded to, but the chief practical point before us arises out of
the fact that we have here, for the first time in England, an
epidemic of trichiniasis. By calling attention to the subject, it
will, to say the least, suggest precautions by which future
epidemics may be avoided."

The above remarks form the substance of a lecture given
on the 24th of April, 1871. A week later I delivered the third
of the Cantor lectures for that year, when I took occasion to
add the following particulars :

"It has been asked whether the so-called muscle-Trichinae,
after they have arrived at their destination within the flesh of
man, are capable of producing any more unfavorable con-
sequences ? The answer is, Certainly not. In the case of man
it would be necessary that his muscles should be eaten in
order for the Trichinee to become sexually-mature worms ;
and in those countries where cannibalism exists, the man-
eater would himself become trichinised, and would certainly
deserve his fate. I was very desirous to follow up the
account of this outbreak by inquiries respecting the particular
animal which had been the cause of the outbreak. I may
therefore mention that my informant, Dr Dickinson, states
that the family, including the man-servant, all fed together, and
that they had for upwards of a fortnight eaten daily, and some-
times twice a day, sausages made from the flesh of the trichi-
nised animal. And he adds : The meat cut from the ham and
flitches, and what is called the spare-rib, was roasted before
the fire or in the frying-pan. Occasionally it was cooked in
the oven. Dr Dickinson ascertained from the mother that she
liked her meat to be underdone, and thus, therefore, there is
very little doubt that the meat was generally undercooked.
The man, a strong labourer, had a good appetite, and would

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PARASITES OP MAN

therefore get a large share. He is improving slowly. Dr
Dickinson adds in a postscript, what is still more to the point,
that the sausages would be most likely undercooked ; they
would be cooked in the frying-pan, and if only brown on the
outside would be eaten. It is probable that the outbreak was
due therefore to eating underdone meat from this pig, cooked
in various ways, and not alone from the ham itself."

If the facts connected with this outbreak be honestly faced,
it must be rendered clear to any unprejudiced observer that
Dr W. Lindow Dickinson was the first person to observe,
recognise, and treat the Trichina disorder in this country. No
other English, Scotch, or Irish physician has encountered any
similar case. If I lay stress upon this fact it is because I
have learned from Dr Dickinson that another person has
asserted priority in this relation. Sir Dominic Corrigan is
stated to have told a gentleman in the House of Commons,
" that he had often met with trichiniasis in his practice in
Dublin," further averring that the disease " was quite common
in many parts of Ireland." If Sir D. Corrigan merely desired
it to be understood that he had repeatedly encountered the
Trichina at post-mortem examinations, then there is nothing
surprising in his statement, but if, on the other hand, the dis-
ease itself has been frequently recognised in the living Irish
human subject, one can only express astonishment that hitherto
no single instance of the kind appears to have been recorded
either in the public or professional journals.

Bibltogeaphy (No. 21). — English Literature. — Allman, G. J.,
" Exhib. of Specimens," ' Micr. Jrn. and Structural Becord/
1842, p. 94. — Althaus, J., ' Essay on Trichinosis/ London, 1864. —
Idem, " On a Suspected Case," ' Med. Times and Gaz./ 1 864,
p. 161 ; see also pp. 362 and 390. — Atwood, see Belfield. — Ballard,
E., "On Diseased Meat," 'Med. Times and Gaz./ 1864.— BeZ-
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Dec. 28, 1878 ; 1 Med Times and Gaz./ Feb. 15, and ' Lancet/
Feb. 22, 1879 —Bellingham, 0. B., 'Dublin Med. Press/ 1852.—
Bowditch, H. J., " Cases of Trichina," 'Boston Med. and Surg.
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the Discovery of Trichina, in relation to the question of Priority,"
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175

1873, p. 16 ; also in the ' Veterinarian/ March, 1874. — Idem,
Remarks in the ' Journ. Soc. of Arts/ 1866, p. 399; also in
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by Dr. Elendenen at Ostend), in the ' Lancet 3 (anonymous
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(trans, by Ogle), 'Med.-Chir. Rev./ 1863, repr. in ' Edin.
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PARASITES OF MAN

the Pig," ' Annals of Nat. Hist./ and fPr. Ac. N. S. Philad./
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" On Benzine in Trichinosis," ' Med. Times and Gaz./ Oct.,
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— Owen, R.} " Description of a Microscopic Entozoon infesting
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Med. Gaz./ 1835; 'Trans. Zool. Soc./ vol. i, 1835.— Idem,
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1866, p. 269. — Rorie, J., Letter, 'Lancet/ Feb., 1864.— 6W/.S--
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the Human Bladder ; in his paper on " Parasitic Forms," in
' Hay's Amer. Journ. Med. Sci./ 1868, p. 376. — Sanders, R.,
' Edin. Month. Journ./ 1853. — Sawer, A., 'Bost. Med. and
Surg. Journ.,' 1865, p. 16. — Sutton, G., Report on Trichinosis;
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Journ./ Jan., 1864, repr. in ' Glasgow Med. Journ./ April, 1864,
p. 116; also letter in 'Edin. Med. Journ./ Feb., 1864. —
Idem, " Rep. on the Parasitic Diseases of Quadrupeds used
as Food/' pub. by the Med. Officer of the Privy Council,
London, 1865. — Idem, "The Diseases of Meat as affecting
the Health of the People," ' Journ. Soc. of Arts/ April 20,
1866. — Idem, "German Sausages and the Trichina Disease,"
'Scientific Opinion' for April 25, 1866. — Idem, 'Lancet' for
Jan. 6, 1866, p. 16. — Turner, W., "On the Trichina spiralis,"
'Edin. Med. Journ./ Sept., 1860; in the 'Year-Book/
p. 109, for 1860; in 'Med.-Chir. Rev./ 1862; and in 'Bost,
M. and S. Journ./ vol. lxiii, p. 294. — Ude, " Rep. on the
Inspection of Pigs," 'Med. Times and Gaz./ Aug., 1868, p. 141.
— Valentin, 'Micr. Journ. and Struct. Rec./ 1842, p. 87. —
Virchow, R., Extr. from his brochure on 'Trichina' (by myself),
in 'Gunther's Record 'for 1864, p. 611.— Idem, "On the Cure
of Trichinosis," 'Brit. Med. Journ.,' April, 1866, p. 368.— I<7r,/,,
from "Comptes Rendus," in 'Qrt. Journ. Micr. Sci./ 1861. —
Idem, from his 'Archiv/ 1860, Bd. xviii, Heft. 4, p. 330; in
' Brit, and For. Med.-Chir. Rev./ vol. xxvi, p. 515, I860.—
Wedl, Report, 'Brit. Med. Journ./ Dec, 1866, p. 618. — Wills,
S., " Letter respecting the Discovery of Trichina," ' Lancet/

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March 10, 1866, p. 269; see also the ' Times/ Feb. 13, 1866.
— Windsor, J., 'Brit. Med. Journ./ March 4, 1866, p. 319.—
Wood, E., case, 'Lond. Med. Gaz./ 1835.

Foreign Literature : — Ardenghi, E., " Sulla Trichina spiralis,"
'Lo Studente Veterinario/ 1876, p. 115.— Behrens, " Em Fall
von Trichiniasis/7 'Deutsche Klinik/ No. 30, 1863 (quoted by
Davaine).— Bette, F., ibid., 1876.— Bis choff, 'Path. Anat. des
meuschl. Korp./ 1845.— Idem, 'Med. Annalen/ 1840.— Bohler,
'Die Trichinenkrankheit in Plauen/ 1863. — Boudin, "Des
epidemies de Trichina spiralis observees en Allemagne dans ces
dernieres annees," ' Journ. de Med. Yet. Milit./ August and
September, 1864 (quoted by Davaine). — Glaus, ' Wurzb. nat.
Zeitschr./ 1860. — Idem, ' Ueber die Trichine ' (a discourse),
1877.— Golberg, ' Deutsche Klinik/ 1864.— Davaine, G., ' Traite'
(1. c, Bibl. No. 1), 1860, p. 672, 2nd edit., p 732— 768.— Idem,
" Faits et Considerations sur la Trichine," ' Memoires de la
Societe de Biologie ' for the year 1862, torn, iv, ser. 3, 1863;
in 'Gazette Medicale de Paris/ 1863; in 'British Medical
Journal 3 for April 25, 1863 ; and in my ' Bntozoa/ p. 349. —
Idem, "La Trichine" (popular exposition), in 'Revue des
Deux Mondes 3 for May, 1865. — Dujardin (1. c, Bibl.
No. 1), p. 24. — Fiedler, ' Vir chow's Archiv/ 1864. — FlecHes,
F., ' Die Trichinen und die Trichinenkrankheit ' (popular
exposition), Prag., 1866 (quoted by Davaine). — Friedrich,
N., ' Virchow's Archiv/ 1862. — Fiirstenberg, " Wochenblatt
d. Ann. der Landwirthsch., in d. Konigl. Preuss. Staaten/'
1865. — Gerlach, G., ' Die Trichinen, 3 1866. — Idem, ' Hanno-
versche Zeitschrift/ 1864. — Hagen, in ' Pharmaceutische
Centralhalle/ 1862.— iZewZe, 'Muller's Archiv/ 1835, s. 526.—
Herbst, ' Nachrichten v. d. Georg-Aug. Univ. zu Gottingen/
1852 ; 'L'Institut/ 1852, p. 135. — Eeschl, B. L., 'Ueber
Trichinen, die Trichinenkrankheit und die Schiitzmassregeln
dagegen/ Gratz, 1866 (quoted by Davaine). — Eesiner, "Etude
sur le Trichina," ' Gaz. Med. de Paris/ 1864. — Klusemann,
" Die Erkrankung durch den Genuss von Nahrungsmittel aus
dem Thierreiche," 'Deutsche Klinik/ 1864.— Kobelt, 'Valentin's
Repertorium/ 1841. — Krabbe, " Husdyrenes Indvoldsorme,"
' Tiddsskrift for Vet./ 1872.— Kratz, ' Die Trichinenepidemie
zu Hedersleben/ 1866. — Kiichenmeister, ' Parasiten/ 1855. —
LeticJcart, ' Untersuchungen ueber Trichina spiralis,3 1866.—
Idem, " Die mensch. Par./ Bd. ii, s. 409. — Idem, " Die
neuesten Entdeckungen ueber menschliche Eingeweidewurmer

178

PARASITES OF MAN

und deren Bedeutung fiir die Gesundheitspnege," ' Unsere
Zeit./ 1862. — Lion, 1 Zur Geschichte, Therapie, Prophylaxis,
und Sanitatspolizei der Trichinen ' (quoted by Pagenstecher) . —
Luschha, " Zur Naturgeschichte der Trichina spiralis," ' Zeit-
schr. fiir wisseuschaftl. Zool./ 1851. — Meissner, * Zeitschr. f. rat.
Med./ 1855. — Idem, " TJeber Trichinenkrankheit/' ' Schmidt's
Jahrbiicher/ 1863. — Ordonez, E. L., 'Note sur la Distinction
des Sexes et le Developpement de la Trichina- spiralis des
Muscles/ Paris, 1863 ; and ' Compt. Rend. Soc. Biologie/ p. 61,
1863 (quoted by Davaine). — Pag en steelier, 'Verhandl. cL Natur-
hist.-Med. Vereins zu Heidelberg/ 1864. — Idem (und Fuchs),
'Die Trichinen/ 1865. — Perroncito, "La Trichina spiralis" in
' Italia. Estr. degli Annali R. Accad. d'Agric. di Torino/ vol. xx,
1877. — Beyher, 0., ' Die Trichinenkrankheit/ Leipzig, 1862. —
Bodet, H., ' De la Trichine et de la Trichinose/ Paris,. 1865
(quoted by Davaine). — Bupprecht, B., 'Die Trichinenkrankheit
im Spiegel der Hettstedter Endemie betrachtet/ Hettstedt,
1864.— Seidel, 'Jenaische Zeitschr. f. Med. u. Nat/ 1864.—
Siebert, ' TJeber die Trichinenkrankheit und ihre Yermeidung/
Jena, 1863 — Siebold, art. " Parasiten," ' Wagner's Hand-
worterbuch/ 1844. — Simon, G., "Eine Trichinen-epidemie in
Calbe," 'Preussische Medicinal Zeitung/ 1862. — Tommasi,
' La Trichina spiralis e la Malattia prodotta da esso/ Torino,
1863. — Tungel, ' Archiv von Yirchow/ xxvii, 3, 421, 1863
(quoted by Davaine). — Virchow, 'Deutsche Klinik/ 1859;
' Coinptes Rendus de l'Acad. des Sci./ torn. xlix. — Idem,
'Archiv f. Path. Anat. und Physiol./ Bd. xviii. — Idem, 'Dars-
tellung der Lehre von den Trichinen 3 (fur Laien und
Aerzte), 1864. — Vogel, ' Die Trichinenkrankheit/ 1864. — Wag-
ner, " Eine Trichinenepidemie in Leipzig," ' Arch, der Heil-
kunde/ 1864. — Wunderlich, 0. A., " Sur la diagnose probable
de l'affection trichinale," 'Gaz. Med. de Paris/ p. 311, 1863 ;
from ' Wagner's Archiv der Heilkunde/ ii, 3, p. 269,
Leipzig, 1861 (quoted by Davaine) —Zenker, "Zur Lehre
von der Trichinenkrankheit/' ' Deutsches Archiv. fiir Klin.
Med./ Bd. viii, s. 387. — Idem, 'Virchow's Archiv/ 1855 and
1860.

Trichocephalus dispar, Rudolphi. — This well-known worm
possesses a long filiform neck, occupying about two thirds of the
entire length of the body. The surface of the skin though
smooth to the naked eye is furnished on one side with a longi-
tudinal band of minute wart-like papilla). The tail of the male

NEMATODA

179

is curved, and emits at the extremity a short, tubular penis-
sheath, armed with minute retroverted spines. The tail of the
female is straight and bluntly pointed. The eggs measure ?ig"
to ^f' in their long diameter. The whipworm infests the
csecum, and also the upper part of the colon. Upwards of one
thousand were found by Eudolphi in a woman.

The original name of Tvichuris, given to this worm by
Buttner, could not, of course, be allowed to stand when it
became evident that the so-called tail was in reality the head
and neck. The Trichocephalus is not uncommon in England
and Ireland. It is less frequent in Scotland. On the
continent, however, it is so abundant that M. Davaine calcu-
lates that not less than one half of the inhabitants of Paris are
infested by it. From what Dujardin has said it can be
scarcely less abundant in Northern France, for M. Duval, the
distinguished director of the Eennes School of Medicine,
supplied that helminthologist with numerous specimens on
various occasions. The worm abounds in Italy and Egypt ;
being scarcely less prevalent in the United States. The
lamented Mr Noel, one of my old pupils at the Middlesex
Hospital College, brought me specimens which he found post-
mortem on three or four occasions. Dr Haldane, of Edinburgh,
once or twice obtained large numbers (post-mortem). In
Ireland, Bellingham found the worm in eighty-one out of ninety
post-mortem examinations. Mr Cooper, of Greenwich, met
with it, post-mortem, in. eleven out of sixteen instances. When
treating patients for tapeworm I have repeatedly expelled the _u
whipworm.

The organisation of Trichocephalus dispar has been investi-
gated by Dujardin, Mayer, Von Siebold, Eberth, Bastian, and
others. Prof. Erasmus Wilson and myself have carefully
studied the anatomy of the closely-allied whipworm of ruminants
(T. ajjinis) which is discussed in my ' Entozoa.'

The statement of Kuchenmeister that there are no external
appendages in the female Trichocephalus comparable to those
known to exist in the allied Trichosomata, is incorrect. Leuc-
kart's, and especially Virchow's, researches disproved Kiichen-
meister's and Meissner^s notion that Trichina were the youno-
of Trichocephalus. The experiments of Davaine render it
probable that the young get into the human body in a direct
manner. He finds that the eggs undergo no development
whilst yet lodged within the host's intestines. The esre-s a*«

180

PARASITES OP MAN

expelled per anum in the immature condition in which they. first
escape from the body of the parent worm. It further appears
that, after their expulsion, a period of six months must elapse
before embryonic formation commences. The fully-developed
embryo measures ~" in length, and resembles the parent to a
certain extent.

Whipworms rarely put their bearers to inconvenience; never-
theless, both human and animal hosts occasionally suffer from
their presence. Thus, Felix Pascal quotes a remarkable and
fatal instance of cerebral symptoms from this cause in a girl of
four years of age ; and Mr Gibson has recorded an instance in
which these worms produced paralysis and loss of speech.
According to Professor Axe, sheep suffer severely from the
allied species.

Bibliogeaphy (No. 22). — Bastian, H. C, " On the Anatomy
of the Nematoids," ' Phil. Trans./ 1866, p. 545. — Bellint/ham,
0. B., " On the frequency of Trichocephalus dispar in the
Human Intestines," ( Eep. of Brit. Assoc., in Dubl. Journ./
1838, and in ' Med. Chir. Rev./ 1838; see also Bibliog. No. 33
(and the biography of Bellingham by Dr Mapother, in c Dubl.
Jrn. Med. Sci./ 1877, p. 471).— Bush, G., "Anat.of T. dispar/'
'Ann. Nat. Hist./ vol. vii, 1841. — OMaje, sul Tricocephalo
disparo, &c, 1836. — Cobbold, ' Entozoa/ pp. 69 and 329. —
Idem, ' Worms/ pp. 31 and 67. — Davaine, 1. c, p. 205. — Idem,
' Compt. Rend./ 1858, p. 1217, and 'Journ. de Physiol./ 1859,
p. 296. — Dubini, ' Entozoografia uniana/ p. 83. — Dujardin,\. c,
p. 32. — Eberth, " Die Grenerationsorgane von T. dispar,"
< Sieb. und Koll. Zeitschr./ 1860, s. 384.— Gibson, D., « On a
Case of Paralysis, with loss of speech, from intestinal irrita-
tion (produced by T. dispar), 'Lancet/ Aug. 9th, 1862, p.
139.— Goeze, ' Naturg./ s. 112.— Gwrlt, ' Path. An at./ p. 350.—
Kuchenmeister, 1. c, s. 235 ; Eng. edit., p. 321. — Leidy, fProc.
Acad. Phil./ viii, p. 53. — Leuckart, 1. c, s. 465. — Mayer, Sieb.
und Koll. ' Zeitsch. f. wiss. Zool./ Bd. ix, s. 367 ; Bd. x, s. 233,
and s. 383, 1858-60. — Merat, 'Diet. Sc. Med./ p. 560.— Von
Siebold, 'Wiegm. Arch./ 1845.— Wilson, R, ' The Veterinary
Record and Trans./ vol. ii, p. 47, 1846.

Filaria Bancrofti, Cobbold. — The history of the discovery of
this entozoon is second only in interest to that of Trichina
spiralis. Step by step the facts have been evolved by a slow
process of observation, and from the data thus afforded a toler-
ably connected narrative of the probable life-cycle of this

NEMATODA

181

entozoon may now be offered. To place matters beyond all doubt
much remains to be done ; yet that which has been accomplished
is, or ought to be, of surpassing interest alike to the physician,
the scientific pathologist, the epidemiologist, and the philo-
sophic naturalist. In the case of Trichina, Owen's nomencla-
ture was most properly allowed to stand; but for reasons stated
below I hare not hesitated to employ for this worm, in its
adult state, a name differing from that originally given to the
hEematozoon which turns out to be its representative larval
state. Although the male parasite is at present unknown, the
following characters will in the meantime suffice for a diagnosis
of the species : — Body capillary, smooth, uniform in thickness.
Head with a simple circular mouth, destitute of papillae. Neck
narrow, about one third of the width of the body. Tail of
female simple, bluntly pointed ; reproductive outlet close to the
head ; anus immediately above the tip of the tail. Length of
largest females, 3£ in. ; breadth, ±" ■ embryos, to ^" in
length, by to -L/' in breadth ; eggs, averaging by jfa"
from pole to pole.

The first discovery of this entozoon, in its embryo state, was
made by Wucherer on the 4th of August, 1866. To use Dr
Da Silva Lima's words: — "At the moment when Wucherer
was seeking for the Bilharzia hcematolia, he found instead of it
an unknown worm. Our illustrious collaborator/' adds Dr
Lima, "has made his important discovery known under the
modest title of ' Preliminary Notice on a species of Worm at
present not described and still more modestly Wucherer for-
mulated in the following manner his judicious and prudent
conclusions : — It would be rash on my part to put forth a con-
jecture on the coexistence of these worms of the hseinatochy-
luria, aud on the etiological signification which they might
have. I shall therefore abstain until I have been able to make
more ample investigations, and until I have been permitted
to examine the corpse of a heematuric, which has not yet
been possible." (< Gazeta Medica da Bahia/ Dec, 1868
p. 99.)

In the year 1868 Dr J. H. Salisbury referred certain • ova
which he found in the urine to a new and distinct species of
nematode. Although he had no acquaintance with the adult
parasite, Dr Salisbury at once placed the " species " in the
genus Trichina. Here is what he says :— « TricMna cystica
(Salisbury).— This is a small species which I have found in the

182

PARASITES OP MAN

human bladder. In all my examinations I have met with this
little entozoon in three cases only. In two of these it was
only occasionally met with in the urine. In the other it
occurred in great numbers. Frequently from ten to fifteen ova
were found in a single drop of urine."

It is important to remark, that there was no hematuria in
the last-named case, which Dr Salisbury describes as one of
" cystinic rheumatism," or " severe cystinasmia associated with
rheumatism and paralysis." The patient " had been insane for
several years. Her urine was passed milky, with granular
cystine, and was dense and scanty." It is likewise added :
" No examination was made of the muscles after death to
determine whether this species burrowed in the tissue, like the
(Trichina) spiralis."

So much for the principal facts recorded by Dr Salisbury.
His paper is accompanied by two woodcut figures of the ova
(X 300 diam.), and one representation of the embryo (X 1000
diam.). If these figures give the size correctly, the ova mea-
sure only about —i" in length, by in breadth, whilst the
embryo would be about — ' from head to tail.

On the 17th of May, 1872, I communicated to the Metro-
politan Counties Branch of the British Medical Association a
paper on ' Bilharzia,' and in an Appendix to it I wrote as
follows : — " A most interesting circumstance connected with
this case of 'Bilharzia' from Natal lies in the fact that I
obtained from the patient some other urinary parasites in the
egg-condition (fig. 38). On five separate occasions I obtained
one or more specimens of the eggs or embryos of a minute
nematode. In one instance there were about fifty of these ova
in the urine, their contained embryos being well developed and
in a state of activity. Usually they were all in this advanced
condition; but on the 25th of July, 1870, several were observed
in much earlier stages of development. One of these was of a
triangular form; its shape, granular contents, and clearly
defined limiting membrane, indicating separation from the
rachis within the ovarian tube. Another early form was per-
fectly spherical, with a well marked chorional envelope and
double contour. These forms measured about ^' in diameter.
The fully grown eggs observed at the same time gave a
longitudinal measurement of £ by ^ in breadth. On
adding any stimulus, such as diluted sulphuric acid, the embryos
moved themselves freely within the egg. After allowing the

NEMATODA

183

Fig. 38. — Group of eggs and
embryos in a case of endemic
hsematuria (1870). Original.

urine to stand for forty-eight hours, I found, on the 27th of
July, that the shells of the ripe ova had
dissolved, leaving the embryos dead, but
still coiled within a fine transparent en-
velope. In this state they were easily
separated and examined, when they gave
a measurement of ^" in length, by
in breadth. On two occasions, whilst
engaged in rearing the larvae of Bil-
harzia in water, I noticed single speci-
mens of these embryos lying dead ;
and one of the examples thus ob-
served gave a length of ', by 5~" in
breadth."

Knowing what errors of interpretation have often crept into
helminthological literature I was more than usually cautious in
pronouncing upon the source of these urinary parasites. Ac-
cordingly, I remarked that " future discoveries might enable us
to identify the species of nematode to which these ova are
referable." I also added : — " Notwithstanding discrepancies as
to size, I am inclined to think that Dr Salisbury and myself
have been made acquainted with nematode eggs and embryos
referable to one and the same species of parasite. I do not
care to speculate as to the origin of these ova. Long ago I
gave in my adhesion to the determinations of Schneider in
respect of the so-called Spirojptera hominis, but I am by no
means certain that his position may not be disturbed by fresh
discoveries. It is not a little remarkable that the parents of my
patient should have averred that she passed three small vermiform
entozoa by the urethra, corresponding, to judge from their
verbal statements, very closely with the ordinary appearances of
Filaria jpiscium."

Having written thus much seven years back, it is with natural
pleasure that I find my anticipations already verified. Knowing
that I was dealing with parasites in their earliest larval stages,
it never occurred to me to give a specific name to them, and I
could not possibly approve of Dr Salisbury's nomenclature, for
which there was no good ground.

In the original discovery Dr 0. Wucherer procured the worms
from the chylous urine of a female in the Misericordia Hospital
at Bahia; and on the 9th of the following October, 1866 he
obtained similar worms from another female suffering from

184

PARASITES OF MAN

hematuria. He also afterwards found them in a man whose urine
was slightly chylous, but not haematic. In all cases these
sexually-immature nematodes were alive. In September, 1872,
Dr A. Corre furnished a careful description of similar worms
found by Dr Crevaux in a haBmato-chylurous patient at Guade-
loupe. Dr Crevaux frequently examined the blood of this
patient but found no haematozoa. In like manner in Brazil,
Dr J. Silva Lima sought in vain for worms in the blood of no
less than five patients, all of whom suffered from haematuria,
and whose urine contained numerous nematoid worms.

Towards the close of the year 1872 the biological world was
startled by the announcement of the discovery of minute Filariaa
in human blood. Dr T. R. Lewis had found microscopic worms
in the blood, and also in the urine, of persons suffering from
chyluria. The worms could be obtained from day to day by
simply pricking any portion of the body with a finely pointed
needle. To this hasmatozoon Lewis gave the trinomial term
Filaria sanguinis hominis, which thus fitly distinguished it from
the Filaria papillosa hcematica canis domestici described by
Grube and Delafond. Dr Lewis found the average size of the
parasite to be — ' in length by 53—" in breadth. He observed
that while it exists in the blood the body is enclosed in a
delicate transparent tunic or cyst. The worm was never absent
from urine in chyluria. In a case in which there was a milky
discharge from the eyes the worms were also detected. In one
case Lewis calculated that 140,000 Filarias were present in the
blood — a number certainly not relatively large seeing that MM.
Grube and Delafond estimated the verminiferous blood of their
several dogs to contain numbers varying from 11,000 to 224,000.
Lewis also found Filariae in the kidneys and supra-renal
capsules of a woman who died of chyluria. It did not appear
probable that the worms underwent further development in the
human body. On this point Lewis remarks : — " Not only may
those haematozoa found in man live for a period of more than
three years, but there is no evidence that they have any
tendency to develop beyond a certain stage as long as they
remain in the circulation/'' Dr Lewis judged that the form of
chyluria associated with this condition of the blood was local
and intimately related with a tropical climate. The milky
condition of the urine comes on suddenly, not only at first, but
on succeeding occasions also. It is frequently accompanied by
more or less distinctly marked symptoms of various other

NEMATODA

185

obscure diseases, including temporary swellings in the face or
extremities. From certain appearances of intestinal ulceration
Lewis thought that the parasites might gain access to the
system by the alimentary canal, possibly from the tank-water
or the fish inhabiting it. He considered the state of the urine
to be due to the mechanical inteiTuption offered to the flow of
the nutritive fluids of the body. The accidental aggregation of
the Haematozoa might give rise to obstruction of the currents
within the various channels, or occasion rupture of their
extremely delicate walls, and thus cause the contents of the
lacteals, lymphatics, or capillaries, to escape into the most
conveniently placed excretory channel.

Compressed into a small compass, 1 think the above is a
fair statement of the leading facts and phenomena discovered
by Lewis. The whole subject of haematozoology immediately
received additional impulse, the consequences of which have not
yet terminated. In this country Welch was stimulated to
investigate the structure of Filaria immitis in the dog, whilst
others sought diligently for nematoid haematozoa abroad.

On the 20th of April, 1874, Dr Prospero Sonsino communi-
cated to the Neapolitan Royal Academy his memoir entitled
" Researches concerning Bilharzia hcematohia in relation to the
endemic hgematuria of Egypt, with a notice concerning a nema-
toid found in the human blood." In this brochure he made
known the fact of his having discovered microscopic Filariee in a
young Egyptian Jew, in the following words : — " On the 1st of
February last, having well washed the finger of the boy, I
placed one drop of blood under the microscope, when with
astonishment I discovered a living organism of the form of a
nematode, resembling Anguillula, in the midst of the haematic
corpuscles. The worms glided amongst the globules, which were
tossed about by their lively movements, showing various appear-
ances according as they presented themselves either from the
sides, the edges, or the front of the disk" (''Ricerche/ &c, pp. 11,
12). Dr Sonsino took every precaution to prevent error, subse-
quently verifying his " find " from the same patient. Dr Sonsino
directs attention to two of his own characteristic figures of the
worm, and subsequently states not only that he found examples
of the Filariae in the urine of this same youth, but also " in the
urine of another patient." The parasites from these two sources
being figured side by side, it was clear, from their resemblance,
that they referred to one and the same species of entozoon. Dr

186

PARASITES OF MAN

Sonsino having compared the facts supplied by these cases, was
satisfied that the nematodes in question were specifically identical
with those that I had previously obtained from my little African
patient. However, Dr Sonsino was of opinion that his Filariae
were not precisely the same as those that had been described
by Lewis.

On the 8th of April, 1876, I received from Dr William
Roberts, of Manchester, some capillary tubes, charged with
blood, obtained from a patient suffering from chyluria. The
tubes had been transmitted by Dr Bancroft, of Brisbane,
Queensland, Australia ; and in fulfilment of the donor's request,
Dr Roberts afforded me an opportunity of examining their
contents, he having himself verified Bancroft's statement that
they contained Filariae. It was not until May 22nd that I
found opportunity to confirm the observations of Drs Bancroft
and Roberts. The contents of some of the tubes had by this
time completely dried up ; but in others, to which diluted
glycerine had been added, the blood appeared tolerably fresh.
In what might be reckoned as the sixth part of the contents of
one of the tubes, spread on a glass slide, I detected about
twenty Filarise, three of which I sketched in situ, in order to
compare them with the figures of Lewis, and also with others
that I had procured from my Bilharzia- patient in the year 1870.
There could not, I thought, be any doubt as to the identity of
all these sexually-immature nematoids. One novelty, however,
presented itself in the presence of a solitary and empty egg
envelope, measuring about of an inch in its long diameter,
and thus corresponding precisely with the ova that I obtained
from the urine in my Bilharzia case.

According to Bancroft, chyluria is somewhat common in
Brisbane ; and the case here brought forward was not the only
one of the kind which had already furnished Filariaa in the
blood. The patient was a little girl ten years of age.

Thus stood the facts in the spring of 1876. Having
informed Dr Bancroft that a nematoid egg had been detected
in the Australian blood transmitted to England, he was induced
to make further investigations. These happily resulted in the
discovery of the adult worm ; the circumstances attending the
" find" being recorded by Dr Bancroft in a letter written to
myself and dated from Brisbane, Queensland, April 20th, 1877.
He wrote as follows : — " I have labored very hard to find the
parental form of the parasite, and am glad to tell you that I

NEMATODA

187

have now obtained five specimens of the worm, which are waiting
to be forwarded by a trustworthy messenger.

" I have on record about twenty cases of this parasitic
disease, and believe it will be the solution of chyluria, one form
of hematuria, one form of spontaneous lymphatic abscess, a
peculiar soft varix of the groin, a hydrocele containing chylous
fluid, together with some forms of varicocele and orchitis. These
I have verified. In the colony there are no cases that I can
find of elephantine leg, scrotal elephantiasis, or lymph scrotum ;
but from the description of these diseases in the volume on skin
and other diseases of India by Fox, Farquhar, and Carter, and
from Wm. Koberts' article on the latter in his volume on
urinary diseases, I am of opinion that the parasitic nature of the
same will be established.

" The worm is about the thickness of a human hair, and is
from three to four inches long. By two loops from the centre
of its body it emits the Filarige described by Carter in immense
numbers.

"My first specimen I got on December 21st, 1876, in a
lymphatic abscess of the arm; this was dead. Four others I
obtained alive from a hydrocele of the spermatic cord, having
caught them in the eye of a peculiar trochar I use for tapping.
These I kept alive for a day and separated them from each
other with great difficulty. The worm when immersed in pure
water stretches itself out and lies quite passive. In this condi-
tion it could be easily washed out of hydroceles through a
large-sized trochar from patients known to suffer from Filarias."

In July, 1877, I announced Bancroft's discovery in the
' Lancet/ naming the parasite Filaria Bancrofti, and in the
following September I sent the editor an account of the
results of my study of the adult worms received from Brisbane
in the interval. These examinations supplied me with the
diagnosis already given (p. 181).

On the 29th of September, 1877, Dr Lewis published a paper in
the ' Lancet/ wherein, after alluding to my previous announce-
ment respecting the discovery of Filaria Bancrofti, he describes
under the name of Filaria sanguinis hominis a mature worm,
which was evidently the same parasite. Not unnaturally Dr
Lewis put aside the nomenclature I had employed, on the ground
that the name originally given by himself to the embryonal
form ought to be retained, and that "a new name, if not
necessary on anatomical grounds, would only lead to confusion."

JSS

PARASITES OP MAN

Personally I have no objection to Lewis's specific name, but if
the question of priority is to determine the nomenclature, then I
fear we ought to call the species Filaria Salisburyii. Obviously
the retention of Dr Salisbury's nomenclature {Trichina cystica)
would be unsuitable and misleading.

Fig. 39. — Filaria Bancrofli. a, Female (uat. size); i, head and neck (x 55 dinra.);
c, tail; d, free embryo (x 400 diam.) ; e, egg containing an embryo;/, egg, with
mulberry cleavage of the yolk (x 36U diam.). Original.

When (prior to Lewis's discovery of the hsematozoa) I had
myself encountered larval nematodes of the same character as
those described by Salisbury, I, like Wucherer, was careful not
to employ a special name for an immature form, which might
or might not represent a worm hitherto known to science.
The paper in which I described the adult worm from specimens

NEMATODA

189

supplied by Bancroft appeared in the 'Lancet/ Oct. 6th, 1877,
the facts being stated as follows : —

On the 28th of August, 1877, I received a small collection of
entozoa. The box contained the promised Filariaa, and also
eio-ht bottles filled with various intestinal worms taken from
animals. The FilariaB were enclosed in four small tubes and
preserved in glycerine. Three of the tubes (marked 1, 2, 3)
contained sexually-mature worms, the fourth being labelled
" Sediment from adult Fil. sang. — young and ova." I described
their contents in succession. Thus, on the 6th of September, 1877,
I examined the Filaria in tube No. 3. The specimen was injured
and in four portions, these collectively measuring three inches
in length. Although, to the naked eye, the worm had appeared
to Dr Bancroft to be of the thickness of an ordinary human hair,
yet I found it about — ' at the thickest part. It was a female.
At the same time 1 examined the specimen in tube No. 1. This
was also a female. Towards the centre of the body a hernial
protrusion of the uterine horns and intestine had taken place.
In a lithograph sent by Dr Bancroft this specimen was figured
and described as the " parent worm of the Filaria sanguinis,
emitting young Filaria from two loops." Later on I examined
the contents of tube No. 2. In it I found one tolerably perfect
female Filaria, and also a delicate shred forming part of one of
the uterine horns of another worm. This filament measured
one inch and a half in length, and was coiled round the com-
plete worm. On transferring it to a watch-glass containing
water, hundreds of embryos made their escape. Owing to the
transparency of the tissues I had much difficulty in finding the
reproductive outlet, and the effort to find it was all the greater
because Bancroft's figure had misled me. At length I found
the vagina and its orifice close to the head (about ±" from it),
the anal orifice being placed within the ~" from the extremity
of the tail. The vaginal pouch, Ty long, was crowded with
embryos, and a constriction marked its junction with the uterus
proper, which appeared to divide lower down at a distance
of ,3" from the head. Towards the tail a fold of the tuba
Fallopii was seen to extend to within of the extremity. All
sections of the uterine system were crowded with germs, eggs,
and embryos in their usual relative situations.

My examinations of the ova and embryos were chiefly made
from the " sediment " sent in a special glass tube. The fully
formed embryos were fa" in length by in breadth. They

190

PARASITES OP MAN

each showed a double skin, the outer envelope in the more
advanced specimens leaving clear spaces at either end of the
body, resulting from commencing ecdysis. I saw no trace of
intestinal tube, but a central line of condensation marked an
early differentiation of the somatic granular contents. The less
advanced embryos were mostly enclosed in a chorional envelope,
the smallest free embryos measuring only ^m" in length by
in breadth. These had no double contour. The ova, whose
yolk- contents were still in various stages of cleavage, gave an
average long diameter of ~ to ^ of an inch.

Such are the facts I made out, and they enabled me to
amend the characters of the species.

As regards nomenclature, I associated Dr Bancroft's name
with the sexually-mature worm as being in harmony with the
binomial method and little calculated to mislead; moreover, it
helped to fix both the source and date of the discovery (Bris-
bane, Dec. 21st, 1876). The use of this nomenclature detracts
nothing from the high merits of Lewis, who first named the
immature worm Filaria sanguinis hominis. As it now turns
out, both Dr Salisbury and myself had previously been made
acquainted with the young of Filaria Bancrofti ; but it was
reserved for Lewis to discover the hasmatozoal character of the
embryos of this worm, and actually to take them from the
blood. It was a singular circumstance, that when I was
engaged in treating my little African patient for trematode
hasmatozoa, it never once occurred to me that the numerous
nematoid embryos mixed with the Bilharzia ova were haama-
tozoal. As before remarked, it was alleged that my patient
had passed worms two or three inches long by the urethra. I
therefore concluded that these were the parents of the eggs
and embryos, and that all of them were urinary. The infer-
ence was wrong, but it has instructively shown how near one
may go towards a great discovery without really making it.
As regards the larvas, notwithstanding some slight differences
in regard to size and so forth, I have little hesitation in saying
that all the embryo forms severally described by Salisbury, by
myself, by Lewis, Sonsino, Wucherer, Crevaux and Corre,
Silva Lima, Bancroft, Manson, and others, are referable to one
and the same species.

Into the clinical bearings of this subject it is impossible for
me to enter at any length, but I may remark that these para-
sites appear to be associated with, if not actually the cause of,

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191

several distinct morbid conditions. To one of these Bancroft
has given a separate name (Hehninthoma elastica). This is a
highly elastic form of growth to which I have already alluded
under the title of " lymphatic abscess of the arm." In the
first valuable report on Hgematozoa, by Dr Patrick Manson, of
Amoy, China, this careful observer gives interesting particulars
of no less than fifteen cases in which hgernatozoa were found.
Two of these patients had Elephantiasis scroti, two had lymph-
scrotum, two were lepers (one having scrotal disease), two had
enlarged inguinal glands, one had anasarca; and of the remain-
ing six, spoken of as having no concomitant disease, one had
enlarged glands and abscesses, and another suffered from
marked debility. It would thus appear that what is ordi-
narily termed " good health " is rarely associated with a haema-
tozoal condition of the blood in the human subject. The
cases given by Lewis and Manson, where absolutely no recog-
nisable disease existed, must be regarded as exceptional. Dis-
ease, moreover, may exist without any palpable symptoms being
exhibited by the " bearer," and thus perhaps it was with the
haematozoal dogs of G-ruby and Delafond to which I shall again
have occasion to allude. Even those animals that carried
upwards of two hundred thousand microscopic Filarise in their
blood appeared to suffer no inconvenience whatever.

In the autumn of 1877 Dr Da Silva Lima published an
article in the ' Grazeta Medica da Bahia/ in which he dwelt
upon the labors and merits of Wucherer, and, judging from an
omission in one of my memoirs, he supposed that I had insuffi-
ciently acknowledged Wucherer's claims. A translation of
this article appeared in the ' Archives de Medicine Navale/
with an important appendix by Dr le Eoy de Mericourt. In
this addendum the French savant showed that the omission on
my part was unintentional, and had been corrected by me in a
later memoir. Not only had I been amongst the earliest in
England to enforce Wucherer's claims in respect of the micro-
Filarise, but I had first announced his discoveries in connection
with Anchylostoma duodenale. In my translation of Wucherer's
memoir (' Ueber die Anchylostomum Krankheit ') I spoke of the
melancholy satisfaction I had in knowing that the memoir in
question was " among the last that appeared from the pen of
that gifted and amiable physician." Some notice of Dr Lima's
paper and its appendix by Dr A. le Eoy de Mericourt appeared
in the ' Lancet ' for Jan. 5th, 1878, and I also published a full

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PARASITES OF MAN

translation of it, with explanatory notes, in the ' Veterinarian '
for Feb., 1878. Later on, in the 'Lancet' (March 23rd, 1878),
Dr Da Silva Lima published an interesting letter correcting a
misconception that had incidentally arisen in the mind of a
commentator (on the Helminthological work of 1877), and at
the same time he pointed to the original facts connected with
the discovery of Wucherer's Filaria. As my views are in perfect
accord with those of Dr Da Silva Lima, I can only regret that
errors of interpretation should have crept into the discussion,
Dr Lima honorably recognises the nomenclature {Filaria Ban-
crofti) which I proposed for the adult worm, and only claims
for Wucherer that which is fairly due.

On the 4th of January, 1878, I received from Dr Patrick
Manson a manuscript in which he announced the discovery of
the larvae of Filaria sanguinis hominis in the stomach of
mosquitoes. Already, in April, 1877, Dr Bancroft had in-
formed me of his expectation of finding that these insects
sucked up the larvae of the Filaria whilst engaged in their
attacks on man. Dr Bancroft's supposition was a very natural
one, but it remained for Manson to make the actual discovery
of the existence of human haematozoa, or parasites that had
been such, within the stomach of Culex mosquito. I lost no
time in making the principal facts public (c Lancet/ Jan. 12th,
1878). Dr Manson at the same time forwarded for publication
a record of thirty-five additional cases of haematozoa occurring
in Chinese subjects, together with additional particulars of one
of the cases already published in the ' Customs Gazette/
These were afterwards published "as separate contributions in
the c Medical Times and Gazette/ Dr Manson likewise for-
warded materials for a paper entitled " Further Observations
on Filaria sanguinis hominis." In this communication he
gave an analysis of the cases (sixty-two in all) in which he had
observed the haematozoa, and he added valuable statistical
evidence as to the prevalence of Filariae in the Amoy district,
dwelling especially on the influence of age, sex, and occupation
in determining the presence of the parasite. He also described
the morbid states with which these entozoa were commonly
associated.

On the 7th of March, 1878, I formally communicated to the
Linnean Society a detailed account of Manson's investigations
relating to the metamorphoses undergone by the Filariae within
the body of the mosquito. In this paper Manson pointed out

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193

that the female mosquito, after gorging itself with human
blood, repairs to stagnant water for the purpose of digesting
the blood, and also for the purpose of depositing its eggs.
During this period, which lasts four or five days, the Filariaa
undergo remarkable changes. Subsequently, in a more perfect
state, they escape into the water, and in this advanced stage
they are conveyed to the human body along with the water as
drink. Dr Manson persuaded a Chinese, whose blood was
previously ascertained to abound with Filarise, to sleep in a
" mosquito house." In the morning the gorged insects were
captured and examined under the microscope. A drop of
blood from the mosquito was found to contain 120 Filariae, but
a drop taken from the man's hand yielded only some thirty
specimens. Further stages of development are accomplished
within the human host, ending in the sexual maturity of the
parasite. After fecundation successive swarms of embryos
are discharged by the female worm, a part of whose progeny
eventually gains access "to the blood.

Before I proceed to summarise the whole body of facts I
must in the next place state that Manson and myself con-
tributed a joint communication to the Medical Society of
London ou the 25th of March, 1878. In this memoir I especially
dealt with the question of priority .in connection with the dis-
covery of the adult worm. I then restated that the adult
parasite was discovered by Dr Bancroft on December 21st,

1876. The discovery was verified by Dr Lewis on August 7th,

1877, by Dr Silva Araujo October 16th, 1877, and by Dr F.
dos Santos November 12th, 1877. I gave these dates unhesi-
tatingly, without, however, in any way prejudicing the question
already raised in respect of the identity of the worms found in
each case. My own mind was fully made up on that point,
and affirmatively so. Dr dos Santos' find was made in conjunc-
tion with Dr J. de Moura in a case of lymphatic abscess of the
arm. Clinically viewed, the case published by Dr Araujo must
be regarded as unique. Not only were adult and embryonic
Fil arias found in the same patient, but, what was far more sur-
prising and interesting, the patient displayed in his own person
several of the disorders hitherto found apart j and he was more
than once attacked by one or two of the diseases. He expe-
rienced a first attack of chyluria throe years ago, then attacks
of craw-craw commencing a year ago, the latter being 'attri-
buted to bathing in a particular lagoon. He had a second

19L

PARASITES OF MAN

attack of cliyluria six months back, at which time lymph-
scrotum appeared, and also scrotal elephantiasis. Dr Bourel-
Ronciere pronounced this case to be unique, and attributed
nearly all the disorders to the presence of Wucherer's em-
bryonic Filarife. In a very elaborate analysis of and com-
mentary on Dr da Silva Lima's second memoir, Dr Bourel-
Ronciere warmly claims for Wucherer the supreme honor in
all these discoveries. A number of affections hitherto regarded
as distinct, and all of which appear to be due to the action of
Filarial, are regarded by Dr Bourel-Ronciere as mere phases of
one and the same disorder. This affection he terms Wucherer's
helminthiasis. Dr Manson had indeed arrived independently
at a similar conclusion, and I am confident that Wucherer,
were he alive, would in this particular aspect of the question
be the last to claim priority either to Lewis, to Bancroft, or to
Manson.

In this place I may observe that Dr Pedro S. de Magalhaes,
of Rio de Janeiro, detected free microscopic nematodes in the
potable waters of Rio (agua da Oarioca), which from their
similarity he supposes may have some genetic relation with
Filaria Bancrofti. In this opinion I cannot share.

As regards the metamorphoses of the embryo, Manson states
that for a little while after gaining access to the stomach of the
mosquito the embryo undergoes no change (Fig. 40, a). In a
very few hours changes commence, resulting in wider separation
of the outer skin and an appearance of transverse markings on the
body within (ft). In the next stage oral movements occur;
the striation becomes more marked, and the outer envelope is
cast off (c). Then the striated lines disappear and a dotted
appearance is substituted (d). From this condition the embryo
passes to what Manson calls the chrysalis stage, in which nearly
all movement is suspended and the large spots gradually dis-
appear (e, /, g, h, i, j, h). The tail continues to be flexed and
extended at intervals and the oral motions cease. By the
close of the third day the embryo becomes much shorter and
broader ; but the finely pointed tail retains its original dimen-
sions, projecting abruptly from the sausage-shaped body (m, n).
Large cells next appear in the interior of the body, and by a
little pressure one may detect indications of a mouth (o,p, q, r).
At this period the embryo begins to elongate, and at the same
time to diminish in width ; but the growth takes place chiefly
at the oral end of the body. The mouth becomes four-lipped,

NEMATODA

195

Yiq. 40. Larval Filarim in variou8 stage ot growth from the mosquito ; a to d, repre-
senting the first stage of metamorphosis during the first 36 hours; e to , the changes
occurring during the second stage, to the close of the third day ; p to t, forms seen
during the third stage of metamorphosis from the fourth day onwards. The figs, m, p,
represent the head and tail only, whilst t shows the young Vilaria in an advanced
stage, and drawn to a much smaller scale, than the others which arc here magnified
about 125 diameters. Mucli reduced from Mansou's original figures.

196

PARASITES OP MAN

open, and funnel-shaped, and from it a delicate line can be
distinctly traced passing to an opening near the caudal extre-
mity, the tail itself gradually disappearing (s, t). Speaking of
the most advanced stage Manson says : — " A vessel of some
sort is seen in the centre running nearly the whole length of
the body and opening close to one extremity. This end is
slightly tapered down and is crowned with three or perhaps
four papillae, but whether this is the head or tail, and whether
the vessel opening near it is the alimentary canal or the vagina,
I cannot say." Now it is quite evident, I think, from Manson' s
figures that he has here faithfully represented the head and
tail, the former (u) to the left, the latter (v) to the right. In
his manuscript (from which I am now quoting) there is no
special reference to these two figures ; but it is easy to see that
these terminal sections of the body of the advanced embryo
closely correspond with the head and tail of the adult worm
(Filaria Bancrofti). The curved line passing to the left (u)
evidently indicates the commencement of the partially-formed
vagina.

How completely Manson took the initiative in this part of the
work is evident even from Lewis's own later observations. In
a paper published in March, 1878, Dr Lewis, writing from
Calcutta and speaking of the role of the mosquitoes, says : — " I
had repeatedly examined, in a cursory fashion, these and other
suctorial insects, but had not observed any parasites suggestive
of these einbryo-hsematozoa, hence, when, on receipt of a com-
munication from Dr Manson a couple of months ago, a renewed
search was made, I was surprised to find that four out of eight
mosquitoes, captured at random in one of the servants' houses,
harboured specimens of hsematozoa to all appearances identical
with those found in man in this country. After this, however,
several days elapsed before any mosquitoes could be obtained
which contained these embryo-nematoids, and the specimens
obtained on the next occasion were devoid of the enveloping
sheath, which appears to characterise the kind found in man
out here, and apparently, according to Dr Manson, in China
also." Further on Lewis also remarks, " When the insect is
caught shortly after feeding and the contents of its stomach
examined microscopically, the hasmatozoa, if present, will be
observed to manifest very active movements, which may pos-
sibly continue for several hours on the slide. If the insect be
kept for twenty-four hours before examination it is probable

NEMATODA

197

that tlie movements of the parasites will be more sluggish, and
their form probably altered owing to irregular contractions and
dilatations of their substance — changes which may also occasion-
ally be observed when embryo-haamatozoa are preserved on a
glass slide, and they may sometimes be kept alive thus, if in
suitable media, for two or three days. When the insect is not
examined till the third day, the contained parasites will pro-
bably manifest marked signs of disintegration — and possibly
every indication of life will have disappeared from many of the
specimens. After the third or fourth day I have not seen any
active specimens of these entozoa in the stomach or in any part
of the alimentary canal of the mosquito ; those which remain
have undergone more or less fatty degeneration, and are readily
stained with eosin, which, as far as my experience goes, is not
the case so long as they are alive and active. After the fourth
or fifth day it is very rare that traces of any haarnatozoa-like
objects can be detected at all, so that it must be inferred either
that they have succumbed to the digestive action of the insect's
stomach or been disposed of along with the excreta." An
important addendum by Lewis records a fortunate incident
as follows : — " It was observed that nearly all the mosquitoes
captured in one of the servants' houses contained hsematozoa,
so that the supply of suitable insects in all the stages of their
growth became amply sufficient for all requirements. The
result of the examinations under these favorable conditions has
shown that although, the stomach digests a great number of the
ingested haematozoa, as mentioned above, nevertheless others
actually perforate the walls of the insect's stomach, pass out,
and then undergo developmental stages in its thoracic and
abdominal tissues."

I may here observe that Sonsino has instituted a comparison
between the embryos of this Filaria and those of Anchylostoma,
by which it appears that the former measure 0218 to 0*330 mm.
in length, and those of Anchylostoma 0*430 mm. The heama-
tozoa are about forty times longer than broad, and the larval
anehylostomes only fourteen times longer. The tail of Filaria
is conspicuously longer.

In the 'Lancet' for June 22nd, 1878, an announcement
appeared from the pen of Mr D. H. Gabb, of Hastings, stating
that a patient under his care formed the habitat of Filaria
sanguinis hominis j and in the autumn of the same year a
paper which I read to the Linnean Society in the spring

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PARASITES OP MAN

was published. In that paper the following summary was
offered :

1. Filaria Bancrofti is the sexually-mature state of certain
microscopic worms hitherto obtained either directly or indirectly
from human blood.

2. The minute haamatozoa in question — hitherto described as
Wucherer's Filai'iae, Filaria sanguinis hominis, Trichina cystica,
Filariose dermaihemaca, and so forth — are frequently associated
with the presence of certain more or less well-marked diseases
of warm climates.

3. The diseases referred to include chyluria, intertropical
endemic hematuria, varix, elephantiasis, lymph scrotum, and
lymphoid affections generally, a growth called helminthoma
elastica, a cutaneous disorder called craw-craw, and also leprosy.

4. It is extremely probable that a large proportion, or at
least that certain varieties of these affections are due to morbid
changes exclusively resulting from the presence of Filaria
Bancrofti or its progeny within the human body.

5. It is certain that the microscopic hsematozoa may be
readily transferred to the stomach of blood- sucking insects,
and it has been further demonstrated that the digestive organs
of the mosquito form a suitable territory for the further growth
and metamorphosis of the larval Filariee.

6. The character of the changes undergone by the micro-
scopic Filariaa, and the ultimate form assumed by the larvae
whilst still within the body of the intermediate host (Gulex
mosquito), are amply sufficient to establish the genetic relation-
ship as between the embryonal Filaria sanguinis hominis, the
stomachal Filariaa of the mosquito, and the sexually-mature
Filaria Bancrofti.

In the month of September, 1878, I received a letter from
Dr da Silva Lima announcing the fact that Dr Araujo had
verified the existence of the embryos of Filaria Bancrofti in
mosquitoes, at Bahia. These mosquitoes had, I understood,
attacked a French priest in whose blood Dr Araujo also
detected Filarias. Thus, it fell to the lot of Araujo, through
his untiring zeal, to verify in Brazil all the separate discoveries
of Bancroft, Manson, and Lewis.

In the October issue of the 1 Pathological Society's Transac-
tions ' for 1878 Dr Bancroft records numerous cases of filarious
disease, and he gives a succinct account of the circumstances
connected with his original discovery.

NEMATOPA

199

In a clinical lecture published October 12 th, 1878, Dr
Tilbury Fox seeks to diminish the value of these discoveries,
characterising helminthological investigators as merely "recent
writers." Dr Fox denies that Filarias are a cause of true
elephantiasis, but admits the occurrence of " elephantoid in-
flammation and inflammations due to Filarise." Dr Fox's state-
ment that " Filarias have not been found in uncomplicated
elephantiasis, that is, in disease without chylous exudation,"
seems to me to be directly at variance with Manson' s recorded
experiences. I hold that Manson has confirmed the truth of
Lewis's views, and that he has thoroughly proved that (to use
his own words) " varicose groin glands, lymph scrotum,
elephantiasis, and chyluria are pathologically the same disease."
In the first instance I was myself led to conclude that some of
the forms of elephantiasis might be due to other causes than
obstruction of the lymphatics caused by the presence of Filarias ;
but the explanations of Lewis, of Bancroft, and of Manson
more especially, have almost entirely removed this doubt.
Those who seek to explain away the connection between
genuine elephantiasis and Filarias will do well to study Manson' s
last important memoir. He shows that " elephantiasis and
allied diseases are much more frequently associated with the
parasite than are other morbid conditions." This fact is
brought out very clearly in his table of 670 cases, from which
it appears that 58 per cent, of cases of Filaria are associated
with elephantoid disease.

When this opposition to Manson' s views is likely to cease
(on the part of those who do not happen to have been in any
way instrumental to the discoveries in question) it is not easy
to say. In a brief communication which appeared in the last
number of the 'Medical Times and Gazette' for 1878, Dr
Manson successfully combats the doubts that have been enter-
tained respecting the role of the mosquito. Because Lewis
found that canine hasmatozoa were digested, and thus perished
in the stomach of mosquitoes, it had been argued that human,
heematozoa must necessarily undergo similar processes, and
consequently die. Those who oppose the views of helmintholo-
gists in respect of the intermediary host-function of insects on
such grounds can have very little general, and still less special
knowledge of the phenomena of parasitism. It is the old
story. When any new discovery is made, it must always pass
through the ordeals of denial and doubt before it can b© gene-

200

FA 11 A SITES OF MAN

rally accepted as true ; and, as iu the case of Jenner's immortal
discovery, there will always remain a certain number of peculiar
people who show themselves hostile to every advance in science.
Dr Manson may take comfort from this consideration, and rest
assured that the value of his discovery is quite unaffected by
the opposition referred to.

Since I communicated the results obtained by Manson,
Lewis, myself, and others to the Linnean Society, an even
more exhaustive summary of the facts has been published
by Dr Bourel-Ronciere, in the ' Archives de Medecine Navale.'
The distinguished author does full justice to the writings
of English helminthologists, and dwells, with emphasis, upon
the finds and interpretations of Lewis, Manson, and Bancroft.
Incidentally, also, he comments upon Sir Joseph Fayrer's early
recognition of the etiological identity of hseinato-chyluria and
elephantiasis, on other than helminthic grounds. The frequent
concurrence of the two affections had especially struck Sir J.
Fayrer as pointing to a probable common origin. He had also
surmised that the disorders might be due to parasites.

Dr Bourel-Ronciere, alike with the caution, precision, and
logical reasoning of a cultured savant, concludes his elabo-
rate review in the following terms : — " There are the facts.
Certainly, many points remain obscure, many problems await a
solution, and the last word has not been said on the actual part
which the parasite plays in the pathogenesis of the affections
above enumerated — its mode of action, the importance of its
role, the extent of its pathological domain, the habitat of its
progenitors, their identity, and so forth. All these questions
will only be elucidated by necroscopic researches, which at
present remain absolutely wanting."

" However, notwithstanding the doubts which hover over the
future value of these curious discoveries, it is difficult not to
recognise their importance in the study of certain tropical
diseases — which up to the present time have been attributed to
vague and undetermined causes — haeniato-chyluria and ele-
phantoid affections principally. Apart from the interest which
attaches to the natural history of the nematoids, they raise, in
effect, etiological and prophylactic questions, the extreme im-
portance of which we believe it would be needless to demon-
strate. It is greatly to be desired that the researches should
be taken up in other parts of the globe, where endemicity and
perhaps greater facilities for necroscopic investigation would

NEMATODA

201

render them fruitful — Cochin-China, Tahiti, &c. Fresh ob-
servations are necessary to confirm the first and to fill up
notable gaps. The way has been brilliantly opened by the
English and Brazilian physicians. Let our colleagues in the
French colonies put their shoulders to the wheel ; they have
before them a vast field of study to explore.-"

Since the above remarks were written I have received several
communications from Dr Bancroft, and also others from Drs
da Silva Lima, Araujo, Assis Sousa, Paterson, Hall, of Bahia —
the two last named being English physicians in practice there.
I regret that I can do little more than refer to the writings of
these authors in the Bibliography below ; but I may observe that
Drs Paterson and Hall have ascertained that the proportion of
the population of Bahia affected by Filaria is 8^ per cent. Out
of 309 persons examined, 26 had hsematozoa, which is, roughly,
one in twelve, or more strictly, 8"666 per cent.

Amongst recent memoirs that by Sir J. Fayrer, read to the
Epidemiological Society on the 5th of February, 1879, deserves
especial attention. In regard to its significance, I have only
space to remark that, much as we may regret the little interest
shown by our hospital physicians and surgeons in this subject,
it is particularly gratifying to see experienced Indian officers
like Sir J. Fayrer, Mr Macnamara, and Dr John Murray, com-
ing forward both to aid and render homage to their junior
colleagues in Eastern parts, who are successfully labouring to
advance the cause of helminthology and scientific medicine.

In concluding this subject I may observe, that one of the
greatest hindrances to the due recognition of the remarkable part
played by parasites in the production of human endemics and
animal epizootics arises from the circumstance that no incon-
siderable number of minute worms may infest a host without
obvious injury. This immunity proves nothing. If, for example,
we take the case of Trichina we find that several millions of
entozoa may exist in the human, or, at all events, in the animal
bearer, without producing any symptom of discomfort. In such
cases it is not possible to determine the strict limits of health
and disease ; nevertheless, were we to double the amount of
infection, the imaginary line of demarcation is at once bridged
over and the parasites become acknowledged as directly
responsible for grave symptoms which may even prove fatal to
the bearer. Again, the relative strength and size of the infected
host constitute factors that materially limit the power of the

202

PARASITES OF MAN

parasite for injury. Where the entozoa are of minute size,
and where their injurious action is primarily due to the
mechanical obstructions they set up, it is clear that the
virulence of the helminthiases, or resulting diseased conditions,
will mainly depend upon the number of intruders.

Another consideration of the highest value in relation to
epidemiology generally, and more especially in regard to the
practical question as to the best methods of stamping out para-
sitic plagues, is that which refers to the life-history of the
entozoon itself. It must be obvious that in all cases where the
intermediate host can be captured and destroyed, the life-cycle
of the parasite can be broken and interrupted, and if thus
broken, there is an end to the further propagation of the
species. The knowledge that we have acquired by experi-
mental research in this connection has already enabled us to
set a limit upon the prevalence of certain well-known disorders,
such as Trichinosis, Cestode-tuberculosis, and so forth. In
the case of epizootics, however, which are indirectly due to the
action of intermediary hosts that cannot be readily captured or
destroyed, then our power of arresting the disease is com-
paratively limited. In the present case it is probably not
necessary either that a dead or living mosquito should be
swallowed to insure infection ; but it is necessary that the
parasitic larvae should have dwelt within the mosquito in
order to arrive at the highest stage of larval growth prior to
their re-entrance within the human territory. Undoubtedly,
the larvas are swallowed with potable waters. Perfect filtra-
tion before use would certainly check, if in course of time it
did not totally extinguish several of the many virulent diseases
that now afflict the inhabitants of warm climates.

It is with reluctance that I terminate this article, but in
the closing pages of this work (Book II, Section V) I hope
to add a few more particulars in reference to Lewis's latest
researches.

Bibliography (No. 23). — Aravjo, A. J. P. da Silva, "Me-
moria sobre a Filariose/' &c, Bahia, 1875; see also ' Arch, de
Med. Nav./ 1875 and 1878 .— Bancroft, J"., "Oases of Filarious
Disease/' in 'Pathological Soc. Trans.' for 1878, vol. xxix,
p. 407. — Bourel-Ronciere, "Resume of and Commentary upon
the writings of Silva Lima, Silva Araujo, and others," in
'Arch, de Med. Nav/ for March, 1878.— Idem, " Pathologie
exotique. De Phematozoaire nematoide de Thomme et de son

NEMATODA

203

importance pathogenique, d'apres les travaux Anglais et
Breziliens des dernieres annees ;" ibid., for August and Sept.,
p. 113-134 and p. 192-214, 1878.— Oobbold, T. 8., "Discovery
of the Adult Representative of Microscopic Filariaa," ' Lancet/
July, 1877, p. 70. — Idem, ' On Filaria Bancrofti,' ibid. Oct., 1877,
p# 495. — Idem, "Verification of Hasmatozoal Discoveries in
Australia and Egypt," ' Brit. Med. Journ./ June, 1876. — Idem,
" Obs. on Haematozoa," ' Veterinarian/ October, 1873. — Idem,
" Remarks on the Ova of another Urinary Parasite (in the paper
on ' Bilharzia') from Natal," ' Brit. Med. Journ./ July 27th,
1872, p. 89 ; see also Bibl. No. 12. — Idem, " Bntozoa in Relation
to the Public Health" (various papers), ' Med. Times and Gaz./
Jan. and Feb., 1871.— Idem, 'Worms' (1. c, p. 151), 1872.—
Idem, " Heematozoa ; Fresh Discoveries by Lewis," ' Lancet '
for Feb. 6, 1875. — Idem (brief notice), the ' Veterinarian/
p. 209, March, 1875. — Idem, " On the Discovery of the Inter-
mediary Host of Filaria sanguinis hominis," ' Lancet/ Jan. 12,
1878, p. 69. — Idem, " On the question of Priority of Discovery,"
Rep. of Med. Soc. of Lond., in ' Lancet/ March 30, 1878,
p. 465. — Idem, ' Mosquitoes and Filarige ' (explanatory note), in
'Brit. Med. Journ./ March 16, 1878, p. 366.— Idem, " On the
Life-history of Filaria Bancrofti, as explained by the discoveries
of Wucherer, Lewis, Bancroft, Manson, Sonsino, myself, and
others," " Report of the Proceed, of the Linnean Soc." for
March 7, 1878, in ' Pop. Science Rev./ April, 1878 ; and
afterwards published in extenso in 'Journal Linn. Soc./ Oct. 31,
1878. — Idem, "On Filaria Bancrofti," in Part iv of a series of
papers on the Parasites of Man, in the ' Midland Naturalist/
August, 1878. — Idem, " On Filaria sanguinis hominis," in a
letter to the 'Lancet/ July 13, 1878, p. 64. — Idem, " Filarige
and Leprosy" (case from Bancroft) ; 'Lancet/ Feb. 1, 1879. —
Gorre, A., " Note sur Thelminthe rencontre dans les urines
hemato-chyleuses," 'Rev. des Sci. Nat./ 1872. — Gosse, " Sur
Phelminthe rencontre par Wucherer et Crevaux," &c, ' Rev.
Montpellier/ torn, i, p. 190. — Oouto, A., "These de concourso,"
Bahia, 1872. — Crevaux, J., ' De l'hematurie chyleuse, &c,"
1872 ; also in L'Union Medicale/ 1872 (abs. in 'Brit. Med.
Journ./ July, 1872, p. 100); also in 'Arch, de Med. Nav./
1874; and in 'Journ. de TAnat. et de la Physiol./ 1875 (see
also Silva Lima). — Davaine, 0., ' Traite/ 2nd edit., p. 944;
' HEematozoaires/ supp., 1877. — Fayrer, Sir J., "Filaria sang,
hom.," ' Lancet/ March 16, 1878, p. 376.— Idem, " Elephan-

204

PARASITES OP MAN

tiasis Arabum," ' Med. Times and Gaz./ Dec. 1, 1877, p. 588;
" On the Relation of Filaria sanguinis hominis to the Endemic
Diseases of India/' in the ' Lancet/ Feb. 8 and 15, and re-
printed from the ' Med. Times and Gazette ' (same date), 1879.
— Gabb, D. H., letter in ' Lancet/ June 22, 1878. — Leuckart,
1. c, s. 638, 1876. — Lewis, T. K., "On a Haematozoon in Human
Blood," < San. Comm. 8th Rep./ Calcutta, 1872 ; ' Med. Press/
1873, p. 234; ' Indian Ann. Med. Sci./ 1874; ' Lond. Med.
Rec' (abs. by myself in vol. i, p. 5), 1873. — Idem, " Patho-
logical Significance of Nematode Hasniatozoa," ' Tenth Ann.
Rep./ 1873, Calcutta (reprint), 1874; 'Ind. Ann./ 1875.—
Idem, " Remarks regarding the Hsematozoa found in the
Stomach of Culex mosquito," ' Proc. Asiatic Soc. o£ Bengal/
March, 1878, p. 89. — Idem, " Flagellated Organisms in the
Blood of Rats " (being portion of a paper on " The Microscopic
Organisms found in the Blood of Man and Animals/' in ' 14th
Annual Report of the San. Comm. with the Govt, of India'), in
the ' Quart. Journ. of Micr. Science,' Jan., 1879. — Idem (pub-
lished since the present article was written), " The Nematoid
Haematozoa of Man," ibid., April, 1879. — Lima, J. F. da Silva
(with Grevaux), ' Memoria sobre hematuria chylosa ou gordurosa
des paizes quentes ;' extrahida da ' Gazeta Medica da Bahia/
1876; repr. in 'Arch, de Med. Nav./ Dec, 1878 (see also
Le Roy de Mericourt). — Magalhaes, Pedro 8. de, "Filarias em
estado Embryonario, encontradas n'agua tida como potavel (agua
da Carioca)," ' 0 Progresso Medico/ Dezembro, 1877, p. 57. —
Idem, " Nota sobre os nematoides encontrados no sedimento
deposito pela agua (potavel) da Carioca," ' 0 Prog. Med./ 1 de
Setemb., 1878, p. 577. — Idem, " Caso de filariose de Wucherer/
ibid., 15 de Setemb., 1878, p. 589.— Malcina, M.D., "Filaria in
Chyluria," letter in 'Lancet/ Feb. 22, 1879, p. 286.— Manson, P.,
" Rep. on Hasmatozoa," ' Customs Gazette/ No. 33, Jan.-March,
1877; see also 'Med. Times and Gaz.' for Nov. 10, p. 513, Nov. 17,
p. 538,'and Nov. 24, p. 563; Dec. 1, p. 589, 1877; also Jan.,
1878.— Idem, "Additional Cases;" ibid., March 2, 9, 23, 1878.
— Idem, " On Filaria sanguinis hominis, and on the Mosquito
considered as a Nurse," ' Proc. Linn. Soc ,' March 7, 1878 ;
see also report in 'Nature/ March 28, 1878, p. 439.-1^
" On Filaria sanguinis hominis, clinically considered in refe-
rence to Elephantiasis, Chyluria, and allied Diseases," ' Rep. of
Med. Soc. of Lond./ in 'Lancet/ March 30, 1878.— Idem,
" Further Observations on Filaria sanguinis hominis," " Med.

NHMATOI>.\

205

Kep." for April— Sept., 1877, in ' Castoms Gazette/ Shanghae,

1878. Idem, "The Development of the Filaria sanguinis homi-

nis," 'Med. Times and Gaz.' for Dec. 28, 1878, p. 731. — Meri-
court, A. Le Boy de, in Appendix to an art. entitled " Nouvelle
phase de la question relative a la nature parasitaire de la
chylurie. Decouverte du representant adulte de la ' Filaire de
Wucherer/" par le Dr da Silva Lima, from the ' Gaz. Med. da
Bahia/ Sept., 1877; see also the ' Lancet/ Jan., 1878, p. 22
(editorial notice). — Moura, J. de, 'These de Concourso/ 1877. —
O'Neill, "On Craw-craw/' 'Lancet/ Feb., 1875 — Par eira, A. P.,
" On Bilharzia and Chyluria," ' Gazeta Med. da Bahia/ No. 9,
1877 (noticed in 'Lancet/ Feb. 2, 1878).— Salisbury, J. H.,
" On the Parasitic forms developed in Parent Epithelial Cells
of the Urinary and G-enital Organs/' 'Hay's American Journ./
vol. iv, 1868, p. 376.— Santos, F. dos, in ' Gaz. Med. da Bahia/
March, 1877.— Sonsino, P., ' Eicherche/ &c, 1874; 'Delia
Bilharzia/ &c, 1876; ' Sugla Ematozoi/ &c, 1876 (see Bibl.
No. 12). — Idem, " On the Diagnosis of Embryos of Filaria/' in
his paper ' Sull' Anchylostoma duodenale / ' Estr. dall Impar-
ziale/ 1878. — Sousa, M. de A., 'Memoria sobre a Elephantiasis
do escroto/ Bahia, 1878. — Wucherer, 0., "Noticia Preliminar,"
&c, ' Gaz. Med. da Bahia/ Dec, 1868. — Idem. ' Sobre Hema-
turia no Brazil/ ibid., Sept., 1869 ; see also " Mericourt's trans.
(De l'hematurie intertropicale observee au Brezil)," 'Arch., de
Med. Nav./ p. 141, 1870, and the fuller references quoted in
my memoir ; ' Linn. Soc. Journ., Zool./ vol. xiv, p. 368.

Filaria Loa, Guyot. — Although further examinations of this
worm will probably result in placing it in some other genus
than Filaria, yet it is by no means clear that Diesing was right
in placing it with the genus Dracunculus. I therefore abandon
the nomenclature adopted in my previous treatise. According
to the surgeon, Guyot, who made seven separate voyages to the
coast of Angola, these worms cannot be confounded with the
Dracunculus. They are quite white, and relatively much
thicker than guinea- worms. Under the title of Filaria oculi
Moquin-Tandon has spoken of certain small nematodes as "not
uncommon in the negroes of the. Angola coast ;" and he gives
other localities where it occurs. The worms are identical with
those described by Guyot as dwelling beneath the conjunctivae of
negroes at Congo and in the Gaboon region generally. Tho
parasite is rather more than an inch and a quarter in length,
being pointed at one" end and blunt at the other. It is termed

206

PARASITES OP MAN

Loa by the natives, who state that after a period of several
years the worm voluntarily quits the organ. The disease is
thus naturally cured. This parasite enjoys a tolerably wide
geographical distribution, as it has been observed by Clot Bey
in a negress who had come from the town of Monpox, situated
on the banks of the River Magdalena ; by Sigaud, who saw one in
the eye of a negress in Brazil ; by Blot, at Martinique, who
saw two in a negress originally from Guinea ; by Bajon, who
met with one in a little negro girl who had come from Guade-
loupe ; by Mongin, who found one in a negress who had been
living in the Island of San Domingo ; and by Lestrille, who
removed one from beneath the conjunctiva of a negro who
came from Gaboon.

Bibliography (No. 24). — JDavaine, 1. c, p. 839. — Guy on,
' Gaz. Med. de Paris/ p. 106, 1841, and in ' Micr. Journ. and
Struct. Record/ p. 40, 1842, and in ' Dublin Journ./ vol. xxv,
p. 455, 1839. — Idem, ' Compt. Rendus/ torn, lix, p. 743, 1865. —
Guyot, in 'Mem. par Arrachait/ p. 228, 1805. — Kiichenmeister,
1. c, s. 322. — Lestrille, in Gervais and Van Beneden's 'Zool.
Med./ 1859, also quoted by Davaine, 1. c, 2nd edit., p. 840. —
Leuckart, 1. c, s. 619. — Moquin-Tandon, A., 'Zool. Med./ Hulme's
edit., p. 363, 1861.

Filaria lentis, Diesing. — This is a doubtful species. The
worm was first discovered by Nordmann, in a case of lenticular
cataract under the care of Von Grafe, and it was afterwards
found by Jungken in a similar recorded by Sichel.

There is also the instance described by Gescheidt, in which Von
Ammon operated, and from which brief descriptions of the
worm have generally been taken. In this case there were three
worms, two measuring about g" and the third ~" in length. In
Jungken' s case (exhibited by Quadri, of Naples, at Brussels) the
worm was more than §" long. In another case, reported by M.
Fano, the worm was somewhat less than long. There is no
certain evidence that any of these various worms had developed
sexual organs in their interior. It is true that the reproductive
organs were described in two of the worms observed by
Gescheidt; but after a due consideration of all the facts I
fear we must conclude that all the worms in question were
sexually-immature and wandering nematodes, possibly referable
to Gurlt's Filaria lacrymalis, as Kiichenmeister long ago
suggested.

Bibliography (No. 25). — Oohbold, ' Eutozoa/ p. 332. — Da-

NEMATODA

207

vaine, 1. c, p. 821 et seq.—Viesing , ' Syst. Helm./ p. 625.—
Jbno, ' Traite des Malad. des Yeux/ torn, ii, p. 498 ; and in
' Rec. de Med. Vet./ p. 140, 1869 ; quoted by Davaine, p. 831.
—Gescheidt, Amnion's ' Zeitsch./ 1833, s. 436—LeucJcart, 1. c,
Bd. ii, s. 622.— Nordmann,\. c, Bibl. No. 2, 8. 7, 1832. — Sichel,
'Iconogr. Ophth./ p. 707, 1859.

Filaria labialis, Pane. — This is a filiform cylindrical worm
measuring an inch and a quarter in length. The mouth is
armed with four papillae arranged in the form of a cross. The
tail of the female is blunt, the vaginal outlet being placed at a
very short distance from its extremity, and a little above or in
front of the anus. This parasite was found by a medical
student at Naples. It occupied the cavity of a pustule in the
upper lip, giving rise to considerable irritation. Only the male
worm is at present known.

Bibliogeaphy (No. 26). — Davdine, 1. c, edit, ii, Synopsis,
p. 107. — Leuckart, 1. c. (with a fig.), Bd. ii, s. 616. — Pane,
" Nota di un elminte nematoide," in ' Annali dell' Acad, degli
aspiranti Naturalisti/ Napoli, ser. 3, vol. iv, 1864.

Filaria hominis oris, Leidy. — In the fifth volume of the
f Proceedings of the Philadelphia Academy of Natural Sciences '
(1850, p. 117) Dr Leidy furnishes the following description of
this worm as gathered from the examination of a simple spe-
cimen preserved in alcohol, and labelled as having been
" obtained from the mouth of a child." Body white, opaque,
thread-like ; mouth round, simple ; posterior extremity obtuse,
furnished with a short, curved, epidermal hooklet, in length,
by in diameter at base. Dr Leidy offers some specula-
tions as to its origin, but from whatever source the worm was
obtained by the bearer, it seems to be an immature form. Its
length is five inches and seven lines.

Filaria (Nematoideum) trachealis, Bristowe and Rainey. —
This is another very doubtful worm. It was originally de-
scribed in the 'Pathological Society's Transactions' for 1855.
It evidently represents only a juvenile stage of growth of some
species of round worm. Rainey discovered a considerable
number of these worms in the trachea and larynx of a person
who died from a disease affecting the lower extremities. In-
dividually the parasites measured about the — ' in length.

Struvgylus (Filaria) bronchialis, Rudolphi. — This is a small
nematode. The male measures rather more than half an inch
whilst the female is upwards of an inch in length. The caudal

208

PARASITES OF MAN

appendage of the male is furnished with a bilobed, mem-
branous, half -bell- shaped bursa. This surrounds the cloacal
outlet, the latter concealing a double spiculum. The tail of
the female is sharply pointed, the anal orifice being placed a
little in front or above. The body is filiform, of a pale yellow-
color. It is about ^" broad in the male, and — ' in the female.
The mode of reproduction is viviparous.

The original specimens were discovered by Treutler iu
Germany, during the winter of 1791, in the bronchial glands of
an emaciated subject, whilst those sent to Diesing for descrip-
tion were discovered by Dr Fortsitz at Klausenberg, in Transyl-
vania, in the lungs of a boy six years old. Diesing and
Weinland suggested the identity of Filaria bronchialis and
Strongylus longevaginatus, whilst Kiichenmeister went further,
and pronounced them to be one and the same species.

Bibliography (No. 27). — Cobbold, ' Entoz./ p. 357. — Davaine,
' Synops./ 1. c, ' Synopsis ' cix. — Kiichenmeister, 1. c, Eng. edit.,
p. 381. — Leuckart, 1. c, s. 618. — Treutler, F. A., "De vermibus
filiformibus (Hamularia lymphatica) in glandulis conglobatis
bronchiorum repertis/' in ' Obs. Pathol. Anat./ 1793. — WedL,
' Die im Menschen vorkommenden Helminthen ' (quoted by
Leuckart), Wien, 1862, s. 22.

Eustrongylus gig as, Diesing. — This is by far the largest
nematode known to science, the male sometimes measuring a
foot in length and the female more than three feet, whilst the
breadth of the body reaches half an inch at the thickest part.
Though fortunately very rare in man, this worm is known to
occur in a great variety of animals, especially in weasels.
According to Weinland and Jackson, it is particularly abun-
dant in the kidney of the North American mink (Mustela
vison), destroying the substance of the organ, the walls of
which become the seat of calcareous deposit. It has been
found in the dog, wolf, puma, glutton, raccoon, coati, otter,
seal, ox, and horse.

The body of the adult worm is cylindrical, more or less red
in color, and somewhat thicker behind than in front. The
head is broadly obtuse, the mouth being supplied with six
small, wart-like papillee, two of which correspond with the
commencement of the two lateral lines of the body. These
lines are also distinguishable from other six longitudinal lines
traversing the body from end to end by the presence of very
minute papilla? which are less closely arranged towards the

NEMATODA

209

centre (Leuckart). The tail of the male shows a simple,
thick, cup-shaped bursa, which is destitute of rays, and
partly conceals the simple spiculum. The tail of the female
is blunt and pierced by the centrally placed anal opening.
The vulva is situated near the head in the ventral line. The
eggs are stout and oval, measuring ~5" in length by about 535"
in breadth.

As regards development the recent researches of Schneider
have shown that certain kinds of fish play the part of inter-
mediary bearer. Balbiani preserved the ova in water for more
than a year without their hatching, and all his attempts to rear
the larvae in the intestines of the dog by direct experiment
failed. Similar feeding experiments upon fishes and reptiles
also failed. The embryo, when removed from the egg, measures
TjL" in length. It is vermiform, having a pointed head and
simple mouth. Balbiani describes the buccal cavity as con-
taining a protractile stylet. Notwithstanding the negative
results obtained by Balbiani s experiments on fishes, Schneider
(from anatomical data, which Leuckart confirms) has placed it
almost beyond question that the worm hitherto known as
Filaria cystica is the sexually-immature Eustrongylus gigas.
This worm is found encysted beneath the peritoneal membrane
in Galaxias scriba and Synbranchus laticaudatus. It is worthy
of remark that the genus Galaxias comes nearer to the Sal-
monidaa than to the pike family, whilst the Synbranchi are
tropical oceanic fishes. Probably the sexually-immature worm
occurs in other fishes, especially the Salinonidse.

Kemarkably fine examples of the adult worm may be seen
in the Hunterian Collection, Lincoln's Inn, and in the Museum
of the Eoyal Veterinary College. The human example is
undoubtedly genuine. The dissections in the Hunterian Col-
lection of specimens were made by me in 1865. Objection has
been taken to my description of the oesophagus as "spiral."
In Sheldon's specimen it is certainly twisted upon itself, precisely
in the manner in which Davaine has also figured it (' Traite/ fig.
68) ; but I cannot here give further anatomical particulars.
Drelincourt found two worms sexually united in the kidney.
When once the parasites have gained access to this organ,
rapid destruction of the glandular substance follows. Ulti-
mately the kidney is reduced to the condition of a mere cyst or
bag, which, besides the worms, contains a quantity of sanguinco-
purulent matter. Frequently only one worm is present, but

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PARASITES OF MAN

oftener two or three. In the kidney of a puma D'Azara's
friend, Noseda, found no less than six worms, whilst Klein
obtained eight from the kidney of a wolf.

Bibliography (No. 28). — Azara, F. de, ' The Natural History
of the Quadrupeds of Paraguay/ trans, from the Spanish by
W. P. Hunter; Valpy's edit., p. 43, 1837 ; Black's, 1838;
French edit., p. 313, 1801.— Albers, ' Beitr. z. Anat. &c.,' Bd. i,
s. 115. — Aubinais, 'Revue Med.,' 1846, p. 284. — Balbiani,
" Recherches," &c, ' Compt. Rend.,' 1869, p. 1091; ' Rec. de
Med Vet.,' 1870, p. h.—Bickford, " Spec, of Sir. gigas found in
the Kidney of a Dog," the ' Veterinarian,' 1859, p. 312. —
Blainville, ' Diet, des Sci. Nat.,' tab. 29. — Blanchard, e Ann.
des Sci. Nat.,' 1849, p. 186. — Idem, in ' Cuvier's Regne Animal'
(Masson's edit.), ' Les Intestinaux/ p. 57, pi. 27. — Blasius, ' Obs.,
&c.' (with fig. of Lumbricus in renibus hominis), 1674, p. 125. —
Bobe-Moreau, in ' Journ. de Med.,' torn, xlvii. — Boerhaave,
' Aphorism.,' ] 728.— Bremser (1. c, Bibl. 2), s. 223. —Chabert,
' Traite des maladies verm, dans les Animaux,' 1782. — Chiaje,
' Comp. d. Elmintogr. umana,' p. 106. — Glamorgan, J. de, ' La
Chasse de Loup,' 1583 (quoted by Davaine, the worms being
described as " serpents et betes fort venemeuses "). — Gobbold,
'Entoz./ p. 358. — Idem, ' Catalogue of Entozoa in the Museum
of the Roy. Coll. of Surg.,' " Descr. of preps. Nos. 19—25," p. 3,
1866. — Idem, "Parasites of Man," 'Midland Naturalist,' Dec,
1878. — Collel-Meygret, " Mem. sur un ver trouve dans le rein
d'un Chien," in ' Journ. de Physique,' &c, 1802. — Ouvier, see
Blanchard (supra). — Idem, 'Voyage en Sicile,' and in 'Ann.
des Sci. Nat.,' torn. xi. — Davaine, C, ' Traite,' 1. c, deuxieme
edit., p. 271 et seq. (with full lit. refs. at p. 290). — Diesing, 1. c,
vol. ii, p. 325. — Dujardin, 1. c, p. 113. — Frank, F., "Ein Spul-
wiirm in der Urinblase eines Hundes," ' Hufeland's Journ.,'
Bd. xviii, s. 112. — Jackson, 'Catalogue of the Boston Museum/
1847, p. 317. — Klein, T. K., "Anatomical Description of Worms
found in the Kidneys of Wolves," 'Phil. Trans./ 1729-30,
p. 269—Kuchenmeister, 1. c, Eng. edit., p. 376.— Leblanc (rep.
by Rayer and Bouley), in 'Bull, de l'Acad. de Med.,' 1850,
p. 640 ; in ' Rec. de Med. Vet./ 1862, p. 800 ; and quoted by
Davaine.— Leuckart, 1. c, Bd. ii, s. 353-401, 1876.— Moublel,
" Mem. sur les vers sortis des reins et de l'urethre d'un enfant,"
'Journ. de Med-Chir. et Pharm./ 1758, pp. 244 and 337.—
Otto (Anat.), in 'Mag. d. Gesellsch. naturf./ 1814.— Owen, art.
"Entozoa," iu Todd's 'Cyclop.' — Rayer, 'Trait6 des maladies

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211

des reins, 1841. — Rayger, ' Sur un serpent qui sortit du
corps d'un homnie apres sa mort ' (quoted by Davaine, 1. c,
p. 272), 1675. — Schneider, ' Monographic der Nematoden/
1866, s. 50. — Idem (mit Peters), quoted by Leuckart, 1. c.,
s. 382. — Stratton, in 'Edin. Med. and Surg. Journ.,' p. 261,
1843.

Dochmius duodenalis, Leuckart. — Much time might be occu-
pied and wasted over the nomenclature of this parasite. In
my previous treatise, and for reasons there stated, I placed it
under the genus Sclerostoma. On rather slender grounds
Dubini formed the genus Anchylostoma for its reception, but
Yon Siebold thought that, on account of the absence of sym-
metry in the arrangement of the so-called dental organs,
Dubinins genus might very well be allowed to remain. Bilharz,
Diesing, Kiichenmeister, Wucherer, and others have retained
the genus as either Ancylostoma or Anchylostomum. Schneider
keeps it amongst the Strongyli ; but after all that has been said
and written there can, I think, be no doubt that if Dujardnr's
genus Dochmius is to be retained at all, Dubinins worm
must be placed in it. The comparisons instituted by Leuc-
kart afford sufficient proof of the intimate alliance as between
Anchylostoma and Dochmius. Professor Molin thought to
meet the difficulty by calling the worm Dochmius anchylos-
tomum, but the specific term, duodenale, should certainly be
retained.

This worm was discovered by Dubini at Milan, and though
at first thought rare, it is now known to be tolerably common
throughout Northern Italy. The worm has also been recently
found by Dr Kundrata at Vienna, in an Austrian subject.
According to Pruner, Bilharz, and Griesinger, it is abundant in
Egypt. Grriesinger believed that about one fourth of the
people of that country suffered from anaemic chlorosis, solely in
consequence of the presence of this worm in the small intes-
tines. From Wucherer's observations especially, we know that
Dubinins worm is not limited to the localities above mentioned,
for it occurs in the western tropics, in Brazil, and even in the
Comoro Islands.

The worm may be described as a small nematode, the males
measuring §" or rather more, whilst the females extend to very
nearly \" (12 mm.). The head is pointed and tapering,
and bent forward, having the mouth directed towards the
ventral aspect. The oral opening is armed with four asym-

212

PARASITES OF MAN

Fig. 41. — Male Dochmius duode-
nalis, with bursa separately en-
larged. After Kiicheumeister.

metrically disposed, unequally-sized, horny, conical, converging

teeth. The neck is continuous with
the cylindrical body, which is 5'0" in
thickness. The body terminates in a
straight cone-shaped, or rather sharply-
pointed tail in the female, the caudal
extremity of the male ending in a
partially inflexed, blunt point. In the
male there is a cup-shaped, bilobed
bursa, the membranes of which are
supported by eleven chitinous rays, ten
being simple, whilst the median, or odd
one, is bifurcated at the summit. The
mode of reproduction is viviparous.
Adult males and females occur in the
proportion of one of the former to
three of the latter.
As above mentioned, it was Griesinger who first pointed out
the clinical importance of this entozoon. He first explained
the manner in which the worm produces anaemia, the persons
attacked losing blood as if they were being bitten by innu-
merable small leeches. Like the rest of their kindred, these
worms are veritable blood-suckers. In the first instance the
views of Gi'iesinger met with opposition, but they have since
received abundant confirmation. Whilst Kiichenmeister's
' Manual ' furnishes an excellent account of the disorder as
known in Europe, we are chiefly indebted to Wucherer for
what is known of the disorder in Brazil. The experiences
recorded in the ' Deutsches Archiv fur Klinische Medicin ' for
Sept. 27th, 1872 (s. 379-400), were amongst the last that
appeared from the pen of that gifted and amiable physician.
As little or no notice of his writings appears to have been
taken by professional men in this country, I depart somewhat
from the design of this work when I venture to abstract a few
of the clinical particulars which he has supplied. Their im-
portance in relation to sanitary science is obvious, inasmuch as
these parasites are introduced into the human body by drink-
ing impure water, or, at least, water which either contains the
free larvae of the worm, or the intermediary bearers that harbor
the larvae.

It should be borne in mind that Dubinins original discovery
was made at Milan in 1838, whilst Griesinger's recognition

NEMATODA

213

of the worm as a cause of the Egyptian chlorosis resulted
from a post-mortem examination made on the 1 7th of April,
1851.

In the journal above mentioned, "Wucherer records his own
discoveries as follows (' Ueber die Anchylostomunkrankheit/
&c.) : — « Although Griesinger with well-founded confidence
gave an account of his ' find ' and its significance, yet it remained
for a long time unnoticed and unutilised, till at length a case
led me to corroborate it. During my many years' residence in
Brazil, especially during the first year, I had very frequent
opportunities for witnessing the tropical chlorosis, but seldom to
treat it, as it is one of those diseases for which Brazilians seek
no medical assistance. Its treatment falls to the lot of the
curiosos, curadeiros (quacks), who employ the fresh pulp of a
species of fig as a remedial agent with the best results. On
the 13th of December, 1865, I was called to the Benedictine
monastery in Bahia to see a slave of the order suffering from
hypoxemia. The patient was about thirty years of age, married,
a strongly built mulatto. He was a field laborer on the Ingua
plantation of the order, who exhibited in a conspicuous degree
all the symptoms that occur in hypoeemia except the diarrhoea.
He was well nourished, but strikingly pale, his whole face, but
especially the eyelids, being cedematously swollen, as also were
the feet, legs, and hands. The hands and feet were very cold.
His appearance betrayed the most horrible anguish or low de-
spondency. With difficulty only could he raise himself, being
obliged to lie down again immediately on account of his weak-
ness. Auscultation revealed a diminished respiratory murmur,
and bronchial expiration in both lungs. The pulse was very
rapid and small, the patient complaining of pain in the region
of the heart. He had frequent palpitation when he moved, and
he complained of pain in other parts of the body. His abdomen
was much distended by gases, but not sensitive to pressure from
without, except in the region of the stomach. The urine was
clear, its specific gravity 1007 to 1023^°. Under great diffi-
culties he resided for several months after his marriage at
Inhata. Earlier he had been on the estates of the order at
Rio de S. Francisco. He there suffered for a long time from
intermittent fever, but at Inhata he entirely recovered. At
Inhata the slaves frequently suffered from hypoxemia, but in
S. Francisco not at all. He appears not to have made any
misuse of brandy. The slaves of the order were well cared for

214

PARASITES OF MAN

and supplied with good and wholesome nourishing food. The
patient had already, for a long period, treated himself with
steel wine, yet was continually getting worse and worse. He
had not taken the pulp of the fig. As I was unaware he had
suddenly become so ill, they hastily despatched a message to
the town. There was no good to be expected from the further
employment of iron, and the patient was in such a condition that
from the very first I despaired of his recovery. I immediately
prescribed the pulp of the Gammeleira (Ficus doliaria), but
it could not be easily obtained. Considering that the Gamme-
leira would have a drastic effect, I therefore prescribed two
grammes of elaterium, to be divided into eight doses, of which
he should take one every three hours/'' Dissatisfied with this
advice, however, Dr Wucherer goes on to say that on reaching
home he carefully looked up the literature of the subject. " In
a ' Geologico-Medical Report ' by Professor Hirch, recorded in
the ninety-sixth volume of ' Schmidt's Jahrbucher,' I found
how Griesinger had recognised the Anchylo stoma as the cause
of the Egyptian chlorosis, which was clearly identical with our
hypoxemia. He had employed this commended anthelmintic. I
resolved the more to prescribe the pulp of the Gammeleira
when I found it described as a worm-expelling remedy in
Martin's ' Systema Materiae Vegetabilis Braziliensis/ The next
morning, however, when I arrived at the monastery I learnt
that my patient died about two hours after a slight evacuation.
Only after much resistance would they permit the sectio cada-
veris. I merely opened the abdomen, and was surprised to find
everything as Griesinger had described. During the next
season, through the courtesy of my colleagues attached to the
General Infirmary at Bahia, especially of Drs Silva Lima,
Faria, and Caldos, I was enabled to open more than twenty
bodies of anasmically deceased individuals. All were selected
as miserably poor in condition, but only five were bodies of
persons in whom hyposemia was diagnosed, and in these there
were a great number of Anchylostomes in the small intestine.
The intestines of the other bodies contained either none, one, or a
few." Dr Wucherer next states that he compared the characters
presented by his entozoa with those given by Dubini, Diesing,
and Von Siebold, and found a perfect agreement throughout.
He sent several examples to Griesinger, who also established
their identity, and communicated the results of his investiga-
tions accordingly (' Archiv fur Heilkunde/ 1866, s. 387.

NEMATODA

215

See also Leuckart, ' Die Mensch. Par./ Bd ii, s. 411). Dr
Wucherer also forwarded a number of specimens to Dr Weber,
who published a brief account of them with excellent figures
('Path. Soc. Trans./ vol. xviii, 1867, p. 274). As mentioned
in the text of his memoir (s. 394), Dr Wucherer also transmitted
some strongyloids to myself. " The publication of my observa-
tions," adds Dr Wucherer (' Gazeta Medica da Bahia/ 1866,
p. 27 et seq.), "had a result in that Dr J. R. de Moura, of
Thersepolis, in the province of Rio de Janeiro, sought for Anchy-
lostomes in the bodies of tropical anaemics (RypoeemiJcem). He
at once found these parasites, as stated in the same journal (for
1866, p. 132). As occurred to myself, he saw no enduring
results from the application of the remedies which appeared
to be called for, whilst he well knew that unprofessional persons
(Nichtarzte) succeeded in obtaining marked results by the
exhibition of the pulp of the Gammeleira (Ficus doliaria). The
anthelmintic action of this remedy was also unknown to him."
Dr. Wucherer then records how his discovery of these entozoa
was announced by Dr Jobini to the Rio academy, and how Dr
Moura's observations were subsequently communicated, adding
remarks upon the interesting discussion that followed. The
general opinion was that the Anchylostomata were not the
primary and. necessary cause of this tropical anaemia, but rather
a co-operating agent in its production. Against this view Dr
Wucherer afterwards very properly protested ('Gazeta/ Jan. 15th,
1868). In the mean time, says our author, " Dr le Roy de
Mericourt, prompted by my first communication, had invited
the physicians of the French colony to seek for Anchylostomes.
Drs Monestier and Grenet, at Mayotta (one of the Comoro Isles,
which lies about 12° S. lat. to the north-east of Madagascar),
ascertained the presence of entozoa in hyposemics. Dr Grenet
sent the duodenum and a portion of the jejunum of an hypo-
aemic corpse to Le Roy de Mericourt, who compared the Anchy-
lostomes with Davaine's description, and recognised them as
examples of A. duodenale."

" In the year 1868 Dr Rion Kerangel found Anchylostomes in
the bodies of hypoaemics in Cayenne. Thus, the occurrence of
Anchylostomes in hypoaemics has been authenticated by Primer,
Bilharz, and Griesinger, in Egypt; by myself, Dr Moura,
Dr Tourinho, and other physicians, in Brazil j by Monestier and
Grenet, in the Comoros; and by Rion Kerangel in Cayenne.
It thus also appears, from the wide separation of these several

216

PARASITES OF MAN

localities, that the Anchylostomes, if duly sought for, will be
found in many other countries."

These details given by Wucherer are so precise and instructive
that I could not have further abridged them without injustice
to his record. The bearing of the foregoing facts in relation
to the question as to how we may hope to arrest the fatal action
of many of these nematodes is sufficiently obvious. That
strongyles and their allies prove highly destructive to man and
beast is as well established as any other recognised conclusion
in medical science ; nevertheless, there are those who still doubt
the power of these nematodes in relation to the production of
fatal epidemics. I shall deal with the sanitary bearings of the
subject hereafter. In conclusion, I may mention that Dr da
Silva Lima has forwarded specimens of Anchylostomum to the
Hunterian Museum, where they may be seen.

Bibliography (No. 29). — Bilharz, ' Zeitschr. f. wiss. Zcol./
Bd. iv, s. 55. — Cobbold, ' Entozoa/ p. 361. — Idem " Remarks on
Recent Contributions to our Knowledge of the Parasitic Nematoids,
especially in reference to the Wasting Diseases they produce in
Man and Animals," the ' Veterinarian/ Jan, 1876, p. 1. —
Davaine, 1. c, pp. 118 and 931. — Diesing " Revis. der Nema-
toden," ' Sitzb. d. m.-naturw. cl. d. k. Akad./ 1860, s. 716. —
Dubini, ' Entozoografia/ &c, 1849. — Griesinger (quoted above),
see also f Arch. f. Phys. Heilk./ 1854. — Kiichenmeister, 1. c,
Eng. edit., p. 383. — Leuckart, 1. c, ss. 410-455. — Molin,
' II sottordine degli Acroffali/ p. 61 (quoted by Leuckart). —
Siebold, 'Zeitsch. f. wiss. Zool./ 1852, s. 55. — Sonsino, P.,
L' Anchilostoma duodenale in relazione colF Anemia progressiva
perniciosa/ Egitto, 1877. — Idem, ' Sull.' Anch. duod., 1878 (see
also Bibliog. No. 27, both reprinted from ' Imparziale/) — Weber,
E., 1. c, 1867. — Wucherer (quoted above), 1872.

Dracunculus medinensis, Cobbold. — This parasite is popu-
larly known as the guinea-worm, or Medina-worm. Pro-
bably Lister was the first writer who distinctly spoke of it
as the Dracunculus, 1690, the same title being applied to it by
Kaempfer, 1694. Be that as it may, Gmelin, long afterwards,
placed the parasite in the genus Filaria, at the same time
adopting the specific title medinensis. This had been pre-
viously employed by Linneus, who, however, regarded the worm
as belonging to the genus Gordius. It being clear from the
distinctive characters of the entozoon that it was desirable to
separate it from the Filarite, and that no better generic name

NEMATODA

217

could be devised than Dracunculus, I thought it right to
combine Lister's and Gmelin's nomenclature as above, 1864.
Leuckart pursued a similar course, crediting Linneus with the
titles.

The guinea- worm having been known from the earliest times,
it is not surprising that its true nature long remained a
mystery. Any one who has read Kuchenmeister's elaborate
narrative of the historical significance of the Dracunculus will
hardly have failed to arrive at the conclusion that Moses was
probably the earliest writer on the endemic disorder which is
occasioned by this parasite. There can be no doubt that the
" fiery serpents" which afflicted the children of Israel during
their stay in the neighbourhood of the Red Sea were neither
more nor less than examples of our Dracunculus. It is further
evident that Plutarch spoke of Dracunculi, when in the eighth
book of his ' Symposiacon/ he quotes Agatharchidas as stating
that the people taken ill on the Red Sea suffered from many
strange and unheard-of attacks, amongst other worms, from
" little snakes, which came out upon them, gnawed away their
legs and arms, and when touched retracted, coiled themselves
up in the muscles, and there gave rise to the most insupportable
pains/' In order to render the passage more readable, it will
be seen that I have slightly altered the original version (f Para-
siten/ s. 305).

The guinea-worm may be described as a nematode mea-
suring from one to six feet in length, having a thickness
of T'5th of an inch. The body is uniformly cylindrical, termi-
nating below in a more or less curved and mucronately pointed
tail. The head is flatly convex or truncate, having a central,
simple mouth, which is surrounded by four equi-distantly and
cruciately disposed papillae. The mode of reproduction is
viviparous, the body enclosing a prodigious number of hatched
embryos, which, by distension of the uterine ducts, almost
entirely obliterate the somatic cavity. Notwithstanding the
statements of Owen to the contrary, the male Dracunculus is at
present altogether unknown.

The guinea-worm possesses a comparatively limited geogra-
phical range, for not only is it proper to the tropical regions,
but within intertropical limits it is almost exclusively confined
to certain districts in Asia and Africa. Thus, according to
Kiinsenmuller, as quoted by Busk, it occurs endemically in Arabia
Petraea, on the borders of the Persian Gulf and Caspian Sea, on the

218

PARASITES OF MAN

F;o. 42.— Outline of n fe-
male Dracunculus medi-
nensis. JSat. size. Ori-
ginal.

banks of the Ganges, in Upper Egypt, Abys-
sinia, and the coast of Guinea. " In America
the guinea-worm is unknown, except in per-
sons who have had communication with Africa
or other parts where it is indigenous. The
island of Curacoa is the only locality in the
New World which offers an apparent excep-
tion to this fact, and it would be highly
desirable to ascertain the real state of the case
in this instance." The observations of Chis-
holm showed that the Dracunculus is really
prevalent in several of the West Indian islands,
especially in Grenada, and the still later investi-
gations of Dr Da Silva Lima point to its
former prevalence in Brazil. Now, the worm* is
rarely seen at Bahia. Mr Busk said: — "Though
endemic only in the above-mentioned parts of
the world, it would yet appear that all races
of mankind are obnoxious to the attacks of
the Filaria when exposed to what may be
called the contagion ; that is, when placed in
circumstances under which it might be sup-
posed a contagious seminium could be conveyed
to them." Mr Busk also added : — " I have
known many instances tending to prove that,
in order that a European should become in-
fected with the guinea-worm on the coast of
Africa, it is not necessary that he should have
been on shore at all. It has been quite suf-
ficient for him to have exposed the bare
surface of some parts of his person to the
water in the native canoes alongside, or, it
may be, to the discharge from the sores of
those laboring under the disease. This mode
of its introduction accounts for the frequency
with which the legs and feet are attacked
by the parasite, in preference to other parts
of the body, as it will always, I believe, be
found that the men who have become so affected
have been in the habit of going about with bare
feet, as is common among sailors in warm
latitudes. That the contagious material is

NEMATODA

219

conveyed in water is also farther indicated by the well-known
fact that in India, where it is the custom of the natives to carry
water in skins on their backs, the worm makes its appearance
on the back and shoulders and upper part of the body."
These views were published by Busk in 1846, and I am free to
confess that — confirmed as they appeared to be by subsequent
and independent testimony — they completely dominated my con-
ceptions as to the mode of ingress of the young parasites within
the human bearer. Thus, those of our Indian troops which
were most exposed during the rainy season, subsequently
exhibited evidence of having been invaded by the Dracunculus.
As, moreover, the period of incubation of the entozoon com-
monly extends from twelve to fifteen months, it necessarily
happened that the disease often showed itself in localities far
distant from the spot where the troops originally contracted the
disorder. The statement that the period of incubation of the
worm is not less than a year, is probably incorrect, since Carter
mentions that in a school of fifty boys bathing in a certain pond
at Bombay — the sediment of which swarmed with microscopic
tank-worms (Urobales palustris, Carter) — twenty-one were
attacked with Dracunculus during the year, whilst the boys of
other schools, bathing elsewhere, remained, with few exceptions,
uninfected. This is a remarkable occurrence, and it points to
the possibility of the young Dracunculi being confined to
particular pools. That they should, whether occupying the
bodies of intermediary bearers or not, be more abundant in some
waters than others, is just what might be expected, since such
a distribution is in harmony with a recognised law affecting the
abundance or limitation of species in particular localities.
Much, indeed, has been written respecting the nature of the soil
and geological formations occurring in the Indian worm-
districts, but the speculative views enunciated on this point are
little worthy of credit. Those who desire information on this
head should at all events consult the valuable writings of
Smyttan, Grreenhow, Bird, Forbes, Chisholm, and Aitken,
who, apart from the question at issue, supply abundance of
practical information.

Into the anatomy of the adult Dracunculus I do not enter,
but I may remark in passing, that the structure of the worm
has been exhaustively treated of by Busk and Bastian. A
resume of their views is given in my introductory treatise. Carter
and Lcuckart have also added important details. As regards

220

PARASITES OF MAN

the structure and development of the young worms, I have to
observe that the discovery of the viviparous mode of reproduc-
tion in Dracunculus is due to Jacobson. Nearly a quarter of a
century ago I recognised the fact that the uterine organs of the
adult worm almost completely filled up the perivisceral cavity,
and that they were crowded with microscopic worms. Referring

Fig. 43.— a, b, Head and tail of the adult guinea-worm (magnified 10 and 18 diameters
respectively); c, embryo (magnified 500 diameters). Original.

to this " find/' the late Sir George Ballingall, of Edinburgh,
in his well-known work on ' Military Surgery/ recorded tin1
circumstance in the following terms : — " The Assistant Con-
servator of the Anatomical Museum in our University has
detected in the oviduct of an adult specimen from my collection
myriads of minute and perfectly-developed (embryonic) Dra-
cunculi. They can be very well seen with an half-inch object-
glass, but their structure is best exhibited if the magnifying

NEMATODA

221

power be increased to two hundred and fifty diameters linear."
As already stated in my introductory treatise, these observations
w ere made during the winter of 1853-54. In July, 1854,
M . Eobin made a similar statement after examining a fresh
Dramnculus which had been extracted from the leg of a man
by M. Malgaigne. Eobin, not unsuitably, compared the worm
to a double tube, one tubular sheath, as it were, enclosing the
other. " The second tube/' he distinctly affirms, " is the
oviduct, or, rather, that -part which represents the uterus. The
young still remaining in the uterus were nearly all coiled, some-
times with the tail sallying outwards, at others rolled like the
rest of the body." I have thought it only due to Eobin and
myself to show that from the first we were perfectly well
acquainted with the fact of the " great development of the
genital tube and of its close adherence to the parietes of the
body." To be sure, many discrepancies occurred in our
writings, and in those of Busk and Carter. It was Bastian' s
skill and good fortune to correct these errors. Thus, most of
us agreed in recognising a slightly trilobed or tripapillated
mouth ; but Carter failed to demonstrate the existence of these
tubercles, and spoke of the oral aperture as being simple and
" punctiform." The body throughout its three upper fourths
appeared to me to be cylindrical, but Eobin found that it was
flattened. It is finely striated transversely, except at the part
where it contracts to form the slender, pointed tail. According
to Carter, Eobin, and Davaine, the young attain a length of
about 33 of an inch, but Bastian gives it as about . In
thickness, Carter gives the approximative diameter as ^ ',
Eobin makes it — ' to whilst Bastian gives their breadth

at y^a", and Davaine at ^5//. I estimated their greatest length
and breadth to be ^' by '. Eobin and myself thought we
recognised a distinct, rounded, anal orifice ; and whilst Busk,
on the one hand, saw nothing which, in the slightest degree
indicated the presence of an anal opening, Carter, on the other
hand, described the structure which we called the anus as a
gland, at the same time placing the alimentary outlet on one
side and a little above it. According to Bastian, " the intes-
tinal.tube is about ~" in length, and appears to consist of a
simple canal of varying calibre, pursuing a nearly straight
course, and terminating exactly at about the middle, in length,
of the worm. Like Eobin, Bastian recognised oesophageal and
stomachal divisions, and in a few examples he observed the

222

PARASITES OF MAN

= ==

1-!

cascal or terminal portion of the intestine to be partially reflected

upon itself. In regard to the circular
opening which Robin and myself de-
scribed as the anus, Bastian says there
is a rounded body, " about in dia-
meter, with a dark or light spot in the
centre, according to the varying focal
distance, and which seems to represent
a central aperture. Sometimes, above
this, traces of two or three large cells
may be recognised, whilst behind no-
thing definite can be made out, save that
the cavity of the body is visible for
about In other specimens of the

young worm the central body and spot
are wanting, but, in its stead, two lateral
sacculi are met with, about in dia-
meter, that communicate with the ex-
terior by a minute channel through the
integuments, which can sometimes be
distinctly recognised. At other times
|| the channel is obscured by protrusion,
which appears to have taken place
through it, of a minute bilobed papilla,
projecting from the side of the

body. When the projections are seen,
the sacculi are indistinct."

As Bastian found the young in all
stages of development from the germ
condition in diameter up to the per-
fect embryo, and as, moreover, he, like the

Fig. 44. — Embryos of Bracunculus. .
Minified 500 diameters. After rest OI US, COUld detect DO Sexual Orifice

in the adult Dracunculus, he was led to
express his belief that the young were produced agamo-geneti-
cally. He went so far as to call the germs pseudova. It was with
great reluctance that I dissented from the views of so gifted an
observer as Bastian ; nevertheless, later researches have shown
that I was justified in not hastily concurring in the theory of a
non-sexual mode of reproduction for Dracunculus.

Among the many advances of modern helminthology, the
discovery of the true source of the guinea- worm is not the least
important. To the late M. Fedschenko (the lamented and

NEMATODA

223

accomplished Kussian traveller, who lost his life in a snow-
storm on the Alps), science stands indebted for this memorable
advance. Fedschenko showed that the embryos of Dracunculi,
after quitting the human host, succeed in effecting an entry
into the bodies of entomostracous crustaceans belonging to the
genus Cyclops. Within these intermediary bearers, after
twelve hours' sojourn, the embryos undergo a change of skin,
attended with subsequent growth. Here they remain to com-
plete their larval development, which takes place within a
period of five weeks, or, as Fedschenko himself told me, one
month and six days. At length, as perfected larvae, they are,
together with their crustacean hosts, transmitted to the stomach
of the ultimate or human bearer. It is probable that sexual
maturity is next acquired within the human stomach, copula-
tion following. After this, the females migrate to the situa-
tions in which they are found beneath the skin of the human
bearer, whilst the males perish and pass out with the fasces.
Thus much I gathered from M. Fedschenko himself when he
visited this country, and I possess a sketch of the larvae made
by him at the time (October 23rd, 1873). One of the figures
represents a larva which has undergone ecdysis, the long and
narrow embryonic tail being supplanted by one which is blunt-
and forked at the tip. The somatic contents of the embryo
have at the same time differentiated into a complete intestinal
tube, and a constriction marks the junction of the oesophagus
with the stomach. There is also internally an oval-shaped
mass of cells near the centre of the body. These represent
the commencement of the reproductive organs.

What I had gathered from Fedschenko in conversation thus
epitomises that which has since been much more fully stated
by Leuckart j and it is only fair to add that the Eussian
traveller was led up to his discovery by the previous investi-
gations of Leuckart respecting the young of Cucullanus. The
Leipsic helminth ologist had, indeed, specially instructed Fed-
schenko as to the probable source of Dracunculus.

It is often thus that science makes its clear advances, since
a master-mind is needed to set others on the right track. The
embryos of Cucullanus and Dracunculus bear a close resem-
blance to each other, and the similarity of the types is
continued on, though not in the same degree, in the next stage
of larval growth, after ecdysis. The higher larvae of both have
their tails trifurcate at the tip, the head of the Dracunculus-

224

PAKASITES OP MAN

larva being distinguished by the presence of a pair of papilla).
In the case of Cucullanus the embryos are, according to
Leuckart, passively transferred to the stomach of Cyclops by
the mouth ; but in the case of Dracunculus, Fedschenko saw the
embryo in the act of perforating the bodies of the little Crustacea
at the ventral surface, where the segments are bound together
by a thin and easily penetrated connecting membrane. The
larvae then proceed to coil themselves within the limbs, as many
as six or even a dozen of the parasites being occasionally
found within the body of a single crustacean host. "When they
have reached full larval growth they measure about ^' in length.
Of course, after attaining this stage, it is a matter of conjecture
as to the precise way in which their final destiny is accomplished.
Fedschenko fed dogs and cats with the infected Crustacea, but
failed to rear Dracunculi in these animals. Clearly, these
carnivora were unsuitable hosts. Could Fedschenko have
experimented on man the result would probably have been very
different. Arguing from what happens in the case of Cucul-
lanus amongst fishes, and Trichina in man, there can be little
doubt that all the further and final changes undergone by the
larvae are accomplished within the human host. These changes
are usually, if not invariably, consequent upon a direct trans-
ference of the infested entomostraca along with water used as
drink. Thus, it must at once be evident that the simple
sanitary precaution of filtering water before use is amply suffi-
cient to ensure the prevention of attacks of dracontiasis or the
guinea-worm disease. The theosophical remedy of Moses
against this invasion by fiery serpents, as the worms were called
in his time, and the modern prophylactic measures dictated
alike by science and common sense, thus stand in striking
contrast the one to the other. In the nature of things it must
ever remain that unreason and reason will select diametrically
opposite methods of action, equally, no doubt, with the good
intention of bringing about beneficial results.

From what has now been advanced, it will be seen that as
regards the mode of infection the views categorically expressed
in my previous work (' Bntozoa/ p. 387) cannot be maintained.
What, however, is there stated in respect of treatment still holds
good in the main, even as regards prophylaxis.

Bibliography (No. 30) — Adam, ' Trans. Med. and Surg.
Soc./ Calcutta, 1824— Aithen, W.t 'The Science and Practice
of Medicine/ 6th edit.., vol. i, 1872.— (Anonymous), " Review of

NEMATODA 225

the writings and opinions of Duncan, Johnson, Bird, Mylne,
Kennedy, Chisholm, H. Scott, A. J. Robertson, Smyttan,^
Macgregor, Thomas, Mosely, Morehead, Twining, and others,
on the Dracunculus or Guinea-worm," in < Corbyn's India Journ.
of Med. and Phys. Sci./ vol. ii, p. 118, 1836.-(Anon.), "The
Guinea-worm very Prevalent at Bokhara," ' Boston Med. and
Surg. Journ./ 1843, p. 387.— Balfour, J., ' Ind. Ann. Med.
Sci./ 1859, p. 175.— Ballwgall, 0. (1. c, supra), 1854,— Bashan,
E. C, " On the Structure and Nature of the Dracunculus or
Guinea-worm," ' Linn. Soc. Trans./ vol. xxiv, p. 101, 1863.—
Berncastle, J., in the 'Lancet/ 1851.— Bird, J., ' Calcutta Med.
and Phys. Trans./ 1825, p. lhl.—Bremser (1. c, Bibl. No. 2),
s. 194— Brett, ' Surgical Diseases of India/ 1840; see also
'Med.-Chir. Rev./ 1841.— Bruce, N., ' Edin. Med. and Surg.
Journ./ 1806, vol. ii, p. 145.— Busfc, G., ' Micr. Soc. Trans/
(original series), 1846.— Carter, E. J., " Note on Dracunculus in
the Island of Bombay," ' Bombay Med. and Phys. Soc. Trans/
(new series), No. 2, p. 45, 1853-54 ; see also postscript, p. 252.
—Idem, " Further Observ. on Dracunculus," ' Bomb. Med. and
Phys. Soc. Trans/ (new series), No. 4, p. 215, 1857-58.— Idem,
" On Dracunculus and Microscopic Filaridse," ' Ann. of Nat.
Hist./ vol. iv (third series), 1859,— Idem, "Notes on Dracun-
culus," &c, ' Ann. of Nat. Hist./ vol. ix (third series), 1862.—
Chapotin, ' Bull, des Sci. Med./ 1810.— Oharvet, 'Ann. des Sci.
Nat./ 1834.— Ohiaje (1. c, Bibl. No. 2), p. 99.— Chisholm, C,
"On the Malis Dracunculus or Guinea- worm (in Grenada),"
' Edin. Med. and Surg. Journ./ vol. xi, 1815 ; see also the
'Veterinarian/ vol. ix, p. 508, 1836.— Clark, < Med.-Chir. Rev./
1840.— Clarhson, N. F., "Alleged Case in the Horse," the ' Vete-
rinary Record/ 1845, p. 73. — Clot-Bey, 'Apercu sur le ver
dragonneau observe en Egypte/ 1830. — Cobbold, ' Entozoa/
p. 373. — Cuvier, ' Regne animal/ Orr's Eng. edit., 1849, p. 644.
—Davaine, ' Traite/ 1. c, edit, ii, p. 783 (full lit. refs.), 1878.
— Diclcson, 'Path. Soc. Trans./ 1851. — Drummond, 'Med. Com-
mentaries/ 1793, p. 294. — Dubois, ' Edin. Med. and Surg.
Journ./ vol. ii, 1806. — Duncan, ' Calcutta Med. and Phys. Soc.
Trans./ 1835. — Ewart, J., "Questions relating to Dracunculus,"
in a review of his memoir on the " Vital Statistics of the Mey war
Bheel Corps," in the ' Madras Quart. Journ. of Med. Sci./ vol. i,
1860, p. 462. — Fedschenlw, ' Protocol of the Promoters (Freunde)
of the Natural and Physical Sciences at Moscow ' (in the

Russian language), 1869 and 1874 (quoted by Leuckart).

226

PARASITES OP MAN

Forbes, D.," Observ. on Dracunculus" (extr. from the 'Half-yearly
Reports of the diseases prevailing at Dharwar in the 1st Grenadier
Regiment, in the year 1836 '), ' Bombay Med. and Phys. Soc.
Trans./ vol. i, 1838, p. 215. — Gibson, A., "Note on the Preva-
lence of Dracunculus," in his remarks on the " Diseases of the
Deckan," in ' Bomb. Med. and Phys. Soc. Trans./ vol. ii, 1839,
p. 209. — Gramberg, ' Geneeskundige tijdschrift voor nederl.
Indie/ 1861, p. 632 (quoted by Leuckart). — Greenhow, H. M.,
'Indian Ann. of Med. Sci./ vol. vii, 1861, p. 31. — Grierson, D.,
" Observ. on the Dracunculus, as it prevailed in the 22nd
Eegiment, N.I., from April till September, 1841," ' Bomb.
Med. and Phys. Soc. Trans./ No. 4, 1841, p. 90.— Grundler, in
'Commerc. Litt. Nov./ 1740, p. 239. — Henderson, J., "Note
respecting Four Cases of Dracunculus in the 48th Regiment,"
'Madras Quart. Journ./ vol. iii, 1841, p. 353. — Horton, J. A. B.,
'Army Med. Reports/ 1868, p. 335. — Kennedy, B. H., ' Calcutta
Med. and Phys. Soc. Trans./ 1825, p. 165. — Kuchenmeister (1. c,
Eng. edit.), p. 389— Leuchart (1. c, Bibl. No. 1), s. 644-725.
Lewis, T. B., in ' On a Hsematozoon/ &c. (1. c, Bibl. No. 23),
p. 30 et seq. — Lima, Da S., "Remarks on the Filaria medinensis,
or Guinea- Worm ; on the occurrence of this Parasite endemic-
ally in the Province of Bahia ; on its entrance into the human
body by drinking water," in the ' Veterinarian/ Feb., March,
et seq., 1879.— Lister, ' Phil. Trans./ 1690, p. 417. — M'Glelland,
J., ' Calcutta Journ. of Nat. Hist./ vol. i, 1841, p. 366. —
M'Grigor, J., "On the Guinea- w or m " (in his "Account of the
Diseases of the 88th Regiment in Bombay"), ' Edin. Med. and
Surg. Journ.,'' vol. i, 1805, p. 284. — Morehead, G., ' Calcutta
Med. and Phys. Soc. Trans./ vol. vi, 1833, p. 418 ; also
noticed in ' Edin. Med. and Surg. Journ./ vol. xliv, 1835. —
Idem, part ii, ' Calcutta Med. and Phys. Soc. Trans./ vol. viii,
1836-42. — Murray, J., "Guinea-worm a very Common Disease
at Sattara" (in his Official Report on the Hospital, &c),
' Bombay Med. and Phys. Soc. Trans./ No. 9, art. vi, p. 198,
J 847.— Ohe, W. 8., " Case of Guinea- worm," ' Prov. Med. and
Surg. Journ./ vol. vi, 1843. — Oldfield, " Case of Dracunculus "
(from Laird and Oldfield's "Narrative of an Expedition into
the Interior of Africa"), 'Dublin Jomm./ vol. xii, 1838. —
Baton, " Cases of Guinea- worm," ' Edin. Med. and Surg. Journ./
vol. ii, 1806. — Baddoch, "A Case of Guinea-worm," 'Indian
Med. Gaz./ Oct., 1877, p. 265.— Scott, W., " Remarks on the
Dracunculus," in a letter to the Medical Board, Madras, ' Edin.

NEMATODA

227

Med. and Surg. Journ./ vol. xvii, 1821. — Leverancc, G. IE.,
"History of a Case of Guinea-worm/' from 'Amer. Med. Times/
in the 'Glasgow Med. Journ./ vol. ix, 1861-62, p. 377.—
Smyttan, G., " On Dracunculus/' ' Calcutta Med. and Phys.
Soc. Trans./ vol. i, 1825, p. 179. — Stewart, L. W., 'Indian Ann.
of Med. Sci./ vol. vi, 1858, p. 88.— Twining, W., " Cases of
Dracunculus," ' Calcutta Med. and Phys. Soc. Trans./ vol.
vii, 1835.

Oxyuris vermicularis, Bremser. — Of all the parasites infesting
the human body this is the one concerning which the medical
practitioner is most frequently consulted, partly on account of
its remarkable frequency in children, and more particularly on
account of the difficulty often experienced in getting perma-
nently rid of it. The Oxyuris vermicularis is by no means con-
fined to young persons, seeing that adults are infested even to
old age. It is familiarly known as the threadworm or seat-
worm. The male measures about 5", and the female from ~"
to in length. The female possesses a long capillary tail,
which terminates in a three-pointed end. The extremity is
said to act as a kind of holdfast. The tail of the male is obtusely
pointed. In both sexes the body presents a more or less fusi-
form shape, the anterior end being narrowed to form a some-
what abruptly-truncated head, which is often rendered very
conspicuous by a bulging of the transparent integument sur-
rounding the mouth. This presents in profile the aspect of
winged appendages (fig. 45). The oral opening is tripapillated,

Fig. 45.— Head of Oxyuris vermicularis. Highly magnified. After Busk.

leading into a triangular oesophagus. The integument is trans-
versely striated, and of a silvery-white appearance. The'
spicule is simple, single, and very minute. The eggs are
oblong and unsymmetrical. They measure about fa" from pole
to pole, and transversely.

Many years back (1863) I pointed out that the most advanced

228

PARASITES OF MAN

eggs whilst still within the body of the pregnant female
contained tadpole-shaped embryos, and about the same time
the fact was noticed by Claparede. In his beautiful and
scholarly memoir, ' De la formation et de la fecondation des
ceufs chez les vers Nematodes/ he wrote concerning the ova as

1<IG 46— Section of a female Oxyuris vermieularis, magnified 220 diameters (after Busk);
and also several free eggs (original), a. With an imperfectly formed embryo ; b, c, d, with
three tadpole-shaped embryos, magnified 450 diameters.

follows : " The egg, which exhibits the form of a very narrow

disk in the ovary, acquires the shape of an elongated ellipsoid
in the oviduct, and at the surface differentiates itself into a very

NEMATODA

229

thick vitelline membrane. Then it forms a strong and resist-
ing chorion, which imparts to the egg an outline similar to that
of a bridge's span. It has an oval figure flattened at one of its
sides. This chorion is very fragile; it frequently gives way
under slight pressure from the thin plate of glass which covers
the object. It extends itself considerably under the action of
acetic acid, acquiring a size three or four times greater than
that of the egg. The constitution of this chorion is perfectly
identical in the eggs both before and after impregnation. It
is, nevertheless, easy at first sight to know whether or not we
have to deal with a fecundated egg. In the impregnated females
the uteri are filled with thousands of ova, each one of which
encloses an embryo already well formed. The ventral surface
of the embryo and the tail are, without exception, applied to
the flattened side of the egg. The embryo is very broad in the
body, and occupies all the interior space. An embryo such as
Kiichenmeister has represented under the form of a small fili-
form worm folded on itself, and only occupying a very small
part of the cavity of the egg, is never to be seen. In the non-
fecundated females, on the other hand, the uteri are filled with
eggs, which, instead of the embryo, enclose a non-segmented
yolk furnished with a large germinal vesicle. This vesicle is
not visible so long as the eggs have the form of thin disks ; it
only shows itself when the eggs begin to acquire an elliptical
form in the oviduct. It is, however, probable that this vesicle
is the same which was originally visible in the ovary.-" The
chorion itself is homogeneous, but in an allied species (Oxyuris
spirotheca) Gyoery and Claparede found that this egg-covering
consists of spirally-coiled bands resembling the tracheal spiral
fibre of an insect. Under suitable conditions the tadpole-
shaped embryos rapidly assume a vermiform character. The
investigations of Leuckart have shown that " one only needs to
expose the eggs to the action of the sun's rays in a moistened
paper envelope when, at the expiration of five or six hours, the
tadpole- shaped embryos will have already become slender
elongated worms." According to Heller, the simplest way to
rear the vermiform stage of Oxyuris is to put a number of the
eggs in a glass tube filled up with saliva. The tube should
then be placed in the arm-pit, in which situation it can be
carried about with little inconvenience. In a few hours the
transformations will commence and go on continuously until the
vermiform condition is attained. If, as remarked in mv

230

PARASITES OP MAN

' Lectures/ it be asked whether the embryos which have
escaped into the bowel are capable of arriving at the vermiform
stage, the answer is in the affirmative ; for, as Leuckart says,
" the elongated embryos are to be found not only in the fasces
but also in the mucus of the rectum above and around the anus."
Yix has also asserted that free vermiform embryos are occa-
sionally to be detected in the intestine of the human bearer
along with the eggs ; this hatching within the lower bowel,
however, must, in my opinion, be regarded as exceptional.
Heller is of the same opinion. According to Leuckart, the
escape of the embryos from the eggs " ordinarily takes place
under the action of the gastric juice, also primarily in that
condition when they have by some means or other gained
access to a new bearer." Prof. Leuckart and three of his
pupils courageously infected themselves by swallowing the eggs,
and had the satisfaction of observing young Oxyurides in their
stools fifteen days afterwards.

From the united labors of Professors Zenker and Heller it is
now rendered certain that all the further changes necessary to
bring the larvse to sexual maturity are accomplished within the
small intestines of the human bearer ; and it is not necessary
that a change of hosts should occur at any time during the life of
the parasite. Infection ordinarily takes place by the accidental
and direct conveyance of the eggs that are lodged in the neigh-
bourhood of the victim's anus to the mouth. Since the victim
may accomplish this during sleep, it is not in all cases fair to
charge infected persons with uncleanliness. On the other hand,
it too often happens that due care in this respect has not been
exercised, and from such persons you may remove the eggs of
Oxyurides from the margins of the finger nails. One aristo-
cratic person, who was infested by myriads of these entozoa,
confessed to me that in his extreme distress, and consequent
rage, he had freely bitten the live worms in halves between his
teeth. He had thus exposed himself to a terrible revenge,
since multitudes of the ova entering his mouth subsequently
found their way into the stomach and intestines. By whatever
mode the eggs are conveyed to the mouth their subsequent
passage to the stomach ensures their being hatched. In the
duodenum and other divisions of the small intestines, as Zenker
and Heller have shown, the embryos undergo transformation,
casting their skins, and growing with great rapidity. Probably
not more than three weeks or a month is necessary to complete

NEMATODA

231

their growth. Heller obtained mature worms from an infant
only* five weeks old. Finally the worms are transferred to the
caecum, which constitutes, so to speak, their headquarters. It
is an error to suppose that the lower bowel or rectum forms
their especial habitat, nevertheless the most approved manuals,
vade mecums, and general treatises have for a long time sup-
ported this erroneous view. The error had been pointed out by
Stricker in 1861.

The symptoms produced by Oxyurides are occasionally very
serious. In the mildest cases they have a tendency to under-
mine the health. As remarked
in my 1 Bntozoa/ the unpleasant
sensations chiefly develop them-
selves in the evening and at night,
consisting for the most part of
feelings of heat and irritation within
and around the margin of the anus.
The symptoms may become ex-
tremely distressing and almost in-
tolerable, especially when the itch-
ing extends to the genito-urinary
passages, in consequence of the
escape and migration of the para-
sites about these parts. By-and-
by various sympathetic phenomena,
such as restlessness, general ner-
vousness, itchings at the nose, in-
voluntary twitchings, grinding of
the teeth during sleep, chorea, con-
vulsions, and even epileptiform
seizures, may supervene. At the
age of puberty special local dis-
orders arise, the nature of which will

be readily understood when merely Fig. 47- — Adult male Qxyuris vermicularis.
1 o . i 1 • -| i Magnified. After Kiichenmeister.

spoken 01 as the morbid pheno-
mena of sexual irritation. In the female the occurrence of
pruritus and leucorrhcea is not uncommon, accompanied or
not, as the case may be, with hysteria in various forms. There
is usually general asthenia, with more or less emaciation. The
anaemia is sometimes remarkable, but in place of anorexia,
which is, however, an occasional symptom, one frequently finds
a most voracious appetite, especially in young people. Some-

232

PARASITES OP MAN

times there are obscure symptoms simulating those of local
organic disease.

About the treatment of the disorder I have nothing to say-
here, further than to urge the benefits of the preventive
measure of cleanliness. Like Zenker and Heller, I have ob-
tained the eggs of oxyurides from beneath the finger-nails of
young people. In one lad all the nails had been carefully
bitten down to their roots, but from beneath a minute project-
ing portion that was left on the right fourth-finger I procured
two eggs. Their demonstration under the microscope con-
vinced both parent and child of the necessity of frequently
employing local and general ablutions. Personal cleanliness is
essential. In this connection an able biologist has ventured to
hazard a statement to the effect that " probably any infected
person who adopted the requisite precautions against reinfection
from himself or others would get well in a few weeks without
treatment by drugs." ' Dr Ransom bases his belief on the
known facts of the life-history of this entozoon, as recorded
more especially by Leuckart. I regret that I cannot fully
share Dr. Hansom's views, and still less should I think it right
by my silence to seem to endorse his statement to the effect
" that every person who is shown to be infested with those very
common entozoa, Oxyuris vermicularis and Trichocephalus dispar,
is thereby demonstrated to have swallowed minute portions of
his own or another person's fasces." This is putting the case
too strongly. No doubt the eggs of oxyurides swallowed by
ourselves must have previously passed through some person's
rectum ; as such, either separately or mayhap collectively, in
the body of the maternal parasite. That does not, however,
justify the statement, that we " have swallowed " part of our
own or of some other person's excrement. The eggs ought not
to be regarded as constituent portions of the faecal matter. Per-
haps Dr Eansom will say that the surfaces of these eggs, being
in contact with faecal matter, must carry infinitesimal particles
on their surfaces, and it is to such that he refers. As, however,
a large proportion of the ova escape with their parents, whilst
they are still lodged within the maternal worm, it cannot be held
that these intra-uterine ova carry fa3cal matter on their shells.
Commonly the eggs are swallowed in the separate, free, and dry
state. In water they perish quickly. The act of eating with
unwashed hands is a fertile source. of infection, more especially
if the meal be taken either in bed or in the bedroom.

NEMATODA

233

Bibliography (No. 31).— Alexander, J., "On Vermination,"
< Lancet/ 1833.— Anderson, W., " On Santonine, with especial
reference to its use in Koundworm and Threadworm/' ' Brit.
Med. Journ./ April, 1864, p. 443 j also in Braithwaite's ' Re-
trospect of Medicine/ vol. xlix (synopsis, p. 20), 1864.— Barry,
J. M., " On the Origin of Intestinal Worms, particularly the
Ascaris vermicular is, ,} ' Trans. Assoc. of Fell, and Licent. of
King's and Queen's Coll. of Phys. in Ireland/ vol. ii, 1878, p.
383.— Bremser, 1. c, s. 79. —Buckingham, " Ascarides causing
Erotomania/' from 'Bost. Journ., U.S./ in ' Med. Gaz./ 1857.
—Glajparede, E., " On the Formation of^the Egg and Fertilisa-
tion in the Nematoidea," from the ' Zeitsch. f. w. Zool./ trans-
lated by Dallas in ' Ann. Nat. Hist./ vol. i (third series), 1858. —
Idem (memoir quoted in the text above), Geneve, 1859. — Gob-
bold, T. S., ' Worms/ Lect. xii-xvy 1872 — Idem, ' Entozoa/
p. 362 — Idem, 'Brit. Med. Journ./ Aug., 1873. — Idem, 'Tape-
worms and Threadworms/ 2nd edit., 1872. — Idem, ' Lancet/
1866. — Idem, " On the Development and Migrations of the
Entozoa/' ' Brit. Assoc. Bep./ 1864, p. 116. — Date, W.,
' Lancet' for Feb., 1872, p. 185. — Davaine, 'Traite/ 1. c, 2nd
edit., p. 211, and ' Synops./ p. 95. — Dickinson, " Case of Epi-
lepsy in Children relieved by the expulsion of Worms," ' Med.
Times and Gaz./ Jan., 1863. — Dickson, B., art. "Anthelmintics,"
rep. from the ' Penny Cyclopaedia/ in Knight's ' Eng. Cyclop.
Arts and Sci. Div./ vol. i (column 365), London, 1859. —
Dreyfus, " Irritation of the Bladder from Ascarides," from
'Journ. de Med./ in ' Lond. Med. Gaz./ 1847. — Elliotson, J.,
"A Lecture on Worms," ' Lond. Med. Gaz./ 1833.— Idem, "On
Worms in the Intestinal Canal," ' Lancet,' 1831. — Idem, " On
a Case of Threadworms," ' Lancet/ 1831. — Idem, " On Intes-
tinal Worms," ' Lancet/ 1830. — Heller. A., " Darmschmarotzer,"
in von Ziemssen's ' Handbuch/ Bd. vii, s. 632 (see also Anglo-
American edit.), 1876 — Kiichenmeister, 1. c, Eng. edit., p. 356.
— Ransom, in ' Reynolds' ' Dictionary of Medicine.' — -Smith, A.
(and others), ' Lancet/ April 29th, 1865, p. 468.— Strieker, W.,
in 'Virchow's Arehiv/ xxi, 1861, s. 360. — Tatham, 'Lancet/
April, 1867, p. 457 ; see also p. 519. — Vix, E., ' Ueber Ento-
zoen/ &c, Berlin, 1860 ; see also " On the occurrence of En-
tozoa in the Insane, particularly with respect to the Oxyuris
vermicularis /' brief notice (' Allg. Zeitsch. f . Psychiatrie ')
in Winslow's ' Journ. of Psycholog. Med./ vol. i, 2nd series,

1861, p. 158. — Zenker, ' Verhandl. d. phys. med. Soc./ H. ii,

231.

PARASITES OP MAN

Erlangen, 1870, s. 20; and in ' Tageblatt der deutschen
Naturforscherversaumilung zu Dresden/ 1868, s. 140 (also
quoted freely by Leuckart, Davaine, and Heller).

Leptodera (Anguilhda) stercoralis, Bavay. — In the summer of
1876 Dr Normand, of the French Marine, discovered this little
entozoon in the fsecal discharges of soldiers who had been sent
home invalided from Cochin-China. The patients in question
were the victims of the so-called Cochin-China diarrhoea or
dysentery. This disorder is endemic in character, and it had
hitherto been regarded as consequent upon a variety of causes
other than parasitic. Dr Normand's discovery, as such, there-
fore takes equal rank with the analogous revelations made by
Bilharz, Harley, Leuckart, Zenker, Weber, Lewis, and Bancroft,
in respect of the particular helminthiases in man with which
their names are severally associated (Bilharzia disease, Endemic
haematuria, Cestode tuberculosis, Olulaniasis, Inter-tropical
anasmia, Trichinosis, Lymphoid affections, Helminthoma, and so
forth), and also, if I may be permitted to say so, with my own
determinations in respect of a variety of endemics affecting
animals (cestode and nematode epizooty in the horse, the so-
called grouse-disease, the pigeon-endemic due to lumbricoids,
&c).

The Leptodera stercoralis is a minute, smooth-bodied, simple,
rhabditiform nematode, measuring when full grown ±" in
length, with an average breadth of ^ of an inch. The
embryos at the time of their extrusion measure only in
length, but by the time at which a rudimentary vesicle repre-
senting the uterus begins to form, the females have already
attained a length of about . The males and females are of
nearly equal size. The transition from the embryonal state to
the higher larval conditions is accompanied by a change of
skin, after which the digestive and reproductive organs are
gradually but rapidly formed and completed. These changes
have been minutely traced and recorded by Professor Bavay,
who also compares the entozoon with the genera Bhabditis and
Leptodera, in either of which genera the worm might be placed.
I have accordingly adopted the nomenclature suggested by
Bavay.

As happens in all the kindred helminthiases that are known
to be dependent upon the presence of small worms, large
numbers of Anguillules are necessary to produce injurious
effects upon the bearer. Thus, the evacuations of the Cochin-

NEMATODA

235

China patients were found to contain such multitudes of the
worms that their numbers could only be adequately estimated
at so many hundreds of thousands passed in twenty-four hours.
Of course they varied in quantity, not only in different patients,

i'io. 48. — Leptodcra inteslinalis. a, Adult female, and separate figure showing a portion of
the body with the ova in sitd. The two outlined figures represent profile and front views
of the tail, respectively, b, c, Eggs with imperfectly formed embryos, d, Larva. HiWilv
magnified. After Bavay. ° J

but in the same bearer, from day to day. They are to be found
in every stage of growth and development, from that of the
intra-ovular embryo and free embryonic state up to sexual
maturity. They occupy all parts of the intestinal canal, from

236

PARASITES OF MAN

the stomach downwards, being also found in the pancreatic and
biliary ducts, and likewise within the gall-bladder. According
to Bavay, five days suffice under favorable circumstances for the
complete maturation of the worm. This readily accounts for
their occasional extreme abundance.

I am indebted to the courtesy of Dr le Roy de Mericourt for
the original memoirs from which these brief abstracts are taken.

Leptodera intestinalis, Bavay. — This is a larger species, now
and then found associated with the above, and, according to
Bavay, " in infinitely less abundance." This species was also
discovered by Dr Normand, and has been carefully described
by Bavay. Possibly the worm may afford us another curious
instance of dimorphism. Be that as it may, it must be pro-
visionally regarded as a distinct form. As its occurrence is by
no means invariable, its role in relation to the Cochin-Ohina
diarrhoea must, as Davaine has likewise remarked, be regarded
as of secondary importance. It is readily distinguished from
A. stercoralis both in the adult and larval conditions. The full
grown worm, although comparatively narrow, is more than
twice as long as its congener; moreover, the larvee, in place of
possessing finely-pointed tails, have blunt or truncated caudal
extremities. Converting M. Bavay's millimetric measurements
into fractions of the English inch, the average length of the
mature worms will be about whilst their breadth does not
exceed in diameter.

Bibliography (No. 32). — Bavay, " Sur UAnguillule ster-
corale," ' Comptes Rendus/ Oct., 1876, p. 694, also in fAnn.
Nat. Hist./ vol. xviii, 4th series, p. 507, 1876, also noticed in
the ' Veterinarian/ Jan., 1877, p. 19. — Idem, " Note sur
VAnguille intestinale," ' Archiv. de Med. Nav./ July, 1877, p.
64, and in ' Ann. Nat. Hist./ 1877, vol. xix, 4th series, p. 350.
—Cobbold, T. 8., " Parasites of Man," in the ' Midland Natu-
ralist ' for January 1st, 1879. — Davaine, ' Traite/ 1. c, 2nd
edit., Supp., pp. 966-976, 1877. — Laveran, in ' Gaz. Hebd. de
Med./ Jan., 1877, p. 42. — Layet and Le Boy de Mericourt, in
' Diet. Encycl. des Sci. Med./ 1875. — Libermann, in ' Gaz. des
Hop./ March, 1877, p. 237, and in ' La France Med,/ 1877, p.
165 (quoted by Davaine). — Mericourt (see Layet). — Norman d,
A., in ' Comptes Rendus ' for July, 1876, p. 316, and Aug.,
1876, p. 386.— Idem, in ' Arch, de Med. Naval e/ 1877, p. 35,
and separately as 'Memoire sur la diarrhee dite de Cochin-
chine/ Paris, 1877. — Idem, " Du role etiologique de FAnguil-

NEMATODA

237

lule dans la diarrhSe de Cochinchine/' in ' Archives de Mede-
cine Navale' for September, 1878, pp. 214-224.

Ascaris mystax, Rudolphi .— This well-known helminth pos-
sesses aliform appendages, one on either side of the head. It
is of a medium size, the male measuring 2±" and the female
usually 3^" to 4" in length. Both as regards the size of the
alee and the length of the body it varies in different hosts.
Thus the variety infesting the dog has long been regarded as a
distinct species {A. marginata), partly from the circumstance
that the alee are less conspicuous, and partly because the indi-
viduals are often longer and thicker. I possess one specimen
from the dog measuring more than six inches in length. From
like causes the Ascaris leptoptera and other varieties infesting
the carnivora have been regarded as distinct species, but the
worm also varies in one and the same host.

As remarked in my elementary treatise, the late Dr
Bellingham, of St Vincent's Hospital, Dublin, published in the
13th vol. of the 'Annals of Natural History/ an extended
catalogue of Irish entozoa, and in this list he recorded the
existence of a new round worm in man. He says of it : — "From
the distinctness of the lateral membranes of the head I have
given it the name of Ascaris alata." The catalogue was con-
stantly referred to by Dujardin, Diesing, and other systematists ;
but some of the continental helminthologists do not appear to
have had access to Dr Bellingham's more extended account of
this parasite as given in the first volume of the ' Dublin Medical
Press/ No. 7, Feb. 20th, 1839. I am led to this inference from
the doubt which some have cast upon the very existence of the
worm, although others, with more candour, supposed that Belling-
ham had only mistaken the species. Thus, Kiichenmeister ('Para-
siten/ s. 464, and in Lancaster's edit., vol. ii, p. 100) says : —
" The Ascaris alata, found in the small intestines of a man, is
probably only a young individual of one of the long-known
nematoda, if, indeed, it be a worm at all !" (The italics are
mine.) This statement was reproduced by Hulme in his
English edition of Moquin-Tandon's ' Elements of Medical
Zoology/ p. 341 ; and the French author himself evidently
shared the doubt expressed by other people. Dujardin (' Hel-
minthes/ p. 156) admitted the species, as also did Diesing
(' Systema Helminthum/ p. 175), but the latter unluckily added
the following very significant suggestion : — " An Ascaris lum-
bricoides capitis epidermide emphysematice inflata V

238

PARASITES OF MAN

Dr Leidy, of Philadelphia, admitted A . alata among his Entozoa
hominis without comment (' Smithsonian Contrib.' for April,
1853), but Weinland, of Frankfort, in his list, prefixed a note
of interrogation, observing also that it had been " once " found
in Ireland ('Essay on Tapeworms/ p. 88). It is quite clear,
therefore, that these authors did not believe that the Ascaris
mystax was a human parasite. Those who doubtfully accepted
Bellingham's A. alata did so under the impression that what-
ever it was, it could not be regarded as the common Ascaris of
the cat. In the new edition of Davaine's 1 Traite/ A. alata is,
to my surprise, still retained as a separate species, and there is
no mention of the occurrence of A. mystax in man. From what
has recently been written by several continental helminthologists
(Leuckart, Heller, and others), I rejoice to think that it is not
necessary for me again to advance the really superabounding
proofs that Bellingham's A. alata was nothing more than A.
mystax. It has at length been admitted by almost all who are
competent to form an opinion, that the memoir originally com-
municated to the c Lancet,'' in 1863, and subsequently introduced
into the text of my introductory work, finally settled the ques-
tion of identity. It was through the donation of Dr Edwin
Lankester and Mr Scattergood that I was enabled at the time
to announce the third instance of the occurrence of this parasite
in man, and since that date several other instances have been
brought under public notice. Not less than seven cases have
now been noticed in which this little lumbricoid of the cat
and dog has been found in man. For one good human specimen
I am indebted to Dr Morton. In the above list I include
Heller's specimen, and the one from Greenland sent by Steen-
strup to Leuckart. According to Hering's observations this
worm grows with remarkable rapidity. Worms obtained from
a puppy only six days old measured from ^" to g" in length. In
a twelve-day-old puppy they reached nearly an inch in length,
and in a month the growth was up to four inches. Females
only \\" in length already contained eggs, and males only §"
long had acquired their spicules. Three weeks therefore,
would be amply sufficient for the completion of sexual maturity
within the feline or canine host. We do not know, however,
whether or not a temporary host is necessary for the larvas prior
to their introduction into the cat or dog. Hering thinks that
a direct infection by the ova is sufficient j but he gives no proof
of the truth of this hypothesis. " Leuckart (as quoted by Heller,

NEMATOPA

239

1. c, s. 615) found numerous embryonal roundworms in the stomach
of a cat, g^" in length, and in addition all the intermediate stages
of growth up to the larger examples found in the small intestine.
They remain in the stomach until they have attained a length
of from re" to ^" ar|d then pass into the small intestine. When
they have attained a length of nearly they cast their skins
and change the tooth-like boring apparatus for the three cha-
racteristic semicircular lips. These observations on Ascaris
■mystax (adds Heller) render it probable that A. lumbricoides is
also introduced into the human alimentary canal while still in the
embryonal state or somewhat further advanced (und wohl auch
grosse)." The subject will be found more fully discussed in
my account of the large species further on. The cat's worm
possesses an historical interest, not only in connection with
Bellingham's original discovery, but also in respect of Nelson's
subsequent determinations as to the precise mode of impregna-
tion in nematodes. The subject is too extended and too special
to be dealt with here at any great length.

For several years after Nelson left the shores of England to
spend a too short life in New Zealand, the points discussed in
his ' Edinburgh Thesis' (and subsequently published in the e Phi-
losophical Transactions ') formed the subject-matter of numerous
memoirs contributed to the leading German scientific journals.
Stated with brevity, it may be said that, according to Nelson,
the essential act of impregnation occurs when the thimble-
shaped spermatozoa of the male penetrate the unimpregnated or
ovarian ovum. This, he maintained, could and did take place
at any part of the surface of the unfertilised ovum, since the
granular mass of which it was composed, though well defined,
did not, at this period, possess a limiting — or true yolk — mem-
brane. Professor Allen Thomson, in a series of papers (some
contributed in the German language), supported Nelson's views
generally.

Amongst Nelson's chief opponents was Meissner, who demon-
strated that the unimpregnated ova really possessed a delicate
limiting membrane, and that consequently the action of the
spermatozoa was restricted to that portion of the ovarian ovum
which became exposed by rupture or separation from the rachis.
This opening he termed the micropyle. The union of the
sexual elements is quickly followed by a condensation of the
yolk-granules, and by tho disappearance of the hitherto cen-
trally placed germinal vesicle. The ovum next assumes a dis-

Fio. 49.— Germs nnd ova of Acaris myslax. Nob. 1 to 3 magnified 330 diameters and
Nos. 4 to 21 magnified 220 diameters. After Nelson.

NEMATODA

241

tinctly oval shape, the true yolk-membrane and the external
chorional envelope now becoming more and more differentiated,
until the latter acquires a regularly tuberculated surface. Co-
ordinating with these changes the granular yolk is seen trans-
forming itself into a single large embryonal cell ; after a time
this cell divides and subdivides by the ordinary process of
yolk-segmentation, until it is finally resolved into the condition
of a short, stout, vermiform embryo. The egg having assumed
its definitive oval shape, the intrachorional embryo remains
coiled within the shell, and does not make its escape until the
egg has passed from the body of the parent worm.

Into the question of the mode of formation of the ovarian
ova, and also into that of the development of the spermatozoa,
I do not enter. However unwillingly, I must, in this matter,
be contented to refer to Professor Allen Thomson's classical
article ovum (quoted below), to Leuckart's elaborate analysis
(1. c, Bd. ii, s. 76-92), and also, especially, to the exhaustive
memoir of Claparede, whose brilliant labors, like those of Henry
Nelson, were too early terminated by death. Shortly after
graduation Nelson suffered a virtually enforced banishment from
his native land.

Blbliogkaphy (No. 33). — Belling ham, 0. B., " On the Genus
to which the Worms known as Ascarides belong," ' Dublin
Journ./ vol. xiv, 1839. — Idem, " Catalogue of Irish Entozoa,"
' Ann. of Nat. Hist./ vols, xiii and xiv, 1843-44; and in the
first part of Charles worth's ' Mag. of Nat. Hist./ vol. iv, 1840.
See also the address by Dr E. D. Mapother on the " Lives and
Writings of O'Ferrall and Bellingham/' in the ' Dubl. Journ.
of Med. Sci./ Nov., 1877, p. 471 et seq.—Bischoff, ' Widerle-
gung (u. s. w.)/ Giessen, 1853; quoted by Claparede, 1. c.
infra, p. 9. — Idem, * Bestatigung (u. s. w.)/ Giessen, 1864. —
Idem, " Ueber Ei- und Samenbildung und Befruchtung bei
Ascaris mystax," Sieb. and Koll. ' Zeitsch./ 1855, s. 377; also
in S. and K. ' Zeitsch./ 1856. — Bremser, ' Icones helminth./
p. 23, tab. iv. — Claparede, E., " Ueber Eibildung und Befruch-
tung bei den Nematoden/' S. and K. ' Zeitsch./ 1857, s. 106.
— Idem, ' De la formation et de la fecondation des ceufs chez
les vers Nematodes/ Geneve, 1859. See also ' Ann. of Nat.
Hist./ vol. i, 3rd series, 1858.— Oobbold, in ' Proceed, of the
Zoological Soc. of London/ Nov., 3 862.— Idem, 'Brit. Assoc.
Eep./ 1862. — Idem, " On the occurrence of Ascaris mystax in
the Human Body/' with figures, ' Lancet/ Jan., 1863 ; and in

242

PARASITES OF MAN

the 'Dublin Med. Press/ Feb., 1863.— Idem, ' Entozoa/ chap.
xi,p. 316, 1864.— Idem, ' Worms/ pp. 72 and 112, 1872.— Idem,
in " Obituary Notice of Dr Henry Nelson," ' Med. Times and
Gaz./ 1865 ("?).— Davaine, ' Traite/ 1. c, 1877.— Diesincj, 0. M.,
'Syst. Helm./ vol. ii, p. 180, 1850.— Dujardin (1. c, Bibl.
No. 2), p. 162. — Froelich, in ' Naturf./ xxiv, s. 141 (As c. fells).
—Funke, 0., * Lehrbuch (u. s. w.)/ 1857, s. 1299.— GmeJin,
' Syst. Nat./ p. 3031.— Gohe, ' Naturg./ 1. c, s. 79.— Gwrlt,
' Path. Anat./ s. 366. — Heller, A., " Darmschmarotzer," in
'Von Zieinssen's Handbucb/ Bd. vii, s. 361. — Idem, ' Sitzungsb.
d. Erlanger phys.-med. Soc./ 1872, s. 73. — Hering, " Ueber das
Vorkommen und die Entwicklung der Ascaris mystax bei jungen
Hunden," quoted by Leuckart from ' Wiirtemb. Naturw.
Jahresbefte/ 1873, s. 305-337.— Kolliker, in ' Mullens Archiv/
1843, s. 68 et seq. — Leidy, ' Proc. Acad. Phil./ viii, p. 50. —
Leuckart, 1. c, Bd. ii, s. 258. — Meissner, G., " Beobachtungen
iiber das Eindringen der Samenelemente in den Dotter," S.
and K. ' Zeitscb./ 1854, s. 208. — Morton, T., " Another Example
of the Occurrence of A. mystax, from a Child of fourteen
months old," in a letter to the 'Lancet/ March 11th, 1865,
p. 278. — Nelson, H., " On the Reproduction of Ascaris mystax,"
' Proc. of the Royal Soc./ in ' Philosoph. Trans./ and in ' Med.-
Chir. Rev./ 1051-52; also in 'Froriep's Tagsbericht./ 1852,
s. 205-207. — Budolphi, ' Synops./ p. 42, 1819. — Schneider,
"Ueber Bewegung an dem Samenkorperchen der Nematoden,"
in 'Monatsb. d. Berliner Akad./ 1856, s. 192. — Idem, 'Mono-
graphic der Neniatoden/ Erste Abth., s. 38, und Dritte Abth.,
s. 263 (" Entwicklungsgeschichte"), 1866. — Siebold, 'Ver-
gleichende Anatomie/ 1848, s. 153, and in Burnett's edit.,
p. 125 et seq., 1854. — Thomson, A., art. " Ovum," in c Todd's
Cyclop, of Anat. and Phys./ supp., 1859. — Idem, " Ueber die
Samenkorperchen, die Eier und die Befruchtung der Ascaris
mystax," S. and K. ' Zeitsch./ 1856, s. 425. — Idem, "Report
of Glasgow Meeting" ('Brit. Assoc. Rep/), 1855, p. 158.

Ascaris maritima, Leuckart. — This is a well-marked species.
Judging from the characters presented by the solitary, sexually-
immature female which supplied Leuckart with his only means
of diagnosis, this worm may be briefly described as a filariform
nematode about §" in length and about in breadth. Although
there are no cephalic aliform membranes, the cuticle imme-
diately below the lips forms small and distinct projections,

NEMATODA

243

one on either side of the head (' Die Mensch. Par./ Bd. ii,
s. 877).

This entozoon was discovered by Dr Pfaff at Jacobshavn, near
Godhavn, West Greenland, in April, 1865. Two years later
he sent the specimen to Krabbe, who afterwards transmitted it
to Leuckart. In the original communication addressed to the
Copenhagen helminthologist, Dr Pfaff states that he procured
the worm from amongst matters vomited by a child, and he
incidentally observes that he had hitherto encountered only
Bothriocephalus cordatus and Oxyuris vermicularis amongst
Greenlanders. As to the source of infection, Prof. Leuckart
not unnaturally refers to the similar conditions of existence
shared by the human and carnivorous inhabitants of that
country. It is well known that bears, polar-bears, seals, and
walruses are largely infested by nematodes (Asc. transfuga,
A. osculata, Ophiostoma dispar, &c), but these various species
are quite distinct from Dr Pfaff's little " spulwurm."

Ascaris lumbricoides, Linneus. — This common parasite was
for a long while regarded as identical with the great lumbri-
coid of the horse, but the question has been finally settled by
Schneider, who has shown that the human worm, although
identical with Dujardin's Ascaris suilla of the hog, is never-
theless quite distinct from the Ascaris megalocephala of solipeds.
The large lumbricoid occasionally found in the ox belongs to
the human worm. Our large human helminth resembles the
common earth-worm in general appearance only. The males
usually measure from four to six inches in length, and the
females from ten to fourteen inches. Some have been reported
up to seventeen or eighteen inches in length. The body is
smooth, fusiform, and elastic, and marked by numerous fine
transverse rings. It is attenuated towards either extremity, the
anterior end terminating in a prominently three-lobed mouth,
The tail is bluntly pointed. The female is much shorter than
the male, having a diameter of nearly a quarter of an inch. The
male is supplied with a double spiculum, its tail being always
more or less curved towards the central surface. The female
reproductive orifice is situated above the centre of the body.
According to Schneider, the tail supports from 138 to 150
caudal papillae, that is, from 69 to 75 on either side of the
median line. Below the anus the papillae are regularly
arranged in pairs, seven in number, the two uppermost pairs
being double.

244

PARASITES OF MAN

Notwithstanding the advantage which the size of this entozoon
affords us in the matter of observation and experiment, we are yet
ignorant as to the precise mode in which the young gain access to
the human body. From what has been said respecting the quick
growth of Ascaris mystax in the dog, and from what has been
observed respecting the rapid growth of the so-called A. suilla
in the hog, we know that the worm requires but a short time to
pass from the larval to the sexual state. The view of Hering,
Mosler, Davaine, and others, who suppose that these worms are
reared in a direct manner by swallowing the ova, is, as Leuckart
observed, not yet proved. We are not in full possession of
the facts of larval development. It is true that Professor
Heller's interesting " find " has shown that when these worms
first gain access to the human body their size is quite insigni-
ficant. At the post mortem of an imbecile, Heller discovered
eighteen young worms, varying in size from about §" to |" in
length (2 "75 to 13 mm.). The sexes were indistinguishable.
As a set-off against this, Leuckart's repeated attempts to rear
Ascaris lumbricoides and A. mystax by means of direct feeding-
experiments with the eggs all failed. Thus, we are yet left in
doubt as to the destiny of the larvae during the period which
elapses between the time of their escape from the egg and the
time of their entry into the human body. So important is the
question as to the mode of origination, growth, and subsequent
development of the larvae, that it may be well to trace, however
briefly, what steps have been taken to clear up the matter.
Leuckart obtained his negative results by the administration of
ripe ova to dogs, rabbits, swine, and mice. The eggs of
Ascaris lumbricoides have been kept alive by Dr Davaine for a
period of more than five years. I have myself watched the
development of their contents in fresh water through all the
stages of yolk- segmentation up to the stage of an imperfectly-
organised, coiled, intra- chorional embryo, and have kept them
in the latter condition for a period of three months. Accord-
ing to Davaine (' Comptes Rendus/ 1858, p. 1217), the fully-
developed embryo is cylindrical, its length being rJ0th of an
inch. The mouth is not furnished with the three characteristic
papillae of the genus, and the tail terminates suddenly in a
point. Davaine administered some of his five-year-old embryos
to rats, and had the satisfaction of finding a few of these eggs
in the rodent's ftcces, with their embryos still liviog, but
striving to emerge. He also gave eggs to a cow, and intro-

NEMATODA

245

duced others into the stomachs of dogs in small linen-covered
flasks. As a general result it may be said that the embryos
escaped from their shells. Those eggs, however, in which the
yolk- segmentation had not arrived at the early embryonal stage
remained unaffected. According to Heller, the embryo of
A. lumbricoides casts its first skin while still within the egg,
and " a subsequent ecdysis probably completes its definitive
form" (1. c, s. 615). So far back as 1853 Yerloren reared
coiled intra- chorional embryos in the eggs of Ascaris marginata
within a period of fifteen days in distilled water. I also reared
the embryos of this species in fresh water, and kept them alive
for a period of nearly a year and a half, at the expiration of
which time, and during the warm weather, some few of them
succeeded in making their escape. According to Davaine, the
eggs of many nematode species will readily retain their vitality
though long exposed to dryness, but their yolk-contents will
not go on developing during this period of exposure. As
regards A. mystax, however, Heller remarks that whilst ee the
eggs have a great power of resisting external influences, their
development is not arrested in spirits of wine, chromic acid, or
oil of turpentine" (1. c, s. 631). In the case of Ascaris
tetraptera of the mouse, embryonic formation goes on in spite
of the absence of external moisture. Davaine has noticed the
same thing in the oxyurides of rodents. Dryness does not
even destroy the eggs of A. lumbricoides and Tricocephalus
dispar. It would seem, in short, that the eggs of nematodes
which normally take up their residence in cats, dogs, and in
the carnivora which reside m arid regions, will develop
embryos in the egg without external moisture. As before
remarked, Davaine thinks it is not necessary that these nema-
tode embryos should pass through any intermediary bearer, and
he believes that they are often directly transferred to the
stomach of their " hosts " whilst adhering in the form of an
impalpable dust to the coats of their bearers, whence they are
detached by the animal's frequent habit of licking the fur.
Davaine' s view has received some support from the observations
and experiments of Unterberger with the eggs of Ascaris
maculosa. This observer administered eggs of the worm to
doves (whose faeces were free of eggs), and seventeen days
after found ova in the faeces.

With the eggs of the Ascaris megalocephala of the horse I
performed numerous experiments. I reared the embryos in

246

PARASITES OF MAN

simple fresh water, and found them during warm weather
escaping before the expiration of five months. I also succeeded
in rearing these larvae in pond mud, noticing, at the same
time, that after their escape from the shell they grew more or
less rapidly up to a certain point, after which they ceased
growing. The addition of horses' dung to soft wet mud in one
case, and of cows' dung in another, neither appeared to advance
nor retard the process of embryonal formation, so long as the
embryos were enclosed in their shells. On the other hand,
when I reared embryos in simple horse-dung purposely kept
moist, they attained a higher degree of organisation than did
those in wet mud or water. Having watched hundreds of
these larvae under varying conditions, I came to the conclusion
that, after escape from the egg, their activity, growth, and
strength was most marked when they occupied media which
happened to be impure. Davaine experimented on cows, and
Leuckart also experimented on horses, with the eggs of this
worm without success. Leuckart also failed to rear the larvae
in intermediary hosts. Some eggs passed through the water-
palmer unaltered.

These results, so far as they go, seem to be borne out by
facts of a professional order. Thus, an instance has been
brought under my notice where a considerable number of
peasants and their children, dwelling in a parish in Yorkshire,
were infested with this worm. There was, in short, a local
endemic helminthiasis. Through the parish runs a stream
which supplies the cottagers with all the water they employ for
domestic purposes (washing, drinking, and so forth). Some of
the peasants living by the side of the stream keep pigs, and
the sewage from this source has been allowed to pass into the
stream itself. Now, if Schneider's determination as to the
identity of the lumbricoid of man and the pig is correct (which
I do not doubt), the explanation of the cause of the endemic
becomes a very simple matter. But it does not explain all
that we desire to know about the young worms. Either the
freed embryos before they enter the human bearer accomplish
further changes of form and growth in the sewage or impure
water ; or, what is far less probable, they pass into the bodies
of intermediary hosts (such as insect-larvae, Gammari, Ento-
mostraca, &c.) to undergo the necessary changes. Practically,
no doubt, it comes to the same thing in the end. Even if we
suppose that the Ascaris svilla and A. lumbricoides are not

MEM ATOP A

247

identical species, still it is evident that any person discharging
the eggs of lumbricoids in the vicinity of open waters becomes,
by that fact, a source and centre of infection. To ensure an
endemic it is probably only further necessary that the human
inhabitants should employ the contaminated water for domestic
purposes. But time and an increase of temperature must be
allowed for the bringing about of those known and unknown
larval changes that alike form the necessary antecedents of
infection. In this connection I will only add, that if the
present position of the question be such as I have here repre-
sented it to be, we see that Mosler was not far wrong when he
suggested that " contamination of the drinking water with the
eggs out of privies is to be blamed" as a source of infection.
According to Heller, from whom I quote, Mosler actually
demonstrated the presence of the eggs in water thus exposed.
In like manner it becomes obvious that Davaine's practical
remark (although it was based on the assumption of a direct
infection by the eggs), that filtration will probably be sufficient
to prevent infection, loses nothing of its hygienic value.

The foregoing observations naturally lead one to the question
of frequency and distribution. Davaine holds that the com-
parative infrequency of this parasite in Paris is due to the free
use of the filter. In London, though not uncommon, the worm
rarely occurs in great numbers in one bearer. Those cases in
our hospitals, where considerable numbers have been present,
have usually come up from suburban or country places. Heller
states that these worms were found in 9*1 per cent, of post
mortems conducted at Dresden, in 12 per cent, at Brlangen,
and in 17 per cent, at Kiel. He quotes Huss as stating that
no one is free from this worm in Finland. The prevalence of
large round worms in warm countries generally is well known.
Throughout India and the Bast they are extremely abundant,
and the same may be said of the West Indies, Brazil, and the
adjacent territories. Professor Dyce and others have remarked
on the extreme prevalence of lumbrici in the Mauritius, but
they are comparatively rare along the sea border. In all
situations where there is an abundant fresh-water supply these
parasites are particularly common, as in the lowlands of
Holland and the lake districts of Sweden. The abundance of
water is certainly not alone sufficient to explain the frequency
of the parasite, seeing that the most important factor is that
which rests upon the uncivilised habits of the rural population.

218

PARASITES OF MAN

What, therefore, it may be asked, can be the cause of immunity
enjoyed by Icelanders in this respect ? The answer is not
apparent j nevertheless Krabbe and Finsen have testified to the
fact that Iceland is the only country that is entirely free from
Ascaris lumbricoides.

As remarked in my previous work the number of worms
present in any human bearer is usually small, varying commonly
from one to six or eight. Cases in which scores or hundreds
have existed are comparatively rare. Kiichenmeister mentions
the case of one child who passed 103 examples, and of another
child that harbored from 300 to 400 worms. Dr Gilli, of
Turin, gives a case where 510 were passed by a child, and
Cruveilhier estimated that over 1000 existed in an idiot girl,
whose intesiines he found crammed with them. A remarkable
case has also been communicated to me by Dr Mackeith, of Sand-
hurst, Kent, who, by means of santonine, expelled from a little
girl, five and a half years of age, 300 lumbrici; and i am like-
wise indebted to Dr Cooper Hose for notes of a case in which
about thirty lumbrici were expelled, chiefly in consequence of
the employment of this drug. The most interesting fact, how-
ever, in this case was that the child was only fifteen months old.
In this case the symptoms were severe.

The proper habitat of the lumbricus is the upper and middle
part of the small intestine. From this situation it often wanders
into the stomach, and frequently gains access to the outer world,
not only by the natural passages of the mouth, nostrils, and
anus, but also, occasionally, in a more direct way, by perforating
the intestinal and abdominal walls. Many cases are on record
where lumbrici have passed into the abdominal cavity. In other
instances they have lodged themselves within the abdominal
viscera and pulmonary organs. When they find their way into
the parietes of the abdomen and adjacent parts, they usually give
rise to the formation of abscesses requiring surgical interference.

As regards the symptoms produced by lumbrici, these vary
according to the situation they happen to occupy. The sym-
ptoms are also modified by age and temperament. In the
stomach and intestines they give rise to colic and shooting pains
about the abdomen, followed generally by dyspepsia, nasal itch-
ing, nausea, vomiting, and even diarrhoea. Occasionally death
supervenes suddenly. A singular case of this kind (the particu-
lars of which I only gathered from a local newspaper) occurred
in a boy, thirteen years of age, at the County Gaol at Hertford,

NEMATODA

249

in 1873. From Dr Evans's statement, made at the coroner's
inquest, the sole cause of death appeared to be due to pressure
on the windpipe by a worm lodged in the gullet. Sometimes
there is cerebral disturbance, attended with general restlessness
and convulsive twitchings during sleep. Thus, Dr Woodman
has recorded a serious case of convulsions arising from lumbri-
coid worms, in which, however, a cure was effected by expul-
sion of the worms. An anonymous writer in the ' Medical
Gazette ' records a case of epilepsy from this cause, whilst
another writer in the same journal (1839) mentions an instance
where two lumbrici and one tapeworm were associated in the
production of similar phenomena. But a much more striking
case is also given (anonymously) in the ' Gazette 3 for 1874 (p.
415), where a single lumbricus caused the bearer to be a lunatic
for eight years. The victim suffered from cataleptic fits, which
lasted for two or three weeks at a time. M. Petrequin, in his
' Traite Pratique,' records two cases of amaurosis in young girls
produced by lumbrici. A fatal case is recorded by Petrenz,
where 200 worms produced enteritis, and another fatal case is
given by Eoger from perforation (1848). Cases of perforation
are also given by Young, by Blair (1861), by Mondiere (1839),
by Buchner (1851), by Sheppard (1861), and by Luschka
(1854), the worms in this last-mentioned case occupying the
cavity of the pleura. Cases of severe irritation affecting the
genito- urinary organs are given by Dreyfus, Buckingham, and
others j and one or two instances are reported where these
worms have been discharged from several parts of the body
(Neilson, 1833). I may add that the third fasciculus of a work
illustrating the collection of morbid anatomy in the Army
Medical Museum at Chatham gives a case of lumbrici occupy-
ing the biliary ducts and gall-bladder. I find, moreover, two
additional cases of perforation of the small intestine, one of
which appeared in the 'London Medical Gazette' (1827) and
the other in the ' Lancet' (1836).

During the Franco-German war Dr Reginald Pierson, as he
afterwards informed me, removed a lumbricus from an abscess
formed in the abdominal parietes of a soldier. But amongst
the most curious cases (illustrating the wandering habits of these
parasites) are those severally described by Barwell (1857),
Williams, Prichard, and the Messrs Stockbridge. In Barwell's
case an Ascaris was expelled from a child who had swallowed
the brass " eye " of a lady's dress. Through the circular loop of

250

PARASITES OF MAN

this eye, used as a toy, the Ascaris had partly thrust its'body, and
becoming thus strangulated, it probably perished before it was
evacuated. In Prichard's case (1859) one or two lumbrici had
similarly trapped themselves in the eyes of buttons swallowed
by the patient, and one worm, not contented with a single
strangulation, had succeeded in passing its body, through two
buttons. In 1842 Mr T. G-. Stockbridge gave a similar case,
in which he, not inaptly, spoke of these " hooks and eyes " as
constituting a new remedy or " worm-trap " for lumbricus,
and singularly enough, a namesake (W. Stockbridge), in the
succeeding year, also recorded a like instance of the " mechanical
expulsion of worms " by metallic buttons. Again, a third
correspondent in the ' Boston Journal/ under the initials A. M.,
spoke of an open-topped thimble as constituting another new
" worm-trap," whilst he gave a case of lumbrici penetrating
" metallic suspensor buttons." There is also the case reported by
Williams, who, at a meeting of the Boston Society for Medical
Improvement, exhibited " a lumbricus with a dress-hook
attached" (1857). Lastly, another lumbricus, trapped in the
same way, may be seen in the Museum of the Royal College of
Surgeons at Edinburgh.

Owing to the presence of a peculiar irritating vapour which
is given out by these lumbricoids, particularly when fresh,
several observers have experienced curious symptoms. Thus,
Miram on two occasions, when examining A. megalocephala, was
attacked with sneezing, excessive secretion of tears, with swelling
of the puncta lacrymalia, and Huber also experienced a trouble-
some itching of the hands and neck after examining specimens
of A. lumbricoides. In like manner I have myself had watery
suffusion of the eyes (when collecting the perivisceral fluid for
Marcet's analyses : see Bibliog.), and Bastian has given a detailed
account of the serious effects which the poison produced upon
him. In Bastian' s case even spirit specimens produced irrita-
tion. The attacks of catarrh and asthma were so persistent
and severe that they lasted for six weeks at a time. So sensi-
tive was Bastian to the lumbricoid-miasm that he could not
even put on a coat that he had worn during his investigations
without experiencing fresh attacks of sneezing and other
catarrhal symptoms. The attacks became periodical, occurring
between five and six in the morning, being accompanied by
dyspnoea and a distressing spasmodic cough. Bastian, in short,
was quite a martyr in the cause of nematode anatomy.

NEMATODA

251

Bibliography (No. 34). — Abousson, L., "On the Presence of
Worms (lumbrici) in the Air-passages/' from ' Arch. Gen. de
Med./ in 'Med.-Chir. Rev./ 1836.— (Anonymous), A. M.,
"Another New Worm-trap — an open-topped Thimble in the
Nostril (also notice of metallic suspender buttons penetrated by
Lumbrici)/' ' Bost. Med. and Surg. Journ./ vol. xxvii, p. 121,
1842-43 ; see also T. G. and W. Stockbridge. — (Anon.), "Lum-
brici expelled by Bismuth," 'Bost. M. and S. Journ.' (from
f Gaz. des Hop./ ' Journ. des Connaiss. Med./ and 'Boletin del
Inst. -Med. -Valenc.'), 1859. — (Anon.), " Case of one Tapeworm
and two Lumbrici causing Epilepsy/' from ' Bull, du Midi '
and ' Gaz. Med./ 1839. — (Anon.), " Case of Perforation of
the Ileum by Ascarides," from ' Hufl. and Ossan's Journ./ in the
' Lancet/ 1836. — (Anon.), " Case of Lumbrici in the Biliary the
Ducts and Gall-bladder/' note and fig. in third fasc, illust. the
Coll. of Morb. Anat. in the Army Med. Mus. at Chatham, 1838.
— (Anon.), " A Lumbricus causing Catalepsy, with Fits lasting
two or three weeks ; Cure by Vomiting," ' Lond. Med. Gaz./
1847, p. 415. — Archer, IE., "On a Case of A. lumbricoides pro-
ducing alarming symptoms," ' Lancet/ 1857. — Barwell, " Case
of Ascaris expelled by the swallowing of a foreign body,"
' Lancet/ 1857. — Bastian, H. C, " On the Anatomy and Physi-
ology of the Nematoids, Parasitic and Free," ' Phil. Trans.,'
1866, p. 545 : for the account of his poison-symptoms, see foot-
note, p. 583. — Batterbury, B. L., " Jaundice due to the presence
of Lumbrici," ' British Med. Journ./ Nov., 1878, p. 721.—
Bigelow, H., " Worm in an Abscess," ' Bost. Med. and Surg.
Journ./ vol. xxxiii, p. 486, 1836. — Blatchley, G. 0., "Two Cases
of A. lumbricoides, attended with Abscesses, followed by large
purulent discharges, and Worms therein," ' New York Med.
and Phys. Journ./ vol. i, new series, p. 209, 1829. — Bonfils, E.,
" Lesions and Path. Phenomena caused by Lumbrici in the
Biliary Ducts," from 'Arch. Gen./ in 'Brit, and For. Med.-
Chir. Kev./ 1858, and in e Amer. Journ. of Med. Sci./ vol.
xxxvii, 1859. — Bradford, J. T., " Singular Case of Worms
(Lumbrici)," 'Bost. Med. and Surg. Journ./ vol. xxviii, 1843.
Brigham, A., " Worms in the Bladder simulating ' Stone/ "
'Amer. Journ. Med. Sci./ 1837; ' Med.-Chir. Kev./ 1837;
' Quart. Journ. Calcutta Med. and Phys. Soc./ vol. ii, p. 132
1 838.— Buchner, " On the Perforation of the Intestinal Canal by
Worms (with ref. to two cases)," from 'Med. Zeitung/ 1850
in 'Med.-Chir. Rev./ 1851.— Calderwood, "Treatment," 'Brit!

252

PARASITES OF MAN

Med. Journ./ Jan. 30, 1875.— Chapman, N., " Case of 68
Ascarides causing Pulmonary Disease," in his ' Dis. of the
Thoracic and Abd. Viscera/ p. 263, and in ' Med.-Chir. Rev.,'

1845. — Chiaje, Belle-, in ' Rend, dell' Accad. di Napoli/ 1846
("Anat./' p. 403). — Church, J., " On A. lumbricoides," 'Mem.
Med. Soc. Lond./ vol. ii, 1789.— Claparede (1. c, Bibl. No. 33,
for development). — Clark, P., " Discharge of a Lumbricus
through the Male Urethra/' 1 New York Journ. Med./ 1844,
rep. in ' Lancet/ 1844, and in ' Edin. M. and S. Journ./ vol.
lxiv, 1845. — Cloquet, ' Anat. des vers Intest./ 1824. — Cobbold,
" On Sewage and Parasites, especially in relation to the
Dispersion and Vitality of the Germs of Entozoa," ' Med. Times
and Gaz./ Feb. 25, 1871, p. 215.— Idem, ' Entoz./ p. 302-315.
— Idem, ' Worms/ lect. xvi, p. 3. — Idem, art. " Ascaridae," in
' Maunder' s Treasury/ 1862. — Cohan, J., " Case in which Eleven
Round Worms of the species A. lumbricoides were removed
by Anthelmintics/' ' Dubl. Med. Press/ vol. xxvi, p. 211, 1851. —
Cutler, J. H., " Death by Worms (a large Lumbricus being
found in the Wind-pipe)/' ( Bost. Med. and Surg. Journ./
vol. lxvi, p. 392, 1862. — Czermah, in ' Sitz. d. k. Akad. d.
Wissensch./ 1852 ("Anat.," s. 1bb).—Davaine, in his ' Traite/
1. c.j 2nd edit., syn. xcvii, and p. 122-235 (with details of forty-
five cases) ; see also his memoir " On the Development and
Propagation of the Trichoceph. dispar and A. lumbricoides/'
from ' Comptes Rendus/ in f Ann. Nat. Hist./ vol. ii, 3rd series,
1858 ; also in the ' Journ. of Pract. Med. and Surg./ Eng. edit.,
vol. i, 1858, and in the 'Veterinarian/ vol. xxxii, p. 700, 1859,
from ' Proc. of Acad, des Sci./ in ' Bost. M. and S. Journ./
vol. lix, p. 157, 1858-59. — Idem, art. " Entozoaires," in ' Diet,
de Med. et Chir. prat.' — David, J. B., " Cases of Perforation of
the Intestines by Worms/' from ' Gaz. Med. de Paris/ in ' Dubl.
Med. Press/ 1840, p. 223—Biesing, < Syst. Helm./ ii, p. 166;
and in ' Revis der Nemat./ 1. c, s. 660. — Douglas, J., "Worms
(Lumbrici) evacuated at an Ulcer of the Groin," fMed. Ess.
and Obs./ vol. i, 2nd edit. (vol. i, 5th edit., p. 179), p. 222,
1 737. — Dowler, B., " Case of Worms in the Urinary Bladder,"
from ' New Orl. M. and S. Journ./ in f New York Journ. Med./
new series, vol. xiv, 1855. — Dubini, ' Entozoografia umana'
("Anat.," p. 148). — Dupuytren, "Lumbricus passed by the
Urethra," from "Clin. Lect.," in ' Lond. Med. and Surg. Journ./

1846, p. 14. — Dyce, B., "On Lumbrici and the Causes of their
Prevalence in the Mauritius," • Lond. Med. Gaz./ 1834. — Evans,

NEMATODE

253

T., "Lumbricus causing Death/' rep. of coroner's inquest in
the ' Herts Advertiser and St Alban's Times' for Feb. 8, 1873.
Gcrvais (and Van Beneden), ' Zool. Med./ ii, p. 118.— Gilli,
"Account of a Case in which 510 Worms (Lumbrici) were
voided by a Child/' from ' Giorn. d. Scienze Med. di Torino/
in ' Med.-Chir. Eev./ 1843.— Goojpta, G. D. D., " On Suicide
and Lumbrici/' ' Ind. Med. Gaz./ July, 1874, and ' Lond. Med.
Eec./ Aug., 1874, p. 502. — Heller, A., " Darmschmarotzer," in
'Von Ziemssen's Handb./ s. 612-631.— Holland, G. C, 'A
peculiar Case of Nervous Disease or Derangement of the Nervous
System (associated with A. lumbricoides) ," ' Edin. M. and S.
Journ./ vol. lxiii, 1845. — Howall, "Abscess of the Groin, with
discharge of Lumbrici," ' Lond. Med. Gaz./ 1845, and ' Edin.
M. and S. Journ./ 1846, p. 241. — Johnson, W. G., "Case of
forty Lumbrici in a Boy who died with Traumatic Tetanus,"
" Eep. of South Mid. Br. of Brit. Med. Assoc.," in ' Brit. Med.
Journ./ 1858. — Kell, " Perforation of the Intestines by a
Worm," 'Lond. Med. Gaz./ 1828.— Kilgour, T., "Case in
which Worms in the Nose, productive of alarming Symptoms,
were removed by the Use of Tobacco," ' Med. Comment./
vol. viii, 1783. — Kirhland, " Case of Lumbricus in an Abscess
of the Liver," rep. in his book, entitled 'An Enquiry/ vol. ii,
p. 186 (quoted by Richter and Davaine), London, 1786. —
Kiichenmeister, 'Manual/ Eng. edit., p. 410-427. — Leidy, J.,
'Proc. Acad. Phil./ 1856, p. 50. — Lente, F. _D., " Lumbricus in
the Stomach causing Dyspnoea," in his " Rep. of Cases occurring
in the New York Hosp.," in ' New York Journ. of Med./ vol. v,
new series, p. 167, 1850. — Lettsom, 1 Case of Lumbricus evacu-
ated from an Abdominal Abscess," ' Trans. Med. Soc. Lond.,'
and 'Lond. Med. Repos./ 1817. — Leuchart, 1. c, s. 152-258. —
Lieberliuhn,\u ' Miller's Arch.' ("Anat. of A.suilla"), 1855, s. 331.
— Luschha, " Case of Lumbrici within the Pleura," from ' Virch.
Arch./ in 'Med.-Chir. Rev./ 1854.— Lowne, B. T., "The
Anatomy of the Round Worm," ' Trans. Roy. Micr. Soc./ 1871,
p. 55. — Maesson, " On a Worm found in the Bubo of a Woman
36 years of age /' see ' Entozoa and Parasites, being a ref.
to numerous papers / from " Valentin's Repertorium," in
' Month. Journ. of Med. Sci./ vol. ii, p. 559, 1842 ; also in
' Micr. Journ. and Struct. Rec./ p. 85, 1842. — Marcet, W.,
" Chemical Exam, of the Fluid from the Peritoneal Cavity of

A. mecjalocephala," 'Proc. Roy. Soc./ 1862, No. 72, p. 69.

Martin, D. T., "Large number of Worms (140 examples of

254

rAUASITES OF MAN

A. lumbricoides) discharged from a Child five years old," rep.
from the " Stethoscope/-' in ' Bost. Med. and Surg. Journ./
vol. xliv, p. 301, 1851. — Mattel, B., " On a Case of two Lurnbri-
coid Worms, which had penetrated during life into the liver,
and were demonstrated by Prof. G. Pelizzari to his pupils in
the Sch. of Path. Anat. of Florence," from ' Gaz. Mud. Ital.
Toscana/ in ' Dubl. Quart. Journ./ vol. xxiv, 1857. — Michel,
" Case of Epilepsy in a Girl ten years of age, caused by Lum-
brici," from ' Journ. des Connaiss. Med./ in ' Amer. Journ. of
Med. Sci./ vol. vi, p. 451, 1843. — M'Laggan, J., " Gangrenous
Sore from a large Worm in the Parietes of the Abdomen,"
' Med. Comment./ vol. ii, 1774, p. 80. — Molin, in ' Sitzungsb.
d. k. Akad./ 1859, s. 23. — Mondiere, " On Perforation by Worms
(three cases)," from ' I/Experience/ in ' Med.-Chir. Eev./ 1839.
— Moore, E. D., " Example of A. lumbr. ejected by the Mouth,"
'Prov. Med. and Surg. Journ./ 1852. — Morgan, J., "Case of
Perforation of the Stomach, probably by a (Lumbricus) Worm,"
'Lancet/ 1836. — Mori and, W. W., "Ejection of numerous
Lumbrici from the Mouth, impaction of the small intestine
with Lumbrici, of which 365 were removed post mortem)," ' Bost.
M. and S. Journ./ vol. lvi, 1857. — Idem, an "A. lumbr. of unusual
size (over 17 inches long)," c Eep. of Bost. Soc. for Impi'ov./
in ( Bost. M. and S. Journ./ vol. lviii, p. 62, 1858. — Neilson,
" Discharge of Worms from various parts of the Body," 1 Med.-
Chir. Eev./ and ' Lond. Med. Gaz./ 1833— Omond, B., " Case
of Lumbricus attended with Haemoptysis," ' Edin. Med. Journ./
1856. — Owen, B., art. " Entozoa," 1. c. — Padley, G., "Jaundice
and Lumbrici," f British Med. Journ./ Dec. 14, 1878, p. 877.—
Petrenz, " Case of fatal Enteritis produced by (200) Lumbrici,"
from ' Clarus and Eadius' Beitr. zur Pract. Heilk./ in ' Dubl.
Journ./ vol. xi, 1837 ; also in c Lond. Med. Gaz./ 1837.— Play-
fair, " Case of Lumbricus (69 specimens) cured by the Mudar,"
< Calc. Med. and Phys. Soc. Trans.,' vol. ii, p. 407, 1826.—
Pomeroy, C. O., "Escape of Worms (17 Lumbrici) from the
Navel of a Child," ' Bost. M. and S. Journ./ vol. xxi, 1840 —
Prichard, A., " Case of Lumbricus," ' Eep. of East York and
North Lincoln Br. of Brit. Med. Assoc./ in f Brit. Med. Journ./
1859. — Boyer, " Case of Intestinal Perforation by a Lumbricus,"
report in ' Lancet/ 1856. — Bumsey, N., " Cases of Lumbricus
and Teenia associated with Haemoptysis," ' Med.-Chir. Trans./
1818. — Sandwith, E., " Eemarks on Worms in the Peritoneal
Cavity, with a case," ' Brit. Med. Journ./ 1861.— ScMdfer,

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255

" Case of a Deaf and Dumb Child restored after the discharge
of Worms (87 Lumbrici and innumerable Oxyurides)," from
' CEsterr. Med. Wochensch ,' in ' Amer. Journ. of Med. Sci./
vol. viii, p. 473, 1844. — Schneider, A., ' Monog der Nemat./
s> 36. — Idem, " On the Nervous System of Nematoda," from
the German by Bush, 'Quart. Journ. Micr. Sci./ 1863. —
Schultze, " Case of Stuttering occasioned by Worms/' from
'Med. Zeit./ in 'Med.-Chir. Eev./ 1837 .—Sheppard, "Case of
A. lumbr. extracted from an Abdominal Abscess/' ' Brit. Med.
Journ./ 1861. — Smith, J. N., " Thirty-nine Specimens of Ascaris
lumbricoides in a Child/' ' Bost. M. and S. Journ./ 1856. —
Spalding, P., " Case of Worms (100 Lumbrici), ibid., 1839. —
Stockbridge, T. 0, "Worm- trap (hooks and eyes), a New-
Remedy," ibid., vol. xxvii, p. 73, 1842-43 ; see also Anon.,
A. M. — Stockbridge, W., "Mechanical Expulsion of Worms (by
metallic buttons)," ibid., vol. xxviii, p. 419, 1843. — Van Beneden,
' Animal Parasites/ 1. c, p. 95 ; see also Gervais. — Villemin,
" Case of Death from Worms (about 18 specimens of A. lumbr.),
from ' L'Union Med.,' in the ' Lancet,' and rep. in ' Dubl. Med.
Press/ vol. xxxv, p. 327, 1856. — Weinland, in his 'Essay/ 1. c,
p. 88, and in ' Troschel's Arch./ 1859, s. 283 — Welsh, T.,
" Curious Facts respecting (symptoms produced by) Worms
(A. lumbricoides), " art. ix in the ' Med. Papers communicated
to the Massachus. Med. Soc./ vol. i, p. 87, 1790. — Wendelboe,
" Case of discharge of Worms (Ascarides ?) through the Skin,"
from ' Rep. of Roy. Soc. Copenhagen/ in ( Lancet/ 1836 ; see
also Neilson. — Williams, H. W., " Exhibition of a Lumbricus
with a dress-hook attached," 'Rep. of Bost. Soc. for Med.
Improv./ in 'Bost. M. and S. Journ./ vol. lvi, p. 163, 1857.—
Wilson, J., " On the prevalence of Lumbrici in China," in his
'Med. Notes on China/ London, 1846. — Woodman, W. B.,
" Case of Convulsions, &c, arising from Lumbricoid Worms,"
'Med. Times and Gaz./ 1863.— Young, W., "Cases in which
Lumbrici were evacuated by Ulceration through the Parietes of
the Abdomen," 'Lond. Med. Gaz./ from * Glasgow Med. Journ./
1828; rep. in 'Lond. Med. and Surg. Journ./ vol. i, p. 564,
1828.

25G

PARASITES OF MAN

SECTION IY.— Part I. Acanthocephala (Thornheaded

worms).

Echinorhynchus gigas, Goeze. — There is but one recorded
instance of the occurrence of this entozoon in the human body.
This is the oft-quoted case by Lambl, given in the ' Prager
Vierteljahrschrift' for 1859. Lambl, indeed, described it as a
separate species (E. hominis), but as the worm was a sexually-
immature female, its identification with E. gigas, notwith-
standing Schneider's great authority, can hardly be regarded
as absolutely certain. The worm was found in the small
intestine of a boy of nine years, and measured only rather more
than the fifth of an inch in length. As Leuckart hints, the
worm may be Echinorhynchus angustatus, or possibly the E.
spirula, a species found in various South American apes (Gebus
and Jacchus), and also in the Barbary ape (Inuus).

In 1872, Welch, unaware of Lanibl's case, announced the
discovery of " the presence of an encysted Echinorhynchus in
man." The minute parasite found by him occurred in a
soldier, thirty-four years of age, who died at Netley, but who had
contracted the worm in India. " It was situated in the
jejunum, immediately beneath the mucous coat, and formed an
oval prominence in the interior of the gut." Speaking with
great confidence, this able microscopist further remarks : —
" The character and arrangement of the hooklets unequivocally
shadowed forth a species of Echinorhynchus for the first time
discovered as a representative of the Acanthocephala in the
human body." Along with his elaborate description Welch
gives several figures ; but these, so far from producing con-
viction as to the accuracy of his inferences, have unfortunately
led me to believe that the parasite in question would be more
properly referred to the Pentastomidce. But for Heller's
acquiescence I might have more fittingly noticed this worm
elsewhere. Davaine falls into the same view, and moreover
accepts Lewis's " Echinorhynqne du Chicn," which I have
shown to be a nematode (0 heir acanthus robustus). It is thus
that serious errors creep into the literature of parasitism.

In the adult state the female Echinorhynchus gigas is a
huge species, occasionally reaching two feet in length, with a
breadth of one third of an inch. The male rarely exceeds
three iuchcs. This worm is common iu swine, both wild and

AOANTHOOEPHALA

257

domesticated. According to Schneider the embryos take up
their residence in the grubs or larvae of the cockchafer (Melo-
lontha vulgaris), a discovery which very readily explains the
manner in which hogs become infested. Whether E. gigas be
a human parasite or not, it is certainly very injurious, not to
say destructive, to swine. Although this parasite must be
quite common in England I have experienced great difficulty
in procuring specimens. In the second book of this work I
shall give some interesting particulars furnished by the memoir
of Prof. Verrill and privately by Mr George Wilkins. (See
' Parasites of the Pachydermata'.)

Bibliogeaphy (No. 35). — Blanchard, in ' Cuvier's Regne
Animal/ tab. 35 (good fig.), and in ' Ann. d. Sci. Nat./ ser. xii.
— Bremser, ' Icones/ tab. vi. — Oobbold, " Parasites of the Hog,"
the 'Veterinarian/ 1875. — Idem, 'Manual/ 1. c, p. 123.—
Davaine, 1. c, ' Syn./ p. 83. — Diesing, 1. c, ii, p. 2. — Bujardin,
1. c, p. 503. — Goeze, 1. c, s. 143 (good figs.). — Gurlt, 1. c,
s. 367. — Heller, ' Darmschmarotzer/ 1. c, s. 663. — Lambl, 1. c,
supra, Feb., 1859. — Leuchart, 1. c, Bd. ii, s. 729; also in ' Bibl.
TJniv/ for March, 1863, and in ' Ann. Nat. Hist./ vol. xii, 1863.
— Owen, 1. c, in 'Todd's Cyclop/ (figs, after Cloquet). —
Eudolphi, ' Synops./ pp. 63 and 310. — Schneider, in ' Arch. f.
Anat. und Phys./ 1868. — Idem, in ' Sitzungsb. der Oberhess.
Gesellsch. f. Nat./ &c, 1874 (quoted by Leuckart) ; see also
'Ann. Nat. Hist./ 4th series, vol. vii, p. 441, 1871. — Verrill,
' The external and internal Parasites/ &c, 1. c, p. 109. — Welch,
"The presence of an Encysted Echinorhynchus in Man/'
' Lancet/ Nov. 16, p. 703, 1872.— Westrumb, 'De Helm. Acanth/
(good figs.), 1821.

SECTION IV.— Part II.— Suctoeia (Leeches)

As explained in the Introduction we must regard the Leeches
and many allied forms of Suctorial Annelids as creatures pos-
sessed of semi-parasitic habits. They are, perhaps, something
more than what Van Beneden styles "free parasites"— an expres^
sion which almost looks like a contradiction of terms. I cannot
here, however, stop to discuss questions which lie, as it were on
the border-land of parasitology. Three species of leech are more
or less commonly employed in medicine. These are the grey leech
(Sanguisuga mcdicinalis, Savigny), the green-leech (8. officinalis,

258

PARASITES OF MAN

Sav.), and the dragon-leech (8. interrupta, Moq.-Tandon).
The two former abound in Central and Southern Europe, being
also present in North Africa, the last named inhabiting Barbary
and Algeria. So abundant are leeches in the country bordering
the Mediterranean that during the invasion of Egypt by Napo-
leon the French soldiers suffered seriously from their attacks.
When the men lay down to drink, the leeches (Hamopis sangui-
sorba, Sav.) affixed themselves to their mouths and nostrils,
producing serious distress. They also attacked horses, camels,
and cattle. In like manner the Ceylon and Philippine Island
leeches (8. ceylonica, Moq.-Tand., or 8. tag alia, Meyen), of which
there are several varieties, prove exceedingly troublesome to
Europeans. These leeches, not being aquatic forms, occupy
woods and damp places. Unless the limbs of travellers are
well protected, the presence of the blood-suckers is soon dis-
covered by the trickling of blood from the limbs and lower part
of the body. The leeches even sometimes creep up to the neck
and other adjacent parts. These " free parasites " also attack
horses, causing much loss of blood. Terrestrial leeches
abound more or less in all warm countries. Sir J. Hooker
encountered them in the Himalayas, and they are common in
China, Japan, Java (8. Javonica, Wahlberg), and other eastern
parts. They likewise abound in Brazil and Chili. The Ameri-
can leeches for the most part belong to the genus Hsementeria
(H. Mexicana, H. officinalis, and E. Ohiliani, Filippi). The
last named is common in Brazil, the other two being Mexican
forms. Another species, which is blind, has been found in
Brazil by F. Miiller (Oyclobdella lumbricoides) . Not only the
above-named species, but also many other kinds of leeches are
in the habit of attacking man and the domestic animals, but
the subject is too extended and special to be fully dealt with
in this work. Almost a legion of species are known as exter-
nally parasitic upon Fishes, Chelonian and Batrachian reptiles,
Crustaceans, and Echinoderms.

Bibliography (No. 36) —Blainville, 'Diet, des Sci. Nat.,'
torn, xlvii, p. 257. — Brandt (und Ratzeburg), ' Medicin. Zoo-
logie/ Bd. ii. — Brightwell, ( Ann. and Mag. Nat. Hist./ ix, 1 8 i-.

Biasing, ' Syst./ vol. i, p. 465, and " Eevis. der Myzelminth.

(Abth. Bdellideen)," in ' Sitzungsb. der math.-nat. CI. d. k.
Akad. der Wissensch./ Bd. xxxiii, s. 473— Ebrard, ' Compt,
Rend./ 1856, p. 1012.— Idem, ' Monogr. des sangues Med./
1857 — Filippi, Be, ' Mem. Accad., &c, Torino/ and in ' S. und

AOANTIIOOEPHALA

259

K. Zeitsch./ 1829.— Idem, " Nuovo genere," &c, in ' Gaz.
Med. Lombard/ 1849.— Gwibc, ' Fam. d. Annelid./ s. 109.—
Hofmeister, in ' Burnieister's Zeitung fiir Zool./ 1848— Johnson,
' Treatise on the Medicinal Leech/ — Leuckart, 1. c, Bd. ii,
s. 634-739 (with many reia.)—LeyMg ("Anat."), ' S. und K.
Zeitsch./ Bd. i.—Moquin-Tandon, ' Monogr. de la fam. des
Hirudinees/ 1846. — Idem, in his ' Medical Zoology ' (Huline's
edit.), 1861, p. 137.— Mutter, F., in f Archiv f. Naturg./ 1846.
— Perevra, in his 'Materia Med./ vol. ii, p. 2197, 1853.—
Savigmj, ' Descript. de TEgypte/ 2nd edit.— Idem, ' Syst. des
Annelides/ 1820.— Schmarda, fNeue wirbell. Thiere/ Bd. i
(quoted by Leuckart). — Yirey (and Serullas), in ( Journ. Pharm./
1829, p. 614<—Wagener, in f Troschel's Archiv/ 1858, Bd, i,
s. 244 et seq. — Wahlberg, in f CEfvers. Kongl. Vetensk. Akad.
Forhand./ Stockholm, 1855.

SECTION IV— Paet III.— Arachnida (Pentastomes, Mites,

Ticks).

The Trachearian division of the Arachnida comprises a few-
internal parasites that attack man, and many ectozoa which are
parasitic upon man and animals. The species can only be
noticed very briefly.

Pentastoma tanioides, Rudolphi. — In the system of classifica-
tion adopted by Diesing, this entozoon and its allies are placed
in the division Gephalocotyleen and therefore, in association
with the Cestodes, with which, however, it has no structural
affinity. It was long ago pointed out by Yan Beneden, T. D.
Schubart, Leuckart, and others, that the pentastomes were
Acarine and Lernasan Arthropods; the genus being osculant
between the Acaridge and Lernasadas. The whole subject is
discussed in Leuckart' s profound memoir quoted below.

The adult Pentastoma tcenioides is characterised by the
possession of a vermiform, lancet-shaped body, flattened at the
ventral surface, attenuated posteriorly, and marked transversely
by about ninety rings (fig. 50, 1 and 2). The cephalo-thoracic
segments are continuous with the body, each supporting a pair
of strong retractile chitinous claws; four in all. The head is
truncated, furnished with an oval mouth, armed with a horny
lip. The integument of the body is perforated with numerous
respiratory openings or stigmata. These are wanting in the

260

PARASITES OF MAN

cephalic segment. In the larval state {=Pent. denticulatum)
the body is armed with numerous rows of small, sharply pointed

spines. The adult female
measures from three to
four inches in length, but
the male is only about
an inch long. The genital
a,perture of the female is
situated at the extremity
of the tail, that of the
male being placed at the
front part of the abdomen
in the middle line. The
mode of reproduction is
oviparous, accompanied
by a subsequent and com-
plete metamorphosis.

In the mature condi-
tion this parasite infests
the nostrils, and frontal
sinuses of the dog and
wolf, and also, though
more rarely, the nasal
cavities of the horse and
sheep. In the pupal and
larval states it sometimes

Fig. 50.— Tentastoma tanioides. (1) Male and (2) female, of OCCUrS in the abdominal

Stoffi&S" The egg a"d embry° high'y masn,fied- and thoracic cavities of

the human body, but it is
more frequently found in herbivorous mammals, such as the
sheep, deer, antelope, peccary, porcupine, guinea-pig, hare, and
rat. According to Creplin, it infests the domestic cat. In
these animals and in man the young worms occupy little cysts
within or upon the peripheral parts of the liver and lungs.
I have occasionally found them free in the cavities of the
abdomen and pleura.

In the course of the development of this entozoon, Leuckart
recognises four well-marked stages. The first is that of the
embryo with a boring apparatus. In the second stage, the
embryo has become transformed into a motionless pupa. The
third is the ordinary larval condition characterised by numerous
rows of small spines in addition to two pairs of double claws.

AEACHNIDA

261

The fourth is the sexually-developed stage, furnished with a
simple hook-apparatus, and without integumentary denticles.
u Our Pentastomes, there-
fore," says Leuekart, " ex-
hibit two kinds of larval
forms, an earlier and later
one, such as takes place
in other animals ; this also
occurs even in insects
(Strepsiptera and Meloida),
only that, in our case (i. e.
in Pentastoma), both do
not immediately follow one
another, but are separated
by a resting condition,
which I have designated
as the pupa stage. In
choosing this name I do
not mean to express a complete identity of this intermediate
state with the pupal sleep of insects."

So far as my own observations extend, the pupa, in its later
stages, closely resembles the free larva ; but, as Leuckart points
out, the earlier stages are very different. The embryo, after
encystation, repeatedly casts its skin, and during the intervals
of these several successive moultings, the young animal makes
rapid growth, accompanied by a series of structural changes.
Passing through these it at length acquires the perfected larval
state (P. denticulatum) .

As regards the occurrence of this entozoon in the human
body, the best account is that given by Frerichs. As quoted
in my previous work from Murchison's edition of Frerichs' well-
known clinical treatise, the German savant remarks : — " The
Pentastoma is a parasite which has only recently been dis-
covered in the human subject, but it is, nevertheless, far more
common in the human liver than the echinococcus. It is
devoid of clinical importance, because it does not give rise to
any functional derangements. Pruner ('Krankheit des Orients/
1847, s. 245) was the first who pointed out the existence of the
Pentastoma in the human liver. On two occasions he found an
encysted parasite in the liver of negroes at Cairo, the nature of
which, however, he did not accurately determine. Bilharz and
Von Siebold (' Zeitschr. fur Wissench. Zoologie/ Bd. iv, s. 63)

Pre 51. — Upper third of the hody of Pentastoma
denticulatum. Original.

202

PARASITES OF MAN

recognised in it a new variety of Pentastoma, to which he gave
the name of P. constrictum. In Germany the Pentastoma was
found in the human liver by Zenker (' Zeitschr. f. ration. Med./
1854, Bd. v, s. 224) ; it occurs, however, not only in this gland,
but also in the kidneys, and in the submucous tissue of the
small intestine (Wagner). The parasite is by no means rare
with us. Zenker, at Dresden, succeeded in finding it nine
times out of 168 autopsies ; Heschl, at Vienna, met with it five
times out of twenty autopsies ; Wagner, at Leipsig, once in ten.
According to Virchow, it is more common in Berlin than in
Central Germany. During six months at Breslau I met with it
in five out of forty-seven dead bodies. The Pentastoma-endemic
in Germany is not identical with that which occurs in Egypt ;
the former is the P. denticulatum of Rudolphi." This clear
statement of Frerichs is valuable ; but, as Murchison has also
pointed out, there is some discrepancy between Frerichs and
Kiichenmeister's record of Zenker's experience. According
to Kuchenmeister, Zenker met with the Pentastoma thirty times
in 200 autopsies.

Although from a purely clinical point of view, and speaking
generally, this worm, as Frerichs says, can claim little attention,
yet, as we shall see (when treating of the parasites of the dog),
it occasionally proves fatal to the canine bearer. Not only so,
it may even occasion severe inconvenience to the human bearer.
Quite recently a remarkable instance of this kind occurred in
Germany, some notice of which appeared in the ' Medical Times
and Gazette/ Jan. 4th, 1879, as follows :

" Dr Landon of Elbing (' Berl. Klin. Wochenschrift/ No. 49,
1878) relates the case of a workman, aged forty-two, who soon
after the Franco-German campaign of 1870 was laid up with pain
in the hepatic region, jaundice, and gastric disturbance, which
symptoms persisted more or less until 1874, when he came
under Dr Landon's care with an attack apparently of perihepatitis.
It then appeared that since 1871 he had also suffered from severe
attacks of epistaxis, which occurred often twice in the same day.
The patient complained of a feeling of painful pressure in the
left nasal cavity, but with the speculum nothing but a moderate
degree of inflammatory swelling could be detected. Suddenly,
at Easter, 1878, a parasite was disloged from the left side of
the nose by a violent sneeze, and from that moment the epis-
taxis has not occurred. Its cause proved to be the Pentastoma
tanioides."

AKAOHNIDA

2G3

As the full-grown parasite occupies the nasal chambers of the
dog, it is clear that the act of sneezing will be liable to transport
the eggs and their contained embryos to the face and other ex-
posed parts of persons who fondle dogs. In this way the germs
will readily gain access to the human mouth. Ordinarily, the
2*erms are introduced into the human stomach with uncooked
vegetable food and fruits, to which they adhere after expulsion
from the animal's nostrils. The slimy nasal mucus secures
this attachment, especially when it has become dry by exposure
to the air. On reaching the stomach the embryos escape the
egg-coverings and bore their way directly to the liver and
other viscera, in which organs they becomes encysted and
undergo the pupal transformation. Eventually they acquire a
length of 2 to 2\ lines (P. denticulatum) . After a while the
capsules enclosing the larvEe undergo calcareous degeneration,
the parasite perishing.

In the case of dogs it is easy to perceive that when the
animals are engaged in devouring the flesh of herbivora, the
liberated larvse will often come in contact with their noses. In
this way contraction of the body, aided by the integumentary
denticles, will secure their entrance into the nasal cavities. For
our own security, therefore, we should avoid contact with dogs
which frequent butchers' shops and knackeries, and be sure that
our market-garden fruits and vegetables are carefully washed
before they are brought to table.

Pentastoma constrictum, Von Siebold. — This parasite is at
present only known to us in the immature condition ; unless,
indeed, as is by no means improbable, the adult worm has been
described under some other name. It was first discovered by
Pruner on two occasions in negroes, and he also subsequently
found two specimens of the worm preserved in the Pathological
Museum at Bologna, which had been removed from the human
liver. Pruner also found it in the giraffe. Bilharz afterwards
frequently detected it in the livers of negroes at Cairo. It
differs from the larval form of P. tcenioides in not possessing
integumentary spines ; moreover, it is a much larger parasite.
The cephalothorax is furnished with four foot-claws, and the
elongated abdomen displays twenty-three rings placed at tole-
rably regular intervals. The anterior part of the animal is
obtusely rounded off, the caudal end being conical. The worm
usually attains a length of rather more than half an inch, whilst
the breadth scarcely exceeds a line.

264

rARASITES OF MAN

An extremely interesting account of this worm has been
published by Prof. Aitken, accompanied with illustrations by
Dr H. C. Gillespie, taken from specimens iu the Pathological
Museum at Netley. Two cases are recorded. In one of these the
encysted worms were found in the liver and lungs,
and in the other in the liver only. In Dr Craw-
fo7-d's account of the post mortem in the last- men-
tioned case, Prof. Aitken quotes him as saying :
" These worms varied in length from an inch to an
inch and a half, and were found coiled up like a
watch-spring, in small sacs scattered throughout
the whole organ." The patient was a private
of the 1st West India Regiment, and died at
Bathurst, Gambia, in 1854'. In the other case,
where the lungs and liver were infested, the patient
was an African, about twenty-one years old, who
had enlisted into the 5th West India Regiment
at Up Park Camp, Jamaica. He had, a few months
previously, come from the slave depot at Rupert's
Valley, St Helena. According to the post-mortem
report, furnished by Mr Kearney (staff surgeon),
the lower lobe of the right lung contained one or
two yellow specks. " When cut into, worms were seen regularly
encysted in its substance." The surface of the liver was
dotted over " with about twenty or thirty yellow specks, similar
to those seen in the lung." The longest of these specimens
was a trifle less than three quarters of an inch.

Whether Pent, denticulatum be or be not devoid of clinical
interest, it is quite clear from Aitken's account that P. con-
strictum is a formidable parasite and one that occasionally proves
fatal to the bearer. As his remarks suggest, a parasite that
can produce both pneumonia and peritonitis is not a creature
that either the physician or the sanitarian can afford to ignore.
Lastly, I must again express my belief that the so-called
Echinorhynchus, described by Welch, if it be not the Pentastoma
denticulatum, must either be referred to P. constrictum (in an
early larval condition), or to some other hitherto undescribed
pentastomatoid larva.

Bibliography (No. 37). — AitJcen, W., " On the occurrence of
Pentastoma constrictum in the Human Body as a cause of painful
Disease and Death," repr. from the ( Science and Practice of
Medicine/ 1th edit., 1865. — Bellmgham, in 'Ann. Nat. Hist./

Fig. 52 —Pentas-
toma constric-
tum. Magnified
four diameters.
After Bilharz.

AT? AOHNIDA

265

vol. xiv, p. 162 — Blanchard, in 'Ann. des Sci. Nat./ ser. 3,
t. viii, and in ' Kegn. Anim/ (with figs.). — Oobbold, ' Entoz./
p. 393 et seq. — Idem, in ' Quart. Journ. Med. Sci./ 1859, p. 205.
—Idem ("P. cephalophi"), in 'Linn. Trans./ xxii, p. 357, and
xxiii, p. 350.— Idem, in ' Zool. Soc. Proc./ 1861, p. 124. —
Diesing, ' Syst./ i, p. 609. — Idem, ' Eevis. der Cephalocot./
s. 327. — Frerichs (1. c, in text), vol. ii, p. 276. — Klob (und
Schroff), in ' Gesellsch. d. Aerzte/ Wien, 1860. — Kiichenmeister,
1. c., i, s. 370, Eng. edit., tab. viii. — Idem (with Van Beneden),
in 'Bullet. Acad. Belg./ xxii (with figs.), 1855. — Landon
(quoted in text). — Leuckart, in 'Zeitsch. f. rat. Med./ 1857 ;
see also " Obs. on the development and early condition of the
Pent, tanioides," in ' Ann. Nat. Hist./ vol. iii, 3rd series, 1859 ;
also my translation of his " Further Observations on the
development of P. tanioides," from ' Henle and Pfeufer's
Zeitsch./ in the 'Quart. Journ. of Micr. Sci/ for 1859. — Idem,
' Bau und Entwicklungsgeschichte der Pentastomen, nach Un-
tersuchungen besonders von P. tanioides und P. denticulatum,'
Leipzig, 1860. — Moquin-Tandon, 'Med. Zool/ (Hulme's edit.),
"The Linguatula," p. 329. — Pruner (" Nematoideum ") in
'Krankh. d. Orient./ 1847. — Schubart, ' S. und K. Zeitschr./
Bd. iv. — Welch, see Bibl. No. 36. — Zenker, in ' H. und Pf.
Zeitschr. f. rat. Med./ 1854, s. 212 (with figs.).

The ectoparasitic arachnidans comprise a great variety of
mites and ticks [Acaridae and Ixodidce) more or less proper to
man, and also a number of creatures which, though hardly to
be reckoned as human parasites, are apt to transfer themselves
from animals to man. Little more than an enumeration of the
forms is possible here. The Common Scab or Itch insect
(Sarcoptes scabiei) forms the type of a great variety of arach-
nids, generally spoken of as different species according to the
host they dwell upon. Megnin, however, in his beautiful
memoir, quoted below, regards most of the forms of this genus
(found on the horse, hog, sheep, dog, wolf, and other animals)
as mere varieties. In man the female Acarus burrows beneath
the skin, forming galleries or curved channels, in which she
deposits her eggs. The irritation produced is not alone due to
these excavations, but to the presence also of a poison which
the mite discharges when feeding. The Sarcoptes crustosm of
Fiirstenberg, producing the Norway itch, is a variety, if
indeed, it can be called as much. Under the frightful name of
J lermatophagoides Schetemetewsky two parasites found on an

2GG

PARASITKS OF MAN

herpetic patient have been described as new to science by M.
Bogdanoff, but Megnin points out that these Acari are only
female and young male representatives of his Ghorioptes seti-
ferus (var. bovis) respectively. In Newfoundland, Dr Le Roy de
Mericourt discovered a singular species upon an officer who
had come from Havannah (Tyroglyphus Mericourti, Laboulbene).
It possesses enormous palpi, as in the genus GhyJetus to which
Robin refers it. Another ectozoon, placed by Megnin and
others amongst the lowest types of Arachnida, is the well-
known Demodex folliculorum. It is a gregarious species, a

Fig. 53. — Demodex folliculorum, var. caninus. a, Female; b male. Viewed from below
arid in profile. Magnified 300 diameters. After Megnin.

dozen or more examples often being present in a single dilated
hair follicle. Though disfiguring to the human face it pro-
duces little harm. M. Gruby made it out to be a very common
parasite, infesting forty out of sixty persons ; but Megnin, in
his brochure (1. c. infra, p. 119), shows this statement to be an
exaggeration. It infests on the average not more than one in
ten persons. According to Gruby, moreover, a single follicle
in the dog may contain 200 of these mites, another statement
which Megnin deems unreliable. The Demodex of the dog is only
a variety (fig. 53). Many other human Arachnids have been found,
some of which appear to be genuine species, whilst others are

AlvACHNIDA

267

accidental, so to speak. Of the former kind, perhaps we may
reckon the two species discovered by Hessling {Ocelognathus
morsitans and Entarsus cancriformis) . Of the latter sort, those
found by Busk, Simon, and Bory de St Vincent may be cited.
The mite found in Simon's case was the Dermanyssus avium,
which infests cage-birds. Probably it was the same species
which Bory found on a lady ; but in Busk's negro sailor the
mite may have been D. gallince of the common fowl. Differ-
ing from the mites, proper, and also from the true ticks, are
some bug-like forms called Argades. The two best known are
the Miana bug of Persia (Aryas persicus) and the Chinche of
Columbia (A. chinche). Like their congener infesting pigeons
(A. refiexus) these parasites are terrible blood-suckers. The
bite of the Persian bug is so venomous as to have occasioned
death. Various species of tick have been known to attack man,
but the species have not been well determined. Although a
human form has been described (Ixodes hominis, Koch), yet it
is more probable that the species usually attacking man are the
same as those known to infest the domesticated animals. In
this list we may, therefore, reckon Ixodes nigra, Ix. bovis, Ix.
ricinus, and Ix. reduvius. Cases in which one or other of
these ticks occasioned much pain and distress are recorded by
Hussem, Raspail, and Dr Cosson. Besides these there is a
formidable tick well known at Angola (Ix. monbata). Its
habits are like those of the common bed-bug. Severe pain
comes on two hours after the person is bitten. It likewise
attacks animals. The Ix. carapato is similarly troublesome in
Brazil. Another very disgusting arachnid liable to attack man
is the Galeodes araneoides. This large spider-like creature, two
inches in length, commonly attacks camels and has an extremely
venomous bite. One or more species of the dung-beetle mites
(Gamasidae) have also been known to fasten themselves on
man. According to Latreille, they first get attached to the
clothes of travellers, whence they pass to his body, and there
shift about, producing great torment. Another disagreeable
arachnid is the little harvest bug (Leptus autumnalis) , which
not only excites irritation during its crawling motion on the
human skin, but even succeeds in burying itself near the hairs.
The irritation thus produced is almost unbearable. This mite
attacks various animals, especially dogs and cats. I myself
once suffered severely from this species in consequence of fond-
ling a young wild rabbit which, as I afterwards discovered, was

208

FAItASITES OF MAN'

much infested. When the parasites had reached my left arm-pit
they occasioned extreme torture. I have known these autumnal
spiders to produce small suppurating boils on the abdomen.
I may add that Dr Tilbury Fox has brought under my notice
an instance where the hexapod larva of another species (probably
Trombidium cinereum) was found to have occasioned severe
irritation in a child.

Bibliography (No. 38). — Alibert, ' Maladies de la Peau/ Paris,
1833. — Audouin, V., art. " Arachnida," in 'Todd's Cyclop./
vol. i, 1836. — Beneden, Van (et Gervais), ' Zool. Med./ 1859. —
Bourguignon (et Delafond), in ' Rec. Yet./ 1856. — Idem, in
'Mem. de PTnstitut./ 1862. — Gobbold, "Case of Leptus produc-
ing Boils/' in 'Worms/ p. 140, London, 1872. — Gamgce, ' Our
Domestic Animals in Health and Disease/ Edin., 1861. —
Gerlach, ' Kraetze und Raude/ 1857. — Hebra, in ' Oester. Jahrb./
1864. — Hering, ' Die Kraetzmilben/ Stuttgard, 1845. — Krabbe,
" Husdyrenes paras. Mider.," ' Tidssk. f. Yet./ Rook. 2, Bd. iii.
— Kuchevmeister, 1. c, 1855, s. 412 (good figs.). — Me'gnin,
' Monographic de la tribu des Sarcoptides psoriques.' (This work
contains a full bibliography and numerous beautiful plates ; see
also Review in the ' Yeterinarian/ Aug., 1877, p. 563). — Idem,
" Memoire sur un nouveau Symbiote (Ohorioptes, Gerv.),"
' Journ. de TAnat. et de la Physiol./ 1872. — Idem, "Mem. sur
un nouvel Acarien," ibid., 1873. — Idem, " Mem. sur les Hy-
popes," ibid., 1874. — Idem, "Mem. sur ^organisation et la
distribution zoologique des Acariens de la famille desGamasides,"
ibid., 1876. — Idem, "Mem. sur les metamorphoses des Acariens
en general, et en particulier sur celles des Trombidions," 'Ann.
des Sci. Nat./ 1876. — Idem, "Des conditions de la contagion
de la gale des animaux a l'homme/' 'Arch, generales de Med./
1876. — Idem, "Mem. sur le Demodex folliculorum (Owen),"
' Journ. de l'Anat. et de la Physiol./ 1877. — Moquin-Tandon ,
'Elem. de Zool. med/ (Hulme's edit., p. 302-328), 1861.—
Williams, in his ' Veterinary Surgery ' (good figs., reproduced
from Gamgee's translation of ' Gerlach/ &c), 1872.

SECTION IV.— Part IV— Crustacea (Gammarida;).

Although multitudes of small Crustaceans are parasitic upon
fishes, and some few of them adhere to oceanic mammals
(Cetacea), I am sceptical as to the parasitism of Crustacea either

CRUSTACEA — IN SECT A

260

in or upon man. Many of the Amphipodous Gammari lead a
sort of free parasitic existence, and they are themselves very
liable to harbor larval parasites. As regards human parasitism
from this source the only records known to me are those quoted
below.

Bibliography (No. 39). — Banon, " Freshwater Shrimp, or
Gamarus pulex (said to have been passed per anum)/' ' Rep. of
Path. Soc. of Dublin/ in ' Dub. Med. Press/ April 6, p. 351,
1864. — Bartels, " Gamarus pulex in the Human Subject, with a
postscript by Troschel/' trans, by Dr E. P. Wright, from < Ver-
handl. des Naturhist. verein. der Preuss. Rheinl. und Westph./
in ' Dubl. Med. Press/ 2nd ser., vol. ix, p. 407, 1864.-1^™^,
E. P., ' Eemarks on Dr Banon's Case 3 (see Bartels) .

SECTION IV. — Part V. — Insecta (Coleoptera, Diptera,
Hemiptera, Aphaniptera) .

Whilst very many flies, bugs, lice, and fleas persecute animals,
not a few of them also attack man. Several of the species are
genuine parasites, others are semi- parasitic, and others, again, are
altogether outside the border-land of parasitism in the ordinary
sense of the term. In fact, it becomes difficult to say where
the line of parasitism should be drawn. I cannot, however,
ignore all notice of the insect tormentors, whether strictly
parasitic or not.

At least fifty different species have been regarded as playing
the role of parasitism in man. Amongst the Coleopterous
parasites none is more authentic than Blajps mortisaga. At
least half a dozen such cases have occurred. Mr Hope's
catalogue of insects producing parasitism in man gives three
examples of scolechiasis from this source. Sir J. R. Cormack
published a fourth case, and I have recorded a fifth. In this
instance I received the living larva from Dr Home, of Barnsley,
who procured it from an infant eleven weeks old. In my ' Intro-
ductory Treatise ' I have alluded to the case of the girl Riordan,
who not only passed per anum upwards of 1200 larvas, but also
several perfect insects. The case was first reported by Pickells,
Thomson, and Bellingham. One of the other authentic cases,
in which only a few larvae were present, was recorded by
Patterson, of Belfast, and the third case by Bateman. Mr Hope's

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rA It A SITES OF MAN

' Catalogue ' originally appeared in the ' Transactions of the
Entomological Society/ being afterwards published in the pages
of the ' London Medical Gazette/ 1837. Patterson's case was
also, I believe, first communicated to the Entomological {Society.

As regards the mode in which the maggot gained access to
the child in Home's case, it was not easy to decide ; but in the
case of the girl Riordan the mode of ingress was sufficiently
explained. The Blapsidcs, as a family, are closely allied to the
meal-worms, and, like most of the Tenebrionidce, are black and
foul-smelling beetles, frequenting dark and damp situations,
from which they escape only at night. The family comprises
numerous species, of which probably not more than three are
found in this country. They are abundant in Africa, especially
in Egypt, where (according to Fabricius, as quoted by West-
wood, Figuer, and others) the women eat Blaps sulcata cooked
with butter in order to make themelves grow fat. The insects
are also employed as specifics against ear-ache and the bite of the
scorpion. The superstitious notion of a " charm " is generally at
the bottom of these domestic remedies. In the girl Eiordan's case,
as Westwood observes (when epitomising Pickell's account), the
parasites, as such, " probably originated in an absurd and super-
stitious practice, which she had for some time followed, of drinking
daily for a certain time a quantity of water mixed with clay,
taken from the graves of two Catholic priests, and eating large
pieces of chalk. One of these beetles was immersed repeatedly
in spirits of wine, but revived after remaining therein all night,
and afterwards lived three years." The intolerance of light
shown by the perfect insect seems to be equally shared by the
larva. Of this fact I had repeated evidence by observing the
behaviour of the living specimen sent to me by Dr Horne. Thus,
when, on February 5th, 1877, I placed the maggot on the
surface of some moist mould, scarcely half a minute elapsed
before it commenced to bore its way downwards, and in less
than a minute all but the tip of the tail had disappeared. In
like manner, when, on the 7th, I raised the lid of the box, and
found the maggot on the surface of the soil, it almost instantly
proceeded to bury itself. Hope's list records no less than nine
instances of parasitism in man from the larvae of Tenebrio
molitor, and he gives a score of other Coleopterous insects
which he regarded as human " intestinal worms." Undoubt-
edly a large number of insect larvae do get introduced into,
and actually live within the human intestines.

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271

Numerous cases of this sort have come under my observation,
but it often requires a profound knowledge of entomology to
determine the species. Several examples of oestridean larvae
occur amongst them. For one example of (Estrus hominis I am
indebted to Mr Higginson, of Liverpool, who obligingly supplied
me with notes of the case. J\r Kirk presented me with a
small bot which he removed from Livingstone's leg. I after-
wards deposited the African bot, in his name, in the Hunterian
collection. Bates speaks of an (Estrus in Brazil producing boils
in human flesh. "Westwood quotes similar instances, Of these,
one was extracted from the thigh by Dr Brick. Mr Doubleday,
the entomologist, extracted one from his own leg, and M. Goudot,
another entomologist, was also victimised in the same way.
Both of these savans were travelling in America at the time.
Two cases from South America were also recorded by Howship.
In one the larva lodged in the back ; in the other in the
scrotum. Humboldt noticed that the Indians were much
infested by CEstridae. Three cases are severally recorded by
Eoulin, Guerin, and Audouin. Mr Stroop also mentions a case
in which an (Estrus was removed from an ulcer on the shoulder
of a boy in Texas. Another kind of bot known as the Macaco
worm (Cuterebra noxialis) occasionally attacks man, but more
frequently cattle and dogs. For one example, taken from the
leg of a negro, at Belize, British Honduras, I am indebted to
Dr Dobson (A. M. D., Netley). Hope's list records five cases of
bots of (Estrus hominus, one of (E. Guildingii, one of (E. bovis,
and thirteen others (belonging to the same genus) as having
been noticed in man. Since his time many additional cases
have been recorded by J. M. Duncan and others. In like manner
a legion of cases in which the maggots of various Muscidae have
been noticed, either in, upon, or from the human body. At
a meeting of the American Academy in April, 1859, Dr Leidy
showed some larvae of the bluebottle fly vomited by a child ;
five larvae of the flower-fly (Anthomyia) from a physician's
own person (which had produced choleraic symptoms) ; and
nine examples of Cuterebra noxialis. I have myself encountered
numerous insect larvae in medical practice. Amongst others
I have obtained the larvae of Anthomyia canicularis in six
or eight separate instances. One set of specimens, identified by
Wunderlich, was sent to mo by Dr Brandt, of Oporto. Drs
Duffin, W. Fox, and Learcd have supplied me with others.
The larvae described in Farre's case, not being setose, must be

272

PARASITES OF MAN

referred to one or other of the Muscidae proper. Mr Hope gave
nearly forty cases of this kind, referable to eight different
species of fly, and, as already implied, I have myself collected
a great variety of the maggots of Muscidae passed by the bowel,
besides others obtained either from beneath the skin or from
open ulcerations.

The flies hitherto noticed as supplying parasitic maggots in
man are Musca domestica, M. carnaria, M. larvarum, M. nigra,
and perhaps H. Cibaria, M. stabulans, and M. Gcp.sar. This last,
a handsome fly, is the species which proves so troublesome to
sheep. The habit which flies have of depositing their eggs in
open wounds, when the victims are asleep, is a fertile source of
this kind of parasitism. Some of the instances recorded by
Kirby and S pence are revolting in the extreme. They quote
the cases which came under Mr Sell's notice in Jamaica. In
one instance the flies were hatched in a neglected blister on the
chest ; in another from the gums and inside of the cheek ; and
in a third, from the nostrils of a negro, from whom 235 larvse
were expelled. The case of the Lincolnshire pauper, Page, who
was literally eaten up by maggots, is almost incredible. An
equally horrible instance, however, is recorded by Cloquet. It
is said that the Jamaica cases were all due to the larvse of the
bluebottle fly (M. vomitoria) . An instance of the same kind has
been recorded by Mr Knox (A. M. D.). Sufficiently revolting as
these cases are, the horrors attending them are eclipsed by the
habits of the larvae of Lucilia hominivora. The best accounts of
its habits are those by M. Coquerel, M. Saint-Pair, and M.
Yercammer. The insects lay their eggs in the mouth and nostrils,
and when the larvae escape they devour the tissues surrounding
the buccal cavity, the pharynx, glottis, frontal and nasal sinuses,
even reaching the sockets of the eye. Several Cayenne convicts
have perished from the maggots of this fly, which is also pre-
valent in Mexico. These are not, however, the only instances of
maggots gaining access to the nasal chambers. In a case
recorded by Dr Astros, of Aix, 113 were discharged from the
nose of a woman ; and M. Legrand du Saulle records an instance
where a number of larvae occupied the frontal sinuses of a girl,
nine years of age. The larvae produced persistent headache and
convulsions. In the case recorded by Wohlfart, 18 larvae were
discharged from the nose of an old man, and in the example
given by Latham several larvae were obtained from the frontal
sinuses of a woman. Bracey Clark also gives an instance in

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273

which a bot was taken from a woman's jaw. Not improbably
the well-known Indian disease, termed peenash, or worm in the
nose, is due to the presence of OEstridean larvse. Cases by
Eustomjee and Lahory are quoted below. Possibly Stockett's is
another of the same order. The case by Kilgour (Bibliog. No.
34) may be another. I may add that Moquin-Tandon gives an
instance of the discharge of seventy-two bots, resembling those
of the sheep, from a woman's nose (e Journ. de Vandermonde ').
The rat-tail maggots or larvae of Helophilus are parasitic. Two
or three instances have been recorded from the horse. I possess
one from the human intestine. Kirby also quotes an instance
in which Heloph. pendulus was obtained from the stomach of
a woman (' Philos. Mag./ vol. ix, p. 366).

A vast number of non-parasitic insects are injurious to man
and beast. Inasmuch as they subsist at the expense of their
victims and also adhere to his person during their attacks for a
shorter or longer time, they, like the leeches, may be spoken of
as free parasites. The leg-sticker (Stomoxys calcitrans) pene-
trates through thick stockings, causing blood to flow freely.
The clegg of the West Highlands {Hcematopota pluvialis) also
violently attacks man and beast, especially horses. The mosquito
(Gulex anxifer), the gnats (G. pipiens, C. annulatus, and G.
rpulicarisf) , and the midge (Ghironomus plumosus) need only be
mentioned. The creeping gnat (Simulium reptans) is also very
annoying in Sweden. The role of the mosquito, as itself con-
stituting a parasite-bearer, will be again referred to in the
closing pages of this work (Book II, Section V). The bites of
the tsetse (Glossina morsitans), though so destructive to the
horse, ox, sheep, and dog, are not dangerous to man himself.
According to Sir S. Baker, the seroot-fly, or zimb of Bruce,
which is a species of Pangonia, is excessively annoying to
travellers in Abyssinia. Amongst the hemipterous insects the
common bug (Acanthia lectularia) is sufficiently blood-thirsty ;
but there is a far more sanguinary species of this kind in
South America. This is the pampas benchucha (Conorhinus
nrgrovarius). Our distinguished countryman, Darwin, in his
* Voyage,' speaking of these wingless insects, says : — " Before
sucking they are quite thin, but afterwards become round
and bloated with blood. In less than ten minutes the insect
is changed from being as flat as a wafer to a globular form."
This insect somewhat resembles our water-scorpion [N&a
cincrea), a non-parasitic species whose bite causes severe pain

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PARASITES OP MAN

as does also the wound inflicted by the water-boatman
(Notonecta glauca). There are other species of bug, such as
the Acanthia rotundata of Eeunion Island and A. ciliata of
Kasan, the bites of which are worse than that of the common
species. The fly-bugs also, such as the Beduvius personalis, so
common in France, and the B. amcenus of Borneo and Java,
attack man, although their especial habit is to attack and destroy
other insects, including bugs themselves. Other species or
varieties of Eeduvius (B. cruentus, and B. serratus) attack man.
The last named is an Indian form, capable, it is said, of pro-
ducing an electric shock.

Passing to the fleas, the most important and truly parasitic
form is the chigoe or gigger (Pulex penetrans), This abounds
in tropical America and the West Indies. The female insects
only attack man, and this they do for the purpose of securing
a lodgment for their offspring. They attack especially the
soles of the feet, between the toes and near the nails. In bad
cases the whole of the foot becomes affected. After the insect
has penetrated the skin its body swells enormously and becomes
a mere bag of eggs. This swelling causes active inflammation,
which terminates in suppuration and the formation of open
ulcers. The chigoe also attacks various animals. In addi-
tion to the common flea (P. irritans) other species infesting
animals are said to attack man occasionally. As regards those
degraded types of insects known as lice I can only afford space
to remark that five species have been recognised as human.
These are the head-louse (Pediculus capitis) ; the louse of the
eyelids (P. palpebrarum) ; the clothes-louse (P. vestimenti) ;
the distemper louse (P. tabescentium) ; and the pubic louse (P.
inguinalis). The distress these creatures occasion is only fully
known to physicians who deal with the obstinate cutaneous
affections caused by their presence (Phthiriasis) . Some of the
lice found on negroes and Greenlanders are regarded as distinct
varieties. Lice are abundant on animals, and some of the
species are apt to transfer themselves from one host to another.
Thus the Ornithomyia avicularis of cage birds has been found
on man, whilst one or more of the numerous species infesting
the common fowl are, by transfer, apt to produce a severe
phthiriasis in the horse. The lice of the fowl belong to the
genera Leipurus, Liotheum, Menopon, Philopterus, Ooniodes, and
Goniocotes. The unsuitableness of man's person as a habitation
for bird-lice should, however, check the fear which many porsons

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275

have of handling fresh-killed poultry and game birds. Poultry
lousiness in man is probably impossible from this source.

Bibliography (No. 40).—Arture, " Obs. sur le ver nomme
Macaque/' in 'Mem. Acad, des Sci./ Paris, 1753. — Bates,
" (Estrus in Man/' ' The Naturalist on the Amazons.' — Beneden,
Van, ' Animal Parasites and Messmates/ 1876 — Idem (with
Gervais), ' Traite de Zool. Med.'— Blood, M., " Case of Larvae
(Musca sarcophaga) expelled alive in the Faeces/' ' Beale's Arch,
of Med./ vol. iii, p. 134, 1862. — Brinton (similar case), ' Arch, of
Med./ vol. iii, p. 133, 1862.— Bouyer (quoted by Figuer),
* Tour du Monde/ p. 318, 1866. — Clark, Bracey, in ' Linn.
Trans./ vol. iii, 1797 (the jaw-case at p. 323), and vol. xv, 1827.
— Idem, ' Essay on Bots/ 1815. — Cloquet, J., " Case of Blind-
ness from Worms (larvae of Musca) in the Eyes/' from ' Arch.
Gen. de Med.,' in 'Lancet/ 1828. — Oobbold, " On Blaps morti-
saga as a Human Parasite," ' Brit. Med. Journ./ 1877, p. 420.
—Idem, " Entoz." (f Hope's List.'), p. 416— Idem, in 'Worms'
("Leptus," &c), p. 140, 1872. — Goquerel (quoted by Figuer).
— Oormach, J. B., " Exhibition of a Larva {B. mortisaga) passed
by a Child," 'Month. Journ. Med. Sci./ vol. i, 1841. — Crumpe,
S., " History of a Case in which Worms (larvae of a beetle)
were discharged from the Stomach," from ' Trans, of the Roy.
Irish Acad./ vol. vi, in ' Med. Facts and Observ./ vol. viii,
p. 229, 1800. — Denny, 'Monog. Anoplurorum Brittaniae/ 1842.
— Duncan, J. M., " On the Occurrence of Bots in the Human
Subject," ' Edin. Vet. Rev./ vol. i, p. 275, 1858-59.— Idem,
" The Larva of (Estrus bovis in the Human Subject," ' Rep. of
Edin. Med.-Chir. Soc./ in ' Month. Journ. of Med. Sci./ July,
1854. — Farre, A., " On the Larva of Anthomyia canicularis"
'Micr. Journ. and Struct. Rec./ 1841, p. 129, and in 'Trans,
of Micr. Soc. of Lond./ orig. ser., p. 51, 1844. — Figuer, in
' The Insect World ' (good figs.), Janson's edit., 1869.— .FW-
longe, " Anat. of Pulex," in the ' Journ. of the Queckett Club/
vol. iii. — Geer, De, ' Memoires pour servir a l'Histoire des
Insectes/ 1773. — Gervais (see Van Beneden). — Hill, "Account
of the Larva of a supposed OEstrus hominis, or Gad-fly, which
deposits its Eggs in the bodies of the human species, with
particulars of a Case," ' Edin. New. Phil. Journ./ vol. xxii,
p. 284, 1830.— Hilaire (see St Hilaire) .— Hope, "Tables of
Cases of (spurious) Intestinal Worms," ' Lond. Med. Gaz./
1837-38. — Hoppe, " Case of Larvae of Insects {Musca stabulans)
passed by Stool," from ' Bibl. fur Lager/ in ' Med.-Chir. Rev./

276

PARASITES OF MAN

1842. — Hopper, B. S., " Insects (Stratiomis) voided with Urine,"
edit, note in ' Micr. Journ. and Struct. Rev./ p. 160, 1841.—
Joly, ' Recherches sur les CEstrides en general/ &c, Lyons,
1846. — Keferstein, 'Sur TOistros/ Isis, 1827.— Kirhy (and
Spence), ' Introd. to Entomology/ 7th edit., 1856. — Knox, M.,
" Maggots, the larvae of the Bluebottle Fly, in Syphilitic Ulcera-
tion of the Throat," ' Lancet/ Oct. 6, 1877, p. bU.—Lahory,
B. T. 0., " On Peenash, or Worms in the Nose," ' Ind. Ann. of
Med. Sci./ 1855, and ' Edin. Med. Journ./ 18b7.—Leaeh}
" CEstridae," in ' Wernerian Trans./ 1817.— Leidy, in f Proc.
Phil. Acad. Nat. Sci./ 1859, p. 1. —Maclean, " On the Oistros,"
' Linn. Trans./ vol. xiv, 1824, and in ' Zool. Journ./ vol. i and
iv. — Metaxa, "Vom OEstrus (u. s. w.)," in ' Neuen Nord.
Beitr./ Bd. i, and in 1 Mem. de Zool. Med./ Rome, 1835 (quoted
by Westwood). — Moquin-Tandon, ' Med. Zool.; (1. c, Bibl.
No. 38).— Newport, art. "Insecta," ' Todd's Cyclop./ 1839.—
Bustomjee, B., " Case of Worms in the Nose, or ' Peenash/'" in
' App. to Bomb. Med. and Phys. Soc. Trans.,'' No. vii, new ser.,
p. 21, 1861; see also Lahory, on ' Peenash/ — Saint-Hilaire,
11 Sur l'CEstre chez l'horome," in ■ Ann. Soc. Ent. de France/
1833. — Say, " Brick's Case," in ' Trans. Acad. Nat. Sci. Phil./
vol. ii. — Sells, in ' Trans. Entom. Soc. Lond. / see also Lem-
priere's 'Diseases of the Army in Jamaica/ vol. ii. — Stoclcett,
T. H., " An account of a Headache cured by the discharge of a
Worm (?) from the Nose," ' Med. Com./ vol. xix, p. 157, 1794,
and in f Trans. Coll. of Phys. Phil./ vol. i, part i, p. 181, 1793.

— Stroop, St J., " CEstrus," in ' Amer. Naturalist/ vol. vii,
p. 437. — Tanner, " On Lice," in his ' Pract. of Med./ vol. ii,
p. 429, 6th edit., 1869. — Westwood, in his ' Classification of
Insects/ vol. ii, 1840. — Wohlfart (quoted by Moquin-Tandon).

— Yule, "Case of Larvae of Insects in the Human Stomach,"
fEdin. Phil. Journ./ and 'Lond. Med. Repos./ 1825.

SECTION IV. — Part VI. — Protozoa (Psorospermioe, Grega-

rinidas, &c).

The scope of this work does not demand that I should com-
prise within its limits any vegetable parasites ; nevertheless, I
must needs refer, however briefly, to certain confervoid and
sarcodic organisms, which, for the most part, lie on the border-
land of the animal and vegetable kingdoms. Professor Cohn

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277

regards the bacteria as allied to the Oscillitoriaceae He puts
them in his order Schizosporea. It is of little moment,
practically, where these protista forms are placed. Unquestion-
ably many of them are parasitic, as they live in the tissues,
fluids, and secretions of animal bodies, including man. Their
presence in cattle is associated with an anthracoid disease
(charbon), whilst in the human body they have been detected in
connection with zymotic affections. They have been found by
Cohn, Sanderson, Klebs, Chauveau, and others, either in the
lymph of vaccine pustules, or in the miliary eruptions of typhus
fever. Professor Beale, who was one of the first to observe
these special organic particles in vaccine lymph, denies that
they are true Bacteria ; and, indeed, he warmly disputes the
inferences that have generally been drawn from the fact of the
presence of such particles in lymph, blood, and other nutrient
fluids. The best known and defined forms are Bacterium
termo and Bad. lineola, which are concerned in the production
of putrefaction, Bacillus anthracis, found in the blood of animals
suffering from carbuncular disease, Micrococcus septicus, found
in typhus and pyaemia, M. vaccina of cow-pox lymph, and M.
di'phthericus, in diphtheria. As regards their prevalence in
certain forms of relapsing fever, Sanderson states that Dr H. Y.
Carter, of Bombay, examined the blood of 250 fever patients
and found spirilla in nearly every instance. From the inde-
pendent observations of Pasteur, Sanderson, Lister, Tyndall,
Bastian, Eberth, Roberts, Davaine, and many others, it seems
clear that the Bacteria and their allies play an important part in
association with certain morbid states. However, as regards
the etiology of the maladies in which these organisms are found,
it is perhaps too early to speak with absolute confidence. The
subject cannot be dealt with here ; moreover, it is outside the
range of my personal investigations.

Passing to those protozoa which, although retaining some
vegetable affinities, are more or less distinctively animal, I
notice the obscure organisms termed psorosperms. In dealing
with these I shall treat of the forms that infest both man and
animals, confining my remarks to such as happen to have come
under my own observation.

In the year 1865 the public were thoroughly roused to a
sense of danger arising from the consumption of meat. The
panic originated with the outbreaks of trichiniasis in Germany.
During the excitement which subsequently prevailed at the time

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FARASITES OF MAN

of the rinderpest, all sorts of erroneous notions took possession
of the popular mind, and the errors were stimulated by writers
ignorant of helminthology. In January, 1866, I published a
few observations, the purport of which was to show that certain
microscopic organisms found in animals dying from cattle
plague were harmless " parasitic Protozoa," possessing more or
less striking vegetable affinities. About a week previously
some interesting researches on these so-called cattle-plague
bodies had been published by Dr Beale. Those who first saw
these bodies thought they had stumbled upon organisms new to
science. I showed that similar or analogous organisms were
to be met with in a great variety of animals, and likewise in
the human body. They had been called worm-nodules, worm-
nests, egg-sacs, eggs of the common fluke, young "measles,"
corpuscles produced by muscular degeneration, psorospermiae,
stages of growth of gregarinse, amoeboid bodies, and so forth.
In so far as the higher animals were concerned, Bujardin was
the first to describe them. He found these organisms in a
mole. This animal, however, having been fed upon earth-
worms known to harbour such parasites, there was no difficulty
in accounting for the source of the psorosperms.

In 1853 Hessling discovered psorospermial sacs in the mus-
cular substance of the heart, not only of the ox, but also of the
sheep and roe. By him they were regarded as evidences of
muscular degeneration. About ten years previously Miescher
found similar bodies in the muscles of the mouse.

In 1857 Rainey described similar structures taken from the
flesh of swine ; and, in his memoir, he went so far as to main-
tain that these bodies were early stages of development of the
common pork-measle. In the year 1858 Gubler wrote an
important paper on this subject, in which he related a case
where twenty cysts existed in the human liver. The cysts
were of great size, mostly as large as a hen's egg, one of them
being some six inches in diameter. Naturally, the largest had
been diagnosed as an ordinary hydatid. However, on evacuat-
ing their contents (post mortem), they were found to harbor
enormous quantities of minute corpuscles strictly analogous to
those usually obtained from psorospermial sacs. Gubler believed
he had stumbled upon masses of eggs of Distoma hepaticum, but
in this he erred. Shortly after Gubler's discovery similar
bodies from the human liver were described by Virchow ; and
in 1862, the subject was followed up by Dr Dressier, of Prague.

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279

Dressier found in the human liver a number of pea-shaped
bodies, the milky contents (breisubstanz) of which displayed a
multitude of the characteristic corpuscular elements referred
to. These particles, already considered as equivalent to, if not
identical with, the so-called pseudo-navicellas of gregarinas, were
soon encountered by a variety of independent observers. Thus,
Leuckart noticed these bodies in various animals ; but with
caution remarked : — " Concerning the nature of these forma-
tions I will not decide. To be candid, however, it appears to
me to be in no way made out whether the psorospermiae are to
be considered as the result of a special animal development,
whether they, like pseudo-navicellae, are the nuclei of gregari-
niform productions, or whether they are the final products of
pathological metamorphosis." Leuckart found these organisms
in the intestines of a trichinised dog, also in a sheep and pig
fed with Trichinae. He also found them in the muscles of
another pig fed with psorosperms, and likewise in the liver of
various rabbits. He remarks that in swine these parasites are
more abundant than measles. They were present in five of
eighteen pigs, and also in two out of four sheep, whose flesh
was especially examined. The observations of Lindemann at
Nischney-Novgorod are particularly interesting. This medical
officer discovered psorospermial sacs attached to the hair of a
girl who was being treated in hospital for chlorosis. The sacs
in question bore close resemblance to the bodies which we
found in abundance in diseased and healthy cattle. It would
further appear, from Lindemann's observations, that the affec-
tion is not very uncommon amongst the Russian peasants.

In connection with and attached to the same parasitically
affected hairs Lindemann also noticed several movable grega-
rinas ; and partly from this circumstance he was led to believe
in the existence of a genetic relation subsisting between the two
kinds of bodies. He further expressed his conviction that the
people contracted the disease by washing themselves with water
in which gregarinaa abounded. Lindemann moreover refers to
Lebert as having noticed similar parasites in a case of favus,
and concludes that these organisms are of a vegetable nature.
His opinion, though not shared by the majority of parasitologists,
is nevertheless supported by the views of Robin, Ley dig, and
others. Of still higher interest are the observations of Linde-
mann respecting the occurrence of psorospermiaa in the capsule
of the kidney of a hospital patient who died with Bright's

280

PARASITES OF MAN

disease. The sacs in this case were remarkably small ; never-
theless their corpuscular contents indicated their true nature.
The pseudo-navicellae measured only in diameter. Amongst
other contributions of interest I may refer to those of Dufour,
J. Miiller, Creplin, Kolliker, Keferstein, Stein, Drumniond,
Lieberkiihn, and E. Ray Lankester. I doubt if the vegetable
organisms described by Prof. W. T. Gairdner can be referred
to this group of parasites. At all events, by whatever name
these spurious entozoa are called, they were first discovered by
Dufour in insects, by Miiller in fishes, by Miescher in the mouse,
by Dujardin in the mole, by Hessling in the larger quadrupeds,
and by (rubier in man. The results of my own examinations
may be briefly re-stated. In the flesh of cattle I found
psorospermial sacs varying from yL" to — " in length, and in
that of sheep from to ^" '. The bodies were enclosed in
well-defined transparent envelopes, and their contents exhibited
indications of segmentation. In some specimens the segments
displayed themselves as a complete cell-formation, the contents
of each cell being uniformly granular. Under the \" objective
the contained granules were clearly visible, and on rupturing
the sac their peculiar characters were at once manifest, each
granule or corpuscle represented a pseudo-navicel, all displaying
a tolerably uniform size, averaging in diameter. Some of
the corpuscles were round, others oval, several bluntly pointed
at one end, many curved and fusiform, not a few being almost
reniform. Highly refracting points or nucleoli were visible in
their anterior.

Turning to the practical aspect of the subject, I remarked
that these bodies had nothing to do with the cattle plague. No
one who carefully examined the flesh of animals that had died
of rinderpest had failed to discover them ; yet, in one or two
instances they appear to have escaped notice. When it is con-
sidered how long it takes us to examine a few grains weight of
muscle carefully, it is obvious that the body of a large beast
might contain many hundreds of these organisms without our
being able to detect their presence, except by a prolonged
investigation. In the few rinderpest beasts, portions of whose
flesh I submitted to the microscope, I should say there were
not more than 100 of these bodies in one ounce of meat; but in
the heart of a healthy sheep (which I afterwards ate) I calcu-
lated there were about 1000 parasites to the ounce, and in the
heart of a healthy bullock (which likewise served me for a

PROTOZOA

281

meal) their numbers were rather in excess of those in the sheep.
Altogether, at two meals, I could not have swallowed less than
18,000 of these psorosperms. Consumers of beef, mutton, and
pork eat these bodies every day, but they take no harm because
the parasites in question are not true helminths. Fine healthy
beef lias been returned to the butcher when it was as good as
any other meat in the market. I have examined various kinds
of meat, such as veal, pork, and mutton, but in none have I
found so great an abundance of psorosperms as in beef, which
was, notwithstanding, perfectly healthy and sound. I calculated
that in one instance a single ounce of the flesh contained upwards
of 2000 parasites. There is practically no limit to the extent
of this kind of parasitism, and there is no organ of the body in
which psorosperms may not be found. Moreover, the forms
they display are exceedingly various. Psorosperms have been
found by Siedamagrotzky in the muscles of the horse, and not
very long ago, through the help of Professors Simonds and Axe,
I had the opportunity to examine some peculiar worm-like
structures which occupied the mitral valve of a horse. To the
naked eye they looked like coiled nematodes, but I was soon con-
vinced that they formed a peculiar type of psorosperm. A
complete view of these bodies was a matter of great difficulty
owing to the delicate nature of their limiting membrane and to
the confusion of markings produced by the interlacing of the
fibres of the chordee tendinea. At length, by spreading a portion
of the membrane of the valve over a large glass slide, and
by allowing it to dry slowly, I found that the vermiform body
presented neither beginning nor end. The appearances were
curious and puzzling. The organism formed a flattened tube
or sac, almost uniform in width and variously twisted upon
itself. From the main tube there projected several hernia-like
secondary loops or branches, most of them presenting less
than half of the thickness of the former. These peculiarities,
however, can hardly be understood without reference to the
original illustrations. That these . secondary coils were not of
the nature of hernial protrusions was evident, not alone from
the nature of their contents, but also from the fact that they
showed distinct anastomoses. In fact, the parasite was a simple
sac or bag with branches.

On puncturing the main tube with a fine needle a small
quantity of tenacious creamy fluid made its escape. This,
under Ross's |-inch lens, resolved itself into a few excessivelv

282

PARASITES OP MAN

delicate sarcode globules surrounded by fine granules. The
granular matter displayed a tendency to collect itself in the
form of oval masses without showing any trace of a limiting
border. One of these masses, measuring gjj" in length, I
examined under a Wasserlein-objective, when I further ascer-
tained that the elementary particles or granules were uniformly
oval in shape, rather highly refractive, their size scarcely
exceeding ^m" ^n diameter. The sarcode corpuscles, on the
other hand, were of different sizes, ranging between ~' and
•JB1o5// in diameter.

From the facts thus elicited, negative as they were in
respect of helminthic structure, I could see no escape from the
conclusion that we had to deal with a new form of psorosper-
mial bag, whose granular contents consisted of excessively
minute pseudo-navicellas. In the centre of the largest hernia-
like loop there was a clear oval disk, which at first brought to
my mind the nucleus of Monocystis infesting the earth-worm,
but it was merely a vacuole.

The case recorded by Gubler reminds me of another remark-
able instance of psorospermial cysts, in this case associated
with true hydatids. In 1873 Dr Whittell sent me particulars
of a case in which the contents of an hydatid of the liver
(drawn off, during life) consisted of shreds of a true hydatid,
a few echinococcus-hooklets, together with multitudes of
spindle-shaped amoeboid particles of excessive minuteness and
delicacy. The bodies, floating in a transparent fluid, formed a
thick milky or creamy fluid, resembling pus in appearance ; but
there was no trace of pyaemia. Judging from Dr WhittelPs
figures, he must also have found a solitary microscopic nematoid
hsematozoon, the nature of which was not clear to him. I
believe it to have been a specimen of Filaria sanguinis hominis.
Be that as it may, the case is altogether unique and deserves
further elucidation.

As regards the higher forms of protozoa it must suffice to
allude to the Oercomonas hominis of Davaine, found in the
dejections of cholera patients, to the Gere, urinarius of Hassal
and G. saltans of Bhrenberg, to the Trichomonas vaginalis of
Donne, detected in the vaginal mucus, and to the Balantidium
coli of Claparede and Lachmann, originally found by Malmsteu
in the human colon. The Balantidium, or Paramecium coli,
has frequently been observed in the evacuations of fever patients,
and it has also been found by Dr Treille in patients suffering

PKOTOZOA

283

from the Cochin-China diarrhoea. Monads have also' been
found in the stomach and intestines of the hog and various
other animals. Infusorial parasites are particularly abundant
in batrachians, the Bursarim of frogs and toads being familiar
to every helminthologist.

Bibliogeapht (No. 41). — Arloing (and Trvpier), in ' Gaz.-
hebd./ 1873, p. 574 (quoted by Davaine). — Balbiani, ' Compt.
Rend. Soc. Biol./ 1867, p. 103 (quoted by Davaine and Bastian).
— Bastian, " On the Nature of the so-called Sarcina ventriculi,"
' Brit. Med. Journ./ Feb. 3, 1872. — Idem, " On Hefcerogenesis
in its relation to certain Parasitic Diseases/' ' Brit. Med. Journ./
'Feb. 24 and April 20, 1872 (see part iv, p. 417, with figs, from
Balbiani). — Beale, L., " Entozoon-like bodies in Muscles/' in the
' Microscope in Medicine/ 4th edit., p. 485, 1878. — Idem,
"Bacterium Hypothesis of Contagium," ibid., pp. 313—321. —
Burnett, W. T., " On Psorospermia, Mermithes, &c," in a paper
entitled 'Reviews and Records in Anat. and Physiol./ in ' Amer.
Journ. of Sci. and Arts/ vol. xviii, 2nd ser., p. 104, 1854. —
Garter, H. V., " On Spirilla/' quoted by Sanderson in ' Brit. Med.
Journ./ Nov. 17, 1877, p. 700. — Gobbold, " Remarks on Spurious
Eutozoa found in Diseased and Healthy Cattle," ' Path. Soc.
Trans./ vol. xvii, p. 452, 1866, and 'Lancet/ Jan. 27, 1866,
p. 88 ; see also Prof. J. Gamgee's work on the ' Cattle Plague.'
Idem, " On Worm-like Organisms in the Mitral Yalve of a
Horse," 'Veterinarian/ Sept., 1877. — Idem, "On Psorospermiae
in the Eye of the Cod (Morrhua)," ' Linnean Society's Proc./
May, 1862, and in 'Intellectual Observer/ 1862, p. 199.— Gohn,
'Nova Acta/ xxiv, s. 103 (quoted by Leuckart), Bd. i, s. 139.
— Crejplin, ' Wiegmann's Archiv/ 1842, s. 61. — Davaine, 1. c,
2nd edit., " Synops. xxi " (with bibliog. refs.), 1878. —
Donne, ' Cours de Microscopie/ Paris, 1847, p. 157. — Dressier,
quoted by Leuckart, Bd. i, s. 141. — Drummond, ' Edin. Phys.
Rep., 1852, — Dvfour, 'Ann. des Sci. Nat./ 1837. — Dujardin,
'Traite' (1. c, see Bibl. No. l).—Eberth, ' Zur Kentniss Bac-
teritischer Mykosen/ 1872. — Dimer, ' Ueber Psorospermien/
1870.— Gairdner, 'Edin. Phys. Soc. Rep./ 1853.— Gluge, "Cysts
in Sticklebacks," 'Bullet. Acad. Roy. des Sci. de Bruxelles/
1838.— GWer, 'Mem. Soc. Biol./ 1859, p. 657, and in ' Gaz.
Med./ 1858, p. 61.— Earz, G. 0., " Eine neue Mikrococcusform
im lebenden Thierkorper," ' Deutsche Zeitschrift fur Thier-

Medicin und vergleichende Pathologie/ f. Novemb., 1878.

Hcssling, 'Sieb. u. Koll. Zeitsch./ 1853, p. 196. — Eenle, ' Miiller's

281.

PARASITES OP MAN

Archiv/ 1845.— Hollis, W. A., "What is a Bacterium?" repr.
in the ' Veterinarian/ p. 205, 1875. — Kcferstein,' Q'6t\mg. gelehrte
Anzeigen/ 1862. — Kloss, ' Ueber Parasiten (u. s. w.) ' (quoted
by Davaine). — Knock, 'Journ. de Russ. .Kriegs. dep./ Bd. xev,
1866 (quoted by Leuckart and by Davaine). — Kblliker, in
'Zeitsch.' (by Sieb. and Koll.), 1848-49.— Lambl, ' Prager
Vierteljahrschrift/ 1859. — Lanlcester, E. R., " Recent Researches
on Bacteria (with copious references)/' ' Quart. Journ. Micr.
Science/ Oct., 1878.— Lebert, ' Phys. Pathologique ' (quoted by
Leuckart). — Leidy, " Gregarina/' 'Amer. Phil. Trans./ 1851. —
Leisering, " Bericht (u. s. w.)," in ' Sachsen/ 1865. — Leuckart,
1. c, Bd. i, s. 135 and 740, and Bd. ii, s. 842 et seg. — Leydig,
' Miiller' s Archiv/ 1851, s. 221, in 'Micr. Journ./ 1853, p. 206,
and in ' Arch. f. Anat. und Phys./ 1863, s. 191. — Lieberleuhn,
' Mulleins Arch./ 1854. — Lin demann, ' Bullet. Soc. imp. des
Naturalistes de Moscow/ 1863, and in ' Gaz. Med. de Paris/
1870, p. 86.— Lister, J., "Natural History of Bacteria," 'Micr.
Journ./ Oct, 1873. — Malmsten, " Paramecium coli" (quoted
by Davaine, 1. c, 2nd edit., p. 67). — Miescher (quoted by
Leuckart and Siebold). — Miiller, 'Archiv/ 1841, s. 477. —
Rainey, 'Phil. Trans./ 1857. — Bayer, " Singuliere eruption sur
un veron (Cyprinus)/' 'Arch, de Med. Comparee/ Paris, 1842
(quoted by Davaine). — Rivolta, " Psorospermi, &c," trans, in
'Journ. des Vet. du Midi/ 1869, pp. 445 and 521.— Robin, ' Les
Veget. Paras./ 2nd edit., p. 291. — Sanderson, in 'Privy Council
Reports/ 1874. — Siedamagrotzky, in 'Recueil de Med. Vet./
1872, p. 460— Stein, in ' Miiller' s Arch./ 1848, and 'Ann. Nat.
Hist./ 1850. — Ldem, "Abhandl. d. k. Bohmischen Gesellsch./'
x, s. 69, oder Lotos, 1859, s. 57 (quoted by Leuckart, Bd. i, s.
151). — Steinberg, ' Walter's Zeitschr. f. die moderne Medicin/
1862, and in Leuckart, Bd. ii, s. 844. — Stieda, 'Arch. f. pathol.
anat./ Bd. xxxv, and in Leuckart, Bd. ii, s. 846. — Suriray, " Sur
quelques parasites du lombric," 'Ann. des." Sci. Nat./ 1836. —
Virchow, " Zur Kentniss der Wurmknoten," 'Arch. f. Anat. u.
Phys./ xviii, s. 523. — Vogel, ' Path. Anat./ i, s. 404. — Walden-
burg, " Psorospermien," in ' Arch. f. Path. Anat./ s. 435, 1867. —
Windbladh, also Wising, ' On Balantidium coli ' (quoted by
Leuckart, Bd. ii, s. 846-847). — Winkler (see Leisering).

Appendix. — On various occasions I have dwelt upon the
necessity of acquiring accurate information respecting the degree
of mortality due to parasites, and in the present volume (p. 121)

APPENDIX

285

I have referred to the defective evidences supplied by the returns
of the Registrar- General in respect of the echinococcus disease.
My object is not to cast blame upon those whose duty it is to
publish the returns, but rather to call attention to the advan-
tages that would follow if the Registrar- General were supplied
with full and accurate information on this head.

Through the courtesy of Mr Noel A. Humphreys I have been
furnished with the following official statement of the number of
Deaths from Worms in England and Wales, as recorded in the
Auuual Reports of the Registrar- General throughout a decade
of years :

1868

1869

1870

1871

1872

1873

1874

1875 1876

1877

Total .

Including —
Ponigo

Scabies .

Tapeworm

Hydatids

172

148

151

160

154

183

188

227

204

225

15
6
8

13
2
3

9
7
6
33

9
1
3
37

9
4
5
41

14
2
3

5
29

16
5
5

7
2
2
31

13
3
6

Considering the prodigious advances in helminthology during
the last half century, it is certainly remarkable that under the
category of " worms/' as a cause of death, only two kinds of
true helminths should be mentioned in the Registrar's record.
It will also strike the experienced hospital and dispensary
physician as somewhat remarkable that of the two death-pro-
ducing parasites above named one of them should be the " tape-
worm." Now death from Ttenia is certainly a very rare occur-
rence, although grave nervous symptoms are not unfrequently
due to its presence in man. Thus, I am inclined to regard the
46 reported instances of death from this cause as a redundant
estimate. On the other hand, I am surprised to see no specified
instances of death from lumbricoid Ascarides, from Oxyurides,
or even from Cysticerci, which now and then take up their
residence in the human brain.

As regards hydatids I believe the returns to be excessively
deficient. In place of an average of 34 deaths annually from
this cause in the United Kingdom I am of opinion that at least
400 deaths are due to hydatids. This opinion and the data on
which it was founded were communicated by me twelve years
ago to the Linnean Society, and I have since become acquainted
with facts which lead me to conclude that my original estimato
was very much below the mark. The post-mortem registrar

286

PARASITES OF MAN

of one of our large hospitals has told me that of late years as
many as ten deaths might be reckoned as annually due to
hydatids in their institution alone. At a smaller hospital I
ascertained that the average was about four. Obviously, if these
estimates are correct, the Registrar- General's returns for the
United Kingdom do not record a tithe of the annual mortality
due to hydatids. Perhaps another half century will elapse
before the truth of my deductions be confirmed by the official
returns. For me, it must suffice to have pointed to the
desirability of securing more accurate records.

By a curious coincidence I had only just sent to press the
sheet of this work recording the statistics of hydatid disease in
Australia (p. 123), when a paper dealing with the same subject
appeared in the ' Lancet/ I refer to the brief memoir of Dr
David Thomas, of Adelaide, South Australia, which was published
on the 1st of March, 1879. Dr Thomas writes as follows :

" It is well known that Australia presents an extraordinary
prevalence of hydatid disease, but, as far as I know, no definite
statistics have been published to illustrate the fact. Conse-
quently, some months ago, with the kind assistance of the Hon.
W. Morgan, the present Chief Secretary of South Australia, I
endeavoured to procure reports from the Governments of
Victoria, New South Wales, Queensland, New Zealand, Tasmania,
and Western Australia, upon this question. Unfortunately, the
mode of registration of the causes of death in most of these
colonies was such as prevented the necessary replies being
supplied. However, it appears that in Tasmania no deaths
were returned from this disease in the ten years 1867-77.
During the greater part of the same period no separate classi-
fication of hydatid disease had been adopted in New South
Wales; but in 1875 four deaths were attributed to hydatids ; in
1876, eleven were so returned. In Victoria, however, the
record of deaths from this cause is far more complete, and I
append a table based upon the returns from that colony.

" One case, in which hydatid of the kidney was present,
accompanied by malignant disease of bladder, with stricture,
was not returned as a case of hydatid causing death.

"During the decade 1867-77, 2*5 per 1000 deaths were due
to hydatid disease. In 183 out of the 307 cases the liver was
either solely or conjointly with other organs the seat of disease.
Holding the second place in frequency come the lungs in 71
cases, i.e. 53 simple and 18 complicated."

APPENDIX

287

Table of Deaths returned as being due to Kijdatid Disease in the
Province of Victoria during the ten years 1867-77.

Year.

Liver.

Lungs and
pleura

Brain and
spinal cord.

Heart and
Pericardium.

Kidney.

Spleen.

Pancreas.

Omentum.

Ovary.

Abdominal
cavity.

Situation not
recorded.

More than one
organ invaded.

Total annual
deaths.

1868
1869
1870
1871
1872
1873
1874
1875
1876
1877

17
6
10
4
19
17
21
29
23
20

5
8
2
6
3
3
10
7
1
8

1
1
1
1

2
1

i
i

2
2

3
1

7
1

4
1
2
2
6
2
4
2

1(a)
3(6)

1(c)

5(e)
2(/)

2(50
5(A)
2(i)

33
22
17
15
29
29
41
47
36
38

Totals

166

307

in lOyrs

(a) Lungs and kidneys.

(4) In two instances liver and lungs; in the third case liver and brain.

(c) Lungs and liver. (d) Lungs and abdominal cavity.
\e) In three cases liver and lungs; in one kidney aud abdomen; in another liver, lungs, omentum.
(/) Liver and lungs.

(ff) One liver and lungs; the second case lungs and heart.

(h) Three cases liver and lungs; one liver and kidney. (i) Liver and lungs.

Such are the returns as recorded by Dr Thomas. If a com-
parison be instituted between the data supplied by his decade-
report and those supplied by the decade-report which I have
previously adduced (p. 123), it will be seen that as regards the
returns for the years 1868-72, inclusive, both reports are in
perfect agreement. Of still more interest also is the circum-
stance that whilst, on the one hand, out of the total of 307
deaths given in Dr Thomas's table, 116 occurred during the
first semi-decade {i.e. from 1868 to 1872, inclusive), on the other
hand, no less than 181 deaths occurred during the second semi-
decade (i.e. from 1873 to 1877, inclusive). This increase of 45
deaths during the later semi-decade is very significant. It
points either to the fact of more careful returns having been
made, or to an actual increase in the fatality of the disorder.
Possibly both the causes alluded to operated to affect the
returns. Be that as it may, Dr Thomas's record is highly
instructive, and should stimulate the profession in England to
supply our Registrar- General with more precise data wherewith
to construct his annual reports.

BOOK II

PAEASITES OF ANIMALS.

In dealing with this division of the subject it will be impos-
sible to give more than the faintest outline. Let it be borne
in mind that quite as much information might be offered by me
respecting the parasitism of each of the commoner domestic
animals as has been already advanced in regard to the helmin-
thism of man. That would by no means exhaust the subject.
Thus treated, a score of volumes, each equal in size to this, would
barely suffice to cover the whole ground of Parasitology ; and yet
there are not wanting intelligent persons who regard Helmin-
thology as unworthy of their attention. These persons form a
far too prevalent type of educated ignorance, and unfortunately,
it is just this class of people who enjoy the prerogative of edu-
cational responsibility. Even our metropolitan scientific insti-
tutions, expressly raised for the purpose of diffusing useful
knowledge, shrink from the revelations of parasitism. Domi-
nated by the miserable conception which judges that the
life-history of a worm cannot prove either interesting or
instructive to their audiences, they let slip the acquisition of
scientific data, a knowledge of which might enable them to
combat successfully some of the most terrible evils to which
human flesh, in common with that of animals, is heir.

In the following few pages many new points of departure for
fresh scientific work will inevitably be suggested ; and if I only
succeed in conveying to the working student an adequate grasp
of the whole subject, especially in its bearings on the welfare
of the higher domesticated animals and man, I shall have
accomplished all that I can reasonably hope to do within the
restricted limits of space at my command.

QUADRUMANA

289

SECTION I (Mammalia).

In dealing with this class of hosts, exclusive of man, I shall
notice the entozoa of the various orders successively, taking
the arrangement which I employed many years since when
writing the Mammalian Division of a popular treatise called the
'Museum of Natural History/ The internal parasites of those
orders which happen to embrace important domesticated
animals will necessarily receive more attention than the others;
some notice of the ectozoa being likewise added.

Paet I. — Quad rum an a (Monkeys).

Monkeys are less afflicted with flukes than most animals.
The species known to me are Distoma laciniatum, found by
Brongniart in the pancreas of Simia maimon ; D. orbiculare and
Amphistoma emarginatum, from the intestines of Gebus trivir-
gatus ; and Bilharzia JuEmatobia, a single specimen of which I
obtained from, Gercopithecus fuliginosus. The monkeys of the
Old World rarely harbor full-grown tapeworms, but Cysticerci
are abundant (Cyst, tenuicollis, 0. cellulosa, G. pileatus, G.
crispus). The common hydatid {Echinococcus polymorphus) has
been found in many of the Simiae, and by myself in a
Madagascar lemur (L. macaco). Dr Leidy also obtained three
hydatid cysts from a large monkey. On the last day of the
year 1857 I obtained some polycephalous hydatids (Gcenurus
lemuris) from a ring-tailed lemur. They infested the liver,
being more abundant in the lungs. They occupied both sides
of the chest. Loose and detached specimens also existed in the
cavity of the right pleura. Most of those occupying the chest
were connected to the pleura, forming vesicular, semi-transparent
masses, varying in size from a filbert to a large walnut, many
being united in bundles of much larger size. One of these
masses is here drawn (1, fig. 54). It consists of four large
Coenuri, their combined pedicles forming a single stalk. A fifth
hour-glass-shaped rudimentary Ccenurus is also visible. Every
Gcenurus supports a variable number of lobules, each lobe sup-
porting one or more papillae. Here and there the papillae
resemble chains of beads. No trace of tapeworm heads could
be seen, but under a half-inch objective glass, I found some
flat papillae presenting oval depressions at the surface (2, fig. 54).

290

PARASITES OF ANIMALS

On examining some of the larger papilla? they were found to
consist of membranous layers folded one within the other
These were carefully dissected and examined with the aid of
needles, when each one showed in the centre a well-formed

Fig. 54. — Comurus lemuris. 1, Coleny; 2, portion of the ectocyst (enlarged); 3, scoler-
heud (magnified 40 diameters); 4, liouks (magnified 260 diameters). Original.

tapeworm head with four characteristic suckers, and a promi-
nent rostellum supporting a double coronet of hooks, thirty-two
of the latter in all (3, fig. 54). The hooks displayed a marked
disparity of size and form. Each hook showed conspicuous
anterior and posterior root-processes, the larger set of hooks
individually measuring about 5±5" and the smaller in length

QUADRUMANA

291

(4, fio-. 54). There were numerous calcareous corpuscles. The
interior of each vesicle was filled with a fluid, in which there were
no free scolices. On referring to my notes I find that the lemur
had arrived in England about four months previously.

Larval cestodes do not appear to be common in the monkeys
of the New World (Cebidae), nevertheless I found several
Cysticerci in the liver of Macacus radiatus (Feb. 19th 1857),
and a single specimen in the sooty monkey (Dec. 4th 1857).
They were wrongly described by me as Cercariae. The
Cebidas are largely infested with tapeworms {Taenia megastoma
and T. rugosa). A species of Ligula {L. reptans) has likewise
been found beneath the skin of Callithrix sciureus and in
one of the marmosets {Hapale melanurus) . Perhaps the
most common helminth infesting monkeys is the nematode
called Filaria gracilis. I have examined specimens from
the orang, the capuchin, and the spider monkey. This parasite
commonly occupies the abdomen, coiled beneath the perito-
neum, or within folds of the mesentery. It sometimes occurs
beneath the skin, or within the connective tissue of super-
ficial muscles. The female worm has been known to reach
a length of five feet. In 1873 Mr Samuel Smith, of Clifton,
sent me five specimens of this worm. From one of the males,
which measured twenty inches in length, I procured some
spermatozoa, and found their long diameter to average 75~/'.
These corpuscles and other structures, as well as the worm
itself, are figured in my ' Notes on Entozoa ' quoted below,
Next in frequency, perhaps, is the whipworm {Tricocephalus
dispar), which monkeys of all kinds harbor in common with
man. Besides these nematodes, Physaloptera dilatata is found
in the stomach of American monkeys, and Ascaris distans also
(in the large intestine of marmosets more particularly). This
Ascaris has also been found in Gercopithecus fuliginosus and in
Simia sabaea. A small spiroptera is said to infest the walls of
the stomach of Simia maimon. To Dr Murie I am indebted
for a large roundworm {Ascaris lumbricoides) taken from the
intestine of a chimpanzee {Troglodytes niger), and also for a
smaller nematode taken from a green mona-monkey {Gercopi-
thecus). This I have described and named Ascaris cuspidata.
From the intestines of a chacma {Oynocephalus porcarius) M.
Schafhert procured sixteen examples of a small strongyle {Str.
attenuatus, Leidy).

The singular monkey known as Tarsius spectrum is liable

292

PARASITES OP ANIMALS

to be attacked by a filar ia (F. la>vis), which is found beneath the
skin. At least two species of Echinorhynchus (E. spirula and
F. elegans) are known to infest monkeys. 1 have carefully
examined and figured F. elegans in the ' Zoological Society's
Proceedings,'' from specimens supplied to me by Dr Murie.
They had been obtained from a pinche- monkey (Hajpale oedipus)
from New Granada. I am under the impression that Diesing' s
original description of this parasite is the only one that exists.
I have gone over his numerous memoirs contributed to the
Vienna Academy, but can find nothing beyond the specific
characters given in his ' Systema.' All the specimens in the
Vienna Museum, whence his description is taken, were collected
by Natterer. They were procured from the marakina (Midas
rosalia), from two other true marmosets (Ha-pale Ursula and
Ft. chrysoleuca), and from a squirrel monkey or tee-tee (Galli-
thrix sciureus). In the monkey which died at the Zoological
Society's Gardens the cause of death was not clearly due to
the parasites ; nevertheless, the mucous layer of the intestine,
to which the entozoa were attached, showed deep conical pits or
depressions at the spots where the worms had anchored them-
selves. During the perfect retraction of the proboscis of this
Echinorhynchus the centre is represented by a wide opening
which communicates with a cavity beneath. The end of the
neck thus forms a sort of collar, or rosette, made up of rays
arranged like the spokes of a wheel. When the proboscis is
exserted this collar is more or less convex, but it becomes
slightly concave when the proboscis is retracted. Diesing
recognised twenty-four rays ; they -probably vary from that
number up to twenty-eight, at least I counted twenty-seven in
my specimens. During exsertion the proboscis forms a nipple-
like projection. According to Diesing it supports three rows
of hooks, but I certainly saw four rows. When separately
magnified these hooks present very different appearances as to
size and contour. These variations I believe to be due to
growth. The largest hooks measured about in length.

As regards insect parasites, it is well known that monkeys
are largely infested by fleas, but the species have not been much
studied. The same may be said of their mites (Acaridat). A
species of Pentastoma (P. tomatum, Creplin) has been found
occupying little cysts of the peritoneum and omentum in Simia
maimon and 8. cynomulgus. Under a synonym (Linguatula
Biesingii) it has been very fully described by Van Beneden.

OIlEIROPTlfiHA

293

Another species (P. subcylindricum) lias been found attached to
the lungs and liver of a marmoset (Hapale chrysopygus). Dr
Leidy found five specimens of P. euryzonum in cysts on the
subperitoneal surface of the liver of Gynocephalus porcarius.
Leuckart believes that Dr George Harley's P. polygonum is the
adult state of P. euryzonum, and that P. subcylindricum is the
larva of P. proboscideum, found in Boa constrictor and other
serpents.

Bibliography (No. 42). — Beneden (see Van Beneden). —
Gobbold, "On Filaria gracilis and Ascaris cuspidata," in
'Zool. Soc. Proc./ Feb. 3, 1874, p. 124.— Idem, "List of
Entozoa, including Pentastomes, obtained from Animals dying
at the Menagerie in 1857-60 inclusive/' ' Proc. Zool. Soc./ 1861 .
— Idem, 'Entozoa/ p. 119. — Idem, "On Parasite-Larvas (Ccenurus
and Echinococcus of the Lemur)/' ' Intell. Observer/ vol. iii,
pp. 86-96. — Idem, " Bilharzia from Cercopithecus," in ' Synops.
of Distomidse/ ' Proc. Linn. Soc./ 1860, Zool. Div., pt. v, p. 31,
and in " Parasites from Zool. Gardens./' in ' Intell. Obs./ June,
1862, p. 352. — Idem, " On Echinorh. elegans," ' Proc. Zool. Soc./
1876, in ' Notes on Entozoa/ pt. iii, plate xvi, p. 202. — Creplin,
"On P. tomatum," 'Troscbel's Arch./ 1849, s. 54. — Idem,
" F. gracilis/' in 'Wiegm. Arch./ 1851, s. 270. — Diesing,
'Syst./ 1. c, s. 611, and in ' Sitzb. d. k. Akad./ 1854, s. 598,
—Karley, G., in ' Proc. Zool. Soc./ 1857, p. 12.— Leidy, J., " On
Pentastoma," ' Proc. Phil. Acad. Nat. Sci./ 1850, p. 97. — Idem,
' Hydatids/ 1. c, 1856, p. 46. — Idem, c Strong, atten./ 1. c,
p. 54. — Leuckart, ' Bau und Entwicklungsgeschichte der Pentas-
tomen/ 1860, "P. polygonum," ' Synops./ s. 153. — Molin, " On
F. gracilis," in ' Sitzungsb. d. k. Akad. d. Wissensch./ 1858,
Bd. xxviii, s. 376. — Schneider, ' Monag. der Nem./ 1866, " F.
gracilis," s. 87. — Smith, S., " On F. gracilis found in connec-
tion with the Great Omentum of a Spider-monkey/' ' Proc. of
Bristol Naturalists' Soc.,' vol. i, 1873. — Van Beneden, " Ling.
Diesingti," in ' Mem. Acad. Belgique/ vol. xxiii, ' Recherches
sur 1' organisation et le development des Linguatules, suivies de
la description d'une espece nouvelle provenant d'un Mandrill/
1849.

Part II (Cheiroptera).

Until lately not very much attention had been paid to the
parasites of bats, probably on account of the insignificanco of

201.

PARASITES OF ANIMALS

the hosts. However, whilst Dr Dobson has recently been
extending the subject of Cheiropterology, Prof, van Beneden
has added largely to our knowledge of the parasites of bats.
In his beautiful memoir, the Belgian helminthologist asks
whether the parasites quit their hosts during the period of
hybernation, and then proceeds to answer that question in the
negative. Should the bats die, the parasites of necessity share
the same fate. It would appear, however, that the spermatozoa
of the worms are capable of surviving their parents for a
fortnight or even longer. Flukes abound ; the most common
species {Distoma lima) infesting the pipistrelle, noctule, mouse-
colored bat, batterer's bat, parti- colored bat, Daubenton's bat,
whiskered bat, and the greater and lesser horseshoe bats.
Almost as abundant is the Distoma chilostomum found in most
of these bats, and also in Leisler's or the hairy- armed bat.
Dujardin's D. heteroporum is a synonym of the species. Schre-
ber's Monostomum is probably identical with Van Beneden's
Distoma ascidia. This fluke he found in Vespertilio marinus,
V. dasycnemus, V. Daubentonii, V. emarginatus, V. serotinus,
V. mystacinus, V. pipistrellus , V. auritus, and in Rhinolophus
hippocrepis. Another species, distinguished from D. ascidia by
its large ventral sucker, has been found in great numbers in
the noctule (D. ascidio'ides, Van Beneden). The cestodes of
bats are not numerous. The best known species (Tania obtu-
sata) has been found in the serotine, in the mouse-colored bat,
and in Vespertilio lasiurus. Another species (T. decipiens) occurs
in Molossus peroiis, and Ohylonycteris rubiginosus, and a scolex,
forming the type of a new genus and species (Milina grisea,
Van Ben.) has been obtained by hundreds in the intestines of
Vesp. murinus and V. serotinus. Of nematodes we have Ophio-
stomum mucronatum, Bud., and Oph. spinosum, W.-Suhm (from
Vespertilio mystacinus), Tricliosomum specio sum }Ya,n. Ben., Stron-
gylus tipula, Van Ben., Strongylacantha glycyrrhiza, Van Ben.,
Litosoma filaria, Van Ben., Ascarops minuta, Van Ben., and
one or two sexually-immature forms, either found loose in the
tissues or occupying cysts. The Acanthocephala are not known
to infest bats. Bespecting insects, one family (Nycteribiidae)
is exclusively parasitic upon bats. They resemble the forest-
flies in their habits. The best known species is Nyct&ribia
Latreillei, but several others (N. biarticulata and N. Syhesii,
Westw.) have been described. One or more of the Brazilian
bats are infested by Lipoptena phyllostomatis, Nitsch, and

INSEOTIVORA

295

West wood has obtained several allied forms of Hipposcida
(of the genus Strebla), also from bats. As regards the true
Araclmidans (mites and ticks), Van Beneden enumerates Ixodes
lividus, Van Ben., Pteroptus vespertilionis and P. arcuatus,
Koch, Otonissus aurantiacus and Oerathophyllus octactenus, both
of Kolenati, and Caris elliptica.

Bibliography (No. 43) —Audouin, 'Ann. des Sci. Nat./ xxv.
— Beneden, P. J. van, " Les Parasites des Chauves-souris de
Belgique/' fr. vol. xl of the ' Mem. de FAcad./ 1873.— Diesing,
'Syst. Helm./ ii, p. 530.—Dufour, ' Ann. des Sci. Nat./ 1831.
—Dujardin, 1. c, p. 437. — Kolenati, ' Die Parasiten der Chirop-
teren/ 1857. — Latreille, art. " Nycteribie," in 'Nouv. Diet.
d'Inst. nat.' — Miiller, ' Zool. Danica/ ii, p. 43, " Fasciola vesp.
(syn. Dist. lima)." — Perty, " Lipoptena/' 'Del. an. art. BrasiP
(quoted by Westwood). — Budolphi, ' Synops./ p. 117. — TFestf-
wood, "Mem./' in ' Zool. Soc. Trans./ 1835.— Idem, 'Modern
Classif. of Insects/ vol. ii, p. 585, 1840. — Willemoes-Suhm,
B. V., " Helminthologische Notizen," iii, ' Zeitsch. f . wiss.
Zool./ 1873.

Paet III (Insectivoea) .

The entozoa of insectivorous mammals, though sufficiently
numerous, are not important practically. The common hedge-
hog {Erinaceus europceus) is infested by four flukes (Distoma
pusillum, J), trigonocephalum, D. caudatum, D. linguarforme) ,
and also by three thorn-headed worms (Echinorhynchus napee-
formis, E. amphipachus, and E. major). Two tapeworms are
also known (Tania compacta and T. tripunctata) . More atten-
tion has been paid to the round worms. A species of strongyle
(S. striatus) infests the lungs, the male being readily distin-
guished by its nearly round hood. A second species of stron-
gyle has been mentioned by Diesing, but it is more than
doubtful. The lungs are also infested by a small trichosome
(Eucolens tenuis of Dujardin) ; another species of the genus
(Trich. exiguum) infesting the stomach and small intestine. As
the Trichina spiralis has been repeatedly reared by myself and
others in the hedgehog, the little flesh-worm must also be
noticed in this place. Physaloptera clausa occupies the
stomach, and a minute Ascaris (A. pusilla) is found in cysts of
the peritoneum. According to Wedl, the intestine of the
Egyptian hedgehog {Erinaceus auritus) is infested by another

29G

PAliASlTBS OP ANIMALS

worm, which he calls Pterygodermatitis plagiostoma. This is
allied to Froelich's genus Rictularia. As regards the mole
(Talpa europaa), two flukes have been described (Distomum
fiexuosum and Monostomum ocreatum), also two round worms,
namely, Ascaris incisa, occupying the peritoneum, and Spiro-
ptera strumosa in the cavity of the stomach. Dr Schneider
places the latter with the Filarise. A little tapeworm (Tania
bacillaris) infests the small intestines, and a larval cestode
occupies the liver and subcutaneous connective tissues. This is
the well-known Gysticercus talpce, which Leuckart and others
have referred to as being the scolex or juvenile state of Tania
tenuicollis infesting weasles (Mustelidce) . The Cysticercus is
also found mArvicola arvalis. In regard to the shrews, many
species of fluke have been described as occupying the intes-
tines. In Sorex araneus and S. leucodon, the Distoma migrans;
in S. constrictus, the D. exasperatum ; in S. tetragonurus, the
D. corrugatum and D. rub ens ; the last-named fluke, with two
others (D. instabile and D. truncatum), being also found in
Daubenton's shrew. The tapeworms are numerous — Tania
neglecta, T. furcata, T. uncinata, T. pistillum, T. tiara, T.
scalaris, T. scutigera. With the exception of the last named,
all these forms occur in the common shrew. According to the
investigations of M. Villot the cysticercal stage of T. pistillum
is to be found in the glow-worm (Glomeris). This scolex
(Staphylocystis micr acanthus, Yillot) multiplies by proliferation,
and in this way the swallowing of a single intermediate host
may result in the formation of a hundred or more tapeworms.
Another species of Staphylocystis (8. biliarius) is considered by
Yillot to be the larval source of T. scutigera and T. scalaris,
which are perhaps identical species. A small Echinorhynchus
(jE7. appendiculatus) , found in the intestines and also encysted in
the mesentery of the shrew, in like manner becomes transferred
to the stomach of the fox. The nematodes of shrews possess
little interest. In the common shrew the only species known
are Trichosoma splenaceum and an immature worm, whilst in
Sorex tetragonurus we have T. incrassatum, occupying the
tunica vaginalis of the testis, and Strongylus depressus in the
intestines. Not many other insectivora appear to have been
studied in relation to their internal parasites. A larval cestode
has been noticed in the Russian musk rat {My gale), and fIso
a tapeworm [Taenia sphairocephala) in the golden mole (Ghryso-
chloris). Several flukes and a tapeworm have been found in

OARNIVOJJA

297

the water-shrews (Sorex fodiens), but, so far as I am aware,
nothing has been done in connection with the parasites of the
Macroscelidina, of the Banxrangs (Tupaina)., or of the Tanecs
(Centites) and their allies. The entozoa of the star-nosed and .
shrew moles of North America (Condylura and ScaJops) also
deserve attention. From the last-named genus ($. canadensis)
Prof. Leidy obtained a single male spiroptera. It occupied the
stomach and was only half an inch in length.

Bibliography (No. 44). — Leidy, " S. scalopsis canadensis/'
fProc. Phil. Acad./ 1851, p. 156. — Linstow, " D. cordatum,
Einigeneue Distomen (u. s. w.)," ' Arch, fur Anat./ 1873, s. 95.
— Molin, 'Una Monografia del genere Physaloptera,/ Wien,
1860 (p. 7, " P. cla,usa," and p. 31, " P. limbata"). — Idem,
' Nuovi myzelmintha/ Wien, 1859, p. 10, Spec. No. 8. — Idem,
rUna Monogr. del genere Spiroptera/ Wien, 1860, p. 25, Spec.
No. 22. — Schneider, ' Monogr. der Nematoden/ Berlin, 1866,
s. 103, Spec. No. 39.— Stieda, in ' Troschel's Archiv/ 1862,
" Description and figs, of Taenia uncinata and T. furcata of the
Shrew/'' — Thomson, art. " Ovum/' in ' Todd's Cyclop, of Anat.
and Phys./ contains figs, and description of T. pistillum (from
Dujardin), vol. v, p. 28". — Villot, A., " On the Migrations and
Metamorphoses of the Tapeworms of the Shrews/' in e Ann. of
Nat. Hist./ March, 1878, from ' Comptes Rendus/ Nov. 19,
1877, p. 971.— Wedl, K., " Zur Helminthenfauna ^Egyptens,"
' Sitzungsb. d. math.-naturw. Classe ' (u. s. w.), Bd. xliv,
Abth. i, s. 464.

Paet IV (Caenivoba).

Notwithstanding the importance of the entozoa of this large
section of mammals, I must deal with them very summarily,
emphasising my remarks on the parasites of the dog and cat.
Only a few of the ectozoa can be noticed.

The bears are much infested by nematodes, the species being
Ascaris transfuga and Spiroptera (Gongylonema) contorta of
Molin. The latter is found in the oesophagus. Immature
round worms have also been found in cysts. These were
erroneously described- as cestodes by Zeder and Gmelin. The
bear, however, is very liable to be infested by genuine Cysticerci.
Retzius found them in the muscles, and they are described as
examples of the ordinary hog-measle. The museum attached

208

PARASITES OF ANIMALS

to Guy's Hospital contains the heart of a bear which is largely-
infested by Cysticerci. A species of tapeworm has been found
in the polar bear. The coati {Nasua narica) is infested by
Ascaris brachyoptera in the intestine, by Molin's Phy*aloptera
semilanceolata from the stomach, by Echinorhynchus spnrula,
Tcenia crassipora, and Ligula reptans, the latter occupying the
muscles. A second species of Ascaris (A. alienata) is described
from Nasua rufa.

The racoons (Procyon) are infested by a species of pentastome
(P. subcylindricum) , and Prof. Leidy has described a threadworm

(Filaria insignis) obtained from a
cyst in the foot. The gluttons
(Gulo) are liable to be infested by
an Ascaris, a Ligula, and by
Eustrongylus gigas. The giant
strongyle also infests the coati
(Nasua). The skunk harbors
Tcenia crassipora. The Hunterian
Museum contains four specimens
of Sirongylus cruciformis taken
from a bacfger (Meles) . The otters
are largely infested by flukes ;
Distoma trigonocephalum, D. in-
crassatum, and D. rude being found
in Lutra vulgaris, L. solitaria, and
L. braziliensis, respectively, the

Fig. 66.— Sirongylus gigas, coiled within the latter also harboring Hemistoma
kidney of a coati. Alter Leuckart. ....

clathratum. Otters are likewise
infested by Ligulce and Eustrongyli. The weasels (Mustelidce)
are attacked by a legion of entozoa, comprising flukes, tape-
worms, round worms, and thorn-headed worms ; they are also
liable to harbor many ectozoa, a large tick (Ixodes) being
especially troublesome.

Amongst the nematodes is Yan Beneden's Filaroides muste-
larum, for specimens of which I am indebted to Mr Wright
Wilson. This is found in the lungs, trachea, and in the
frontal and nasal sinuses of the common marten (Mustela foina),
in which situations it causes absorption of the cranial bones.
This worm occurs also in the polecat (M. putoriv&)} in the
common weasel (M. vulgaris), and in the pine-marten (M.
martes). The larvse reside in frogs. Weasels are also very
liable to have their kidneys invaded by Eustrongylus gigas. A

CAKNIVOEA

299

species of Ascaris and a Trichosoma (T. entomelas) are not
uncommon in the intestines. In regard to the tapeworms,
Taenia tenuicollis infests the polecat and the common weasel,
and T. intermedia the pine-marten. The most common fluke of
the weasel is Bistoma trigonocephalum. This infests the
intestine, while B. megostomum is found in the stomach. The
stoat or ermine (M. erminea) harbors Strongylus patens and
Taenia brevicollis.

The parasites of the civets, ichneumons and their allies,
(Yiverridae) are of little importance. Many years ago I
described a small fluke [Bistoma comp actum) obtained from the
lungs of the common Indian ichneumon (Viverra mungos). It
is figured in my ' Entozoa/ (p. 16). Two species of tapeworm
(Taenia platydera and T. genettae) have been found in tne
common genet (V. genettae), and also a round worm (Ascaris bra-
chyoptera). From another viverra (V. senegalensis) Dujardin
obtained a strongyloid worm (Bochmins crassus). A species of
mongoos (Herpestes leucurus) is likewise infested by tapeworms
(Bothriocephalus folium) .

Comparatively speaking, very few entozoa infest the hyaenas.
In this family I include the earth-wolf (Proteles lalandi).
Some years back Prof. Flower sent me a large number of
delicate nematodes found loose in the peritoneal cavity of this
singular South African carnivore. The worms themselves were
so peculiar that I was compelled to form a new genus for
their reception (Acanthocheilonema dracunculoides) . From the
intestines of the common Hyaena striata Dr Lautner obtained
Echinorhynchus gigas.

The parasites of the wolf, jackal, and fox family (CanidaB)
have especial interest, as including those of the dog. I can,
however, do little more than mention the names of the various
helminths of the dog, and the sources whence they come. At
the same time, I shall incidentally refer to the wild canine
animals that happen to harbor the same parasites.

The flukes of the dog are few in number.
Perhaps the most important is Bistoma conjunc-
tum, originally discovered by myself in an
American fox (Ganis fulvus) that died at the
Zoological Gardens. Lewis, eleven years after-
wards, found it in the pariah dogs of India,
where it is of frequent occurrence. It infests the F^fLm5Eight%nr"-
bile ducts. As already stated, Prof. McConnell £B ^naT

300

PARASITES OF ANIMALS

subsequently found this entozoon in man (1875), a second

instance being recorded later on (1878).
Another species of canine liver fluke has
been described by Prof. Ercolani (1).
campanulatum) , besides which there is
the winged species occupying the small
intestines (Holostoma (datum). This
latter is also found in Ganis azarce.

The tapeworms of the dog are not
only numerous, but also particularly
injurious, alike to their bearers and to
mankind. By experimental research we
have ascertained the sources of most of
the Tcenice. The serrated species (T.
serrata) is derived from Cysticercus
pisiformis infesting hares and rabbits.
This is common in sporting animals,
owing to the careless practice of allow-
ing gamekeepers and kennel masters to
throw the fresh viscera of the inter-
mediate hosts to the dogs. I have
witnessed this stupid habit in the field.
The cucumerine tapeworm (T. cucume-
rina) is, by most observers, considered
to be identical with the T. elliptica of
the cat. I regard it as a variety. This
delicate species is excessively common
and is now, through Melnikow's dis-
covery, known to be derived from the
louse of the dog (Trichodectes latus).
This circumstance affords a curious illus-
tration of the fact that an ultimate host
may carry the intermediate host upon
its back.

As regards the relative prevalence
of these tapeworms in England, it
may be said that whilst T. serrata
occurs almost entirely in our harriers,
greyhounds, sheep-dogs, and lurchers

C*; SSE^tffimt cucumerina is liable to infest any variety
t^wOZ^A^ of <*og, and probably infests nearly 70

OABNIVOB \

301

per cent. According to Krabbe the prevalence of the last-named
species is 57 per cent, in Iceland and 48 per cent, in Copenhagen,
whereas the T. serrata is almost absent from those countries.
The gid tapeworm (T. coenurus) is derived from the ordinary
gid hydatid infesting the brains of sheep and lambs. The
polycephalous bladder-worm (Coenurus cerebralis), so familiar to
agriculturists and veterinarians, is often confounded with the
ordinary hydatid infesting ruminants. Coenuri infest the soft
parts of rabbits, but it remains to be shown whether they are the
same species. Possibly the Gcenurus cuniculi is merely a variety.
The gid tapeworm is not very abundant in England. In Denmark
it appears to be rare, occurring in 1 per cent, only ; but in
Iceland Krabbe found it in 18 per cent. In common with other
helrainthologists, I have frequently reared this and the serrated
species by worm feedings administered to dogs. The lettered
tapeworm (T. litter ata) is very commonly spoken of as the Tcenia
canis lagopodis. It was so named by Viborg, but I prefer the
more distinctive nomenclature of Batsch. We know nothing,
for certain, respecting the source of this entozoon. It is rare
if not altogether wanting in Denmark, but abundant in Iceland
(21 per cent,). I have obtained specimens from a cheetah
( Ganis jubatus) which died at the Zoological Gardens, and Mr
W. H. Jackson, of Oxford, found it in a cat. The worm is
certainly not confined to the Arctic fox (G. lag opus.).

A well-known tapeworm infests the fox which has not yet
been noticed in the dog. This is the Tcenia crassiceps, whose
scolices (Gysticercus longicollis) reside in the viscera and soft
parts of field mice and voles (Arvicola arvalis, A. terrestris, A.
amphibius). This relationship was pointed out by Leuckart.
Another tapeworm (T. opuntioides) mentioned by Rudolphi as
occurring in the wolf, seems to be of doubtful authenticity. A
formidable and not uncommon tapeworm is Tania marginata.
This large species occurs in at least 25 per cent, of English dogs,
whilst in Iceland its prevalence reaches 75 per cent. In Den-
mark about 14 per cent. only.

It is well known that the larval or scolex stage (Gysticercus
tenuicollis) of the margined tapeworm resides in the sheep and
dog. In a feeding experiment with five examples of this bladder
worm I reared five strobiles of ten days' growth. These imma-
ture tapeworms were each one inch long. By far the most
important tapeworm of the dog, however, is the hydatid-forming
species (T. echinococcus) . This remarkable entozoon is the sole

302

PARASITES OF ANIMALS

cause of the terrible echinococcus disease, so prevalent in Iceland
and elsewhere. Experimental research, initiated by von Siebold,
has explained its origin ; Van Beneden, Zenker, and others have
also experimented successfully. Rarely attaining a length of
3", the perfect strobile is made up of only three proglottides in
addition to the head, the lowermost segment being sexually
mature. As hydatids (Echinococcus veterinorum or E. homii
are found in a great variety of animals as well as in man, and
as these bearers form so many kinds of intermediate hosts, it is
easy to understand how readily dogs and wolves may acquire
the sexually-mature tapeworm. I am in possession of hydatids
from the liver of a clouded tiger (Felis macroscelis) . In
England the Ttenia echinococcus is excessively rare, and has not
been seen in any dog which had not previously been subjected
to a feeding experiment. Mr Nettleship succeeded in rearing
large numbers. In Iceland, Krabbe found dogs to be infested
to the extent of 28 per cent., a proportion fully explaining the
prevalence of hydatid disease in that country. The remaining
tapeworms of the dog belong to the genus Bothriocephalus ; of
these, the broad tapeworm (B. latus) is best known, because it
infests man. Diesing has described a variety found in the
Pomeranian dog as a separate species [Bibothrium serratum).
The museum of the Royal Veterinary College contains a very
perfect specimen of B. latus from an English dog, but the
parasite is of rare occurrence in this country. It is generally
supposed that this tapeworm is derived from the consumption
of fish belonging to the salmon and trout family, but Dr Pock,
of Utrecht, thinks that the bleak (Leuciscus alburnus) is the
usual intermediate host. I have already discussed this question
at some length. Experimental proof is still wanting. In
addition to B. latus the dog is liable to harbor B. cordatus, B.
fuscus, and also two varieties of the last-named species (7?.
dubius and B. reticulatus, Krabbe). Taking the pit-headed
tapeworms as a whole, their prevalence in Iceland is not con-
siderable, amounting to about 5 per cent. only. Lastly, it may
be mentioned that instances are recorded of the occurrence of
the hog-measle (Cysticercus celluloses) in the dog. Though
many have felt sceptical on this point, Grurlt's authority is not
to be lightly set aside, confirmed, as it has been, I believe, by
MM. Megnin and Leblanc.

Passing to the round worms it may be said that Ascaris mar-
ginata is, at the best, a mere variety of A. mystax of the cat,

OA RN I VOU A

:j()3

with which must also be placed A. leptoptera of the lion and
other felines. The lateral appendages not only vary in breadth
in these three forms, but also in the specimens obtained from
each host. I have encountered examples in a dog, which
measured more than six inches in length. The worm is excessively
common in England, occurring in probably not less than 75
per cent., whilst in Denmark it occurs in about 24 per cent.
According to Krabbe it is rare in Iceland. Its presence is at
all times more or less injurious to the bearer, being a frequent
cause of sickness, colic, convulsive fits, and paralysis. Occa-
sionally the worms prove fatal to dogs by wandering into the
trachea. At the Eoyal Veterinary College, in 1864, a litter of
six puppies, of only three weeks growth, died rather suddenly
in consequence of the presence of these worms in the stomach
and small intestines. So far back as the year 1684 Redi de-
scribed round worms from the walls of the oesophagus of a dog.
These were afterwards noticed by various observers in tumours
of the mucous membrane of the stomach. Owing to their red
color, derived from the ingested blood of the host, the species
was named Spiroptera sanguinolenta. In 1867 I suggested
that the minute Filarise found by Grube and Delafond in the
blood of dogs would probably turn out to be referable to this
species. The researches of Lewis have proved that this suppo-
sition was correct. To be sure, other nematoid hsematozoa, of
microscopic dimensions, occur in the dog, but those described
by Grube and Delafond may be referred to Spiroptera.
These authors estimated their number in the canine host
to vary from 11,000 to upwards of 200,000. In one instance
Messrs Grube and Delafond found six worms lodged in a clot
occupying the right ventricle of the heart. Four were females
and two males. Although they were described as representing
an altogether new species, which they termed Filaria papillosa
hamatica canis domestici, I think there can be little doubt that
they were examples of Spiroptera sanguinolenta not fully grown.
The writings of Lewis abound with interesting details respect-
ing the structure and development of this worm, and as much
may be said of the writings of Manson and Welch concerning
the cruel threadworm {Filaria intimitis, Leidy) occupying the
right cavities of the heart. I was first made acquainted with
this entozoon in 1853, by examining specimens in the possession
of Prof. Hughes Bennett of Edinburgh ; at which time also I
was put in possession of a valuable MS. (since lost) describing

304 PARASITES OF ANIMALS

the ravages of this entozoon in the dogs of China. I have

since received numerous verrnini-
ferous hearts both from China and
Japan, and also some heart-worms
from Charleston, U. S., sent by Mr
M'Innes. In a recent communica-
tion, Dr Manson has spoken of this
worm as if it were comparatively
harmless, but all the evidence I
long ago received through the late
Mr Swinhoe, formerly H. B. M.
Consul at Anioy, through Mr Dare's
letters enclosing Dr Orton's valu-
able observations ' (addressed to the editor of the ' Field'),
through the lost MS. above alluded to, through Dr Lamprey's
statements, and through many other sources, lead to the
very opposite conclusion. No doubt the canine hosts do for a
time appear to be little inconvenienced by their nematode
guests, but sooner or later the most distressing symptoms set
in. As in Hoysted's case (quoted below) the convulsive spasms
may occasion death in a few minutes, but frequently they last
for hours or days, with more or less prolonged intervals of relief
before the final struggle.

Some other filariform nematodes have been imperfectly
described. Of these, Gescheidt's Filaria oculi canini (F. tris-
jnnulosa, Diesmg) was probably a sexually-immature worm, and
the same may be said of the encysted worms found by Mi-
Mather in the mucous coat of the intestines and in the liver
ducts and acini (Filaria h&patica, Cobbold). Of more interest
is Leisering's haematozoon (Strongylus subulatus). These
minute worms occupy the veins, the largest females not
exceeding TV in length. They are viviparous, and thus
form another source of embryonic hsematozoa. A single drop
of infected venous blood commonly carries from four to six
mature worms. In this place may be mentioned Dr Osier's
Strongylus canis bronchialis. The largest males measure J" and
the females fully \" . In the worms sent to me by Prof. Osier
I saw no evidence of strongyloid structure, and in his description
he avoids all mention of the presence of any caudal hood in
the male. I regard the worms as Filarise (F. Osleri, Cobbold).
Very great interest attaches to them from the fact that they
produce a destructive canine epizooty, resembling the ordinary

I'ig. bH.— Filaria immilis. Tail of male.
Enlarged. Original.

CARNIVORA

305

"husk" or parasitic bronchitis of calves, lambs, and other
domesticated animals. The only other genuine strongyle known
to infest the dog is Enstrongylus gigas. This is a very common
parasite in wolves. I have already spoken of this parasite at
some length (Book I, p. 207), and can only further refer to
the recently published case by Megnin (quoted below, and at
full length in my paper in the 'Veterinarian' for April, 1879).
The Museum of the Royal Veterinary College contains three
fine examples of this worm coiled within the kidney of a dog, or
rather within the renal capsule, for the substance of the organ
is almost entirely wanting. These are from Bickford's case.

Amongst the many good " finds " made by Lewis in India,
not the least interesting is that appertaining to Cheir acanthus
robustus. Lewis, indeed, supposed that he had detected Echi-
norhynchi in chestnut-sized tumours of the walls of the stomach,
but, as I pointed out at the time, the parasites were entirely
destitute of Acanthocephalous structure. As is well known,
this curious nematode infests various felines, such as the wild
cat, puma, and tiger. In addition to the above canine nematodes
we have the wrinkled threadworm {Trichosoma plica) infesting
the bladder. This is of much more frequent occurrence in the
fox. In the list Trichina spiralis must also be included,
although, so far as I am aware, it has only been seen in dogs
that have been subjected to feeding experiments.

Another nematode common to the fox and dogs, and infesting
the caecum, is the whipworm (Trichocephalus depressiusculus) .
It is very rare in the dog. Lastly, there is the important little
strongyloid worm generally known as Dochmius trigonocephaly.
At the hands of Leuckart the general structure and development
of this entozoon have received complete elucidation. It infests
the small intestines, and is found alike in the fox, wolf, and
cheetah. It has also been obtained from Canis lagopus and C.
azarte. Dr Krabbe did not encounter this worm in Icelandic
dogs ; nevertheless, he obtained it in a blue fox which died in
Kjoerbolling's menagerie, and which had come from Iceland.
In Danish dogs it occurred in less than 2 per cent. The
embryos of this worm are rhabditiform and possess three
long bristle-like teeth, the slender tail being furnished with a
distinct appendage at the tip. They develop in moist situations,
where they feed freely, grow rapidly, and change their skins,
throwing off the caudal tip with the first month. It seems
evident that they do not require a change of hosts, since Leuc-

300

PARASITES OF ANIMALS

kart succeeded in roaring the sexually-mature Dochmii by
introducing the rhabditiform larvae into the stomach of the dog ;
moreover, his experiments upon water-snails belonging to the
genus Physa gave negative results.

Of Arachnidan parasites (Trachearia) infesting the dog, by
far the most interesting is the well-known Pentastoma tcenioides,
shown by Leuckart to be the adult condition of the still
better known Pentastoma denticulatum. It resides in the nasal
sinuses. As already mentioned in a former part of this work,
these parasites present four marked stages of growth, namely
(1), the embryo, (2) the pupa, (3) the active larva (P. denticu-
latum), and (4) the sexually-mature worm. As the eggs and their
embryonic contents are lodged in the nasal mucus of the dog,
and are commonly distributed by the act of sneezing on the part
of the animal, the sources of infection are not far to seek.
Clearly the larvae usually get introduced to the bodies of man-
kind and herbivorous animals by the ingestion of unclean vege-
table matter. The embryos set free in the stomach bore their
way to the liver and other viscera, in which organs encystation
and moulting subsequently take place. The fondling of dogs
infested by pentastomes may prove dangerous by a more direct
transference of the eggs to the hands and mouth. As regards
the dog, the adult parasite has been known to prove fatal. A
very striking instance of this kind was recorded by Prof. Dick,
where the worms wandered into the trachea producing asphyxia.

The ectozoa of the dog, though not numerous as species, are
of importance in relation to mange. The follicle-mites form a
family by themselves (Demodicida) , and, as already observed,
those infesting the dog and cat are, alike, mere varieties of the
human species (Demodex folliculorum, var. caninius and var. cati).
Whilst the human parasite restricts itself to the face, the canine
variety (fig. 52) will occupy any part of the dog's body. The
follicle-mite of the cat, however, usually confines itself to the
ear. According to Megnin, to whose beautiful monograph we
owe so much, two or three dozen of these parasites may be
found occupying a single follicle of the dog. Acne-like pustules
are thus formed, and when they are very numerous death may
result from the excessive irritation, which is usually accom-
panied with depilation. An interesting example of this kind
recently occurred at the Royal Veterinary College. In regard
to the ordinary mange-mite (Sarcoptes canis, Gerlach) M. Megnin
points out that it is in all respects identical with the human

CARN1V0KA

307

itch -insect. In the wolf and fox, however, the same species
forms well-marked varieties (8. scabici, var. lupi and var.
vulpis). As regards true insect parasites and tormentors of the
dog, I can only allude to a few of them. In tropical America
dogs are said to be attacked by the larvae of a species of gad-
fly (CEstrus cards), whilst in Africa they are often fatally bitten
by the tsetse (Glossina morsitans). In addition to the flies
(Diptera), several kinds of fleas (Aphaniptera) frequently prove
troublesome (Pulex canis, P. martis, and P. penetrans), and the
same may be said of certain lice (Hemiptera) . The common
louse of the dog (Trichodectes lotus) proves especially noxious
to young puppies. Of the two other species, namely, Eamato-
pinus piliferus and H. canis, the former is tolerably common,
whilst the latter is comparatively rare. This species is also
found on the ferret. A new form of mite (Ghorioptes ecaudatus),
infesting the ears of the ferret, has recently been described by
M. Megnin.

I have already referred to several of the parasites of the cat-
tribe (Felida), but some others require notice. Only two
flukes (Amphistoma truncatum and Hemistoma cordatum) have
been described as infesting the cat. Of the tapeworms, Tcenia
crassicollis is the best known. This is derived from Cysticercus
fasciolaris of the mouse and rat. It is not uncommon to find
this scolex in the sexually- immature tasnioid state in the liver,
measuring six or seven inches in length. An exceedingly
interesting communication by Dr Romano, of Gremona (Frioul),
demonstrates the possibility of severe feline epizooty as due to
this entozoon. As I gather from an account given in the journal
quoted below, " during the summer of 1876, Dr Romano was
informed by his confrere Dr Leoncini, a physician practising at
Osoppo, that for about a fortnight most of the cats in a certain
hamlet of the town had died without appreciable cause after
presenting the following symptoms : — Gradual wasting, with
complete loss of appetite, retracted abdomen, slight diarrhoea
at first, then constipation, abundant saliva, contraction of the
elevating muscles of the upper lip in some subjects, great pros-
tration of strength, loss of the visual faculty. Some of the
feline patients no longer heard or appeared no longer to hear
their master's voice ; some vomited and seemed to experience
relief, for the appetite improved, but they soon died like the
others. Nervous phenomena, epileptiform convulsions, and more
frequently colic, also showed themselves. Having visited the

308

PARASITES OP ANIMALS

locality (of the outbreak), Dr Eomano could not at first pro-
cure any corpses for the post-mortem examination, for the
children had thrown them into the Tagliamento, which flows at
the foot of the fortress of Osoppo. It was only after the lapse
of some days that he was able to open one of the animals
which had just succumbed. The principal evils were remarked
in the stomach, the walls of which were retracted and
formed the seat of a catarrhal inflammation, from the products
of which a long, white, flat worm was removed with care for
examination. All the other organs were in good condition.
The examination of the helminth in the stomach, made with the
help of Dr Leoncini and Fachini, showed that the flat worm
(white, and with the body divided into rings, 12 centimetres
long, and 5 or 6 millimetres broad) had all the characters of
the taenias, and this was confirmed by a microscopic examina-
tion of the head. A few days later Dr Romano made an
autopsy of two other cats. In one of the corpses he noted the
alterations described above, and found a tgenia smaller than the
first ; in the other the same lesions without any helminth.
This negative circumstance very naturally disconcerted Dr
Eomano, but several people of the place came to assure him
that they had seen their cats, during the course of the malady,
after violent and repeated efforts at vomiting, throw up a sort
of white cord, which they recognised as corresponding with the
ta3nia he showed them. Thus confirmed and reassured in his
diagnosis, Dr Romano sought to identify the species." In this
connection it is specially interesting to note that " during the
whole summer the inhabitants of Osoppo had been over-run by
bands of rats proceeding from the fortress. They were com-
bated by means of cats, and it was the best hunters among
the felines that succumbed. Here was, therefore, a striking
relation of cause and effect which could not be gainsayed."
Dr Romano communicated his observations to the National and
Royal Veterinary Society, but by an error in the report
the species appears to have been described as Tcenia tenuicollis
instead of T. crassicollis. In this connection I have only
further to add that the wild cat is infested by a tapeworm
scarcely an inch in length (Tania Uneata). A species of
Bothriocephalus {B. deci%)iens) likewise infests the domestic cat,
in common with most of the wild felines, such as the tiger,
puma, ounce, and jaguar. Dr Bancroft brought me a specimen
from an Australian cat. The nematodes of the cats are very

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309

abundant. Dr Bellingham found a trichosome (T. felis eati) in
the urinary bladder of the wild cat. This is probably identical
with T. plica. A tolerably common nematode is Dochmius
tubaformis, which occurs not only in the cat but also in the
leopard, puma, jaguar, ounce, panther, and also in Felis tigrina
and F. mcllivora. In the last-named and in other Brazilian
felines an echinorhynchus (E. campanulatus) was found by
Natterer. A strongyle (Sclerostoma dispar) infests the lungs
of the puma. A species of spiroptera (S. subtequalis, Molin)
infests the oesophagus and stomach of the lion and tiger. Redi
also noticed a species of Filaria beneath the skin of the lion.
PJiysaloptera terdentata (Molin) and P. digitata (Schneider)
infest the stomach of the puma. I can only allude also to Taenia
latieollis of the lynx, Pentastoma recurvatum, occupying the
frontal sinuses and air-passages of the ounce, and Ligula
reptans the subcutaneous tissues of the leopard. Lastly, there
is the Olulanus tricuspis of the domestic cat. I have examined
the lungs of three cats containing this parasite, which was first
described as an entirely new species by Leuckart. The adult
worms, only in length, occupy the walls of the stomach.
Thence they are apt to migrate or stray into the lungs and
liver, where they encyst themselves. When myriads of them
are thus encysted a kind of nematode tuberculosis is set up.
This disease I have elsewhere called olulaniasis. Ordinarily,
however, the encysted condition of olulanus is to be found in
the muscles of mice, which are thus said to be olulanised.
Clearly, as Leuckart's experiments substantially prove, the
domestic cat acquires the adult worm by catching and devour-
ing olulanised rodents. Every now and then the disorder thus
created produces a virulent and fatal feline epizooty.

Bibliography (No. 45). — (Anonymous), "Curious Note on
two Taenia from the Dog (asserting that neither strong whisky
nor boiling water would kill them)/' 'Med. Commentaries/
vol. xvi, p. 370, 1791.— Araujo, A. J. P. S., "A Filaria
immitis e F. sanguinolenta no Brazil/' ' Gazeta Medica da
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Worm found in the Heart and Vessels of a Dog/' from ' Journ.
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Dogs in China," ' Proc. Linn. Soc./ vol. ix, Zool. Div., p. 296,
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Beneden, below).— Bickford (1. c, Bibl. No. 28).— Chapman,

310

PARASITES OF ANIMALS

" On Ascaris mystax (leptoptera) in the Tiger and American
Wild Cat," ' Proc. Acad. Philad./ 1875, pp. 14 and 17.— Cherry,
F. F., " Worms in the Stomach of a Dog," in the ' Farrier and
Naturalist/ 1829, vol. ii, p. 303. — Glamorgan, J. de, " Serpents
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on Entozoa, with experiments in regard to Tcenia serrata and
T. cucumerina," ' Linn. Trans./ 1858. — Idem, " On some new-
Forms of Entozoa," ' Linn. Trans./ 1859. — Idem, " Further
Observ. on Entozoa, with Experiments," 'Linn. Trans./ 1861.
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new generic type of Entozoon (Acanthocheilonema) from the
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vol. xxii, p. 363, 'Proc. Zool. Soc./ March, 1861. — Idem, various
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on the Life-epochs (biotomes) of T. coenurus and T. echinococcus,"
in a paper on 'Animal Individuality/ 'Journ. Linn. Soc./ Zool.
Div., vol. viii, p. 163. — Idem, " Worms in the Heart (Mr.
M'Imies' specimen from Charleston)/' the ' Veterinarian/ Feb.,
1875. — Idem, "Note of Lewis' Discoveries," in 'Nature/
March 11, 1875, vol. ii, p. 363. — Idem, "Kemarks on Fustron-
gylus gig as " in the 'Veterinarian/ April, 1879. — Coles, F. C,
" Worms in the Heart and (Esophagus of a Dog," ' Path. Soc.
Kep./ March, 1878. — Dare, J. J., " Death of Dogs from Worms in
the Heart," the 'Field/ Feb. 24, 1872.— Davaine, 'Traite/ 1. c,
2nd edit., p. 290 (for refs. to lit. of Strongylus gigas). — Delafond
(with Grube), " Note on a Verminiferous kind of Blood of a Dog,
caused by Hsematozoa of the genus Filaria," from 'Ann. de
Chimie et de Physique/ in ' Ann. of Nat. Hist./ vol. xi, 1843, and
' Comp. Rend./ in 'Lond. Phys. Journ./ p. 28, 1843 ; also from
' Gaz. Med./ in 'Med.-Chir. Eev./ vol. xxxv, p. 393, 1843-44 ;
see also ' Edin. New. Phil. Journ./ vol. lii, p. 233, 1852, and
'Veterinarian/ vol. xviii, p. 216, \Uh.—Dich, "Worms the
cause of Sudden Death in a Dog," the ' Veterinarian/ vol. xii,

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p. 42, 1840 (Pentastomes). — Diesing, ' Syst. Helm./ vol. ii,
p. 327 (Eustrongylus). — Ercolani, G. B., " Osservazione elmin-
tologiclie sulla dimorfobiosi nei Nematodi, sulla F. immitis e
sopra una nuova specie di Distoma dei cani," ' Mem. Accad.
Bologn., 1874-5. — Frank (see Bibl. No. 28). — Gay, J., Speci-
men of Olulanus in Hunterian Museum, and marked in the
'Catalogue' "No. 1814a, Lung of Cat, pneumonic, from the
presence of parasites." — Gay, "Nematodes of Panther," in
'Hering's Repert./ 1873, from 'II. Med. Vet/— Herbst, "On
Trichinse in the Badger," from ' Ann. des Sci. Nat./ in ' Assoc.
Med. Journ./ 1853, vol. i, p. 491. — Eoysted (and Sir J. Fayrer),
"On Filaria sanguinis (really F. immitis)," 'Lancet/ March 1,
1879, p. 317. — Jamieson (see Manson). — Jardine (see Manson). —
Klein, J. T., " Worms found in the Kidneys of Wolves,"
'Phil. Trans./ vol. xxxvi, 1729. — Krabbe, " Husdyrenes In-
voldsorme," 'Tidsskrift for Veterinairer/ 1872. — Leblanc,
" Subcutaneous Tumour in the Dog, due to the presence
of Strongylus gigas," from ' Eecueil de Med. Yet./ in ' Bdin.
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Megnin). — Legros, "Haematozoa of Dogs," in ' Rec. de Med.
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Archiv/ 1865. — Leuclmrt (see Bibl. Nos. 28, 29). — Idem,
(for development of T. serrata, &c), ' Die Blasenbandwiirmer
und ihre Entwicklung/ Giessen, 1856. — Idem, ' Bau und Bnt-
wicklungsgeschichte der Pentastomen (especially P. tanioides
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norhynchus, &c, from the Dog," in his oft-quoted ' Memoir/
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und Seitenfelder von Filaroides mustelarum, v. Ben./' ' Arch,
f. Naturg./ xl, s.. 135 (with figs.). — Idem, " Einige neue Nema-
toden," &c, 'Arch. f. Naturg./ p. 293, Bd. xxxix. — Manson,
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figs.), Jan.-March, Shanghai, 1877. (N.B.— The same journal,
Rep. No. 12, Gaz. No. xxx, contains remarks on worms in
the heart of dogs by Dr A. Jamieson. The subject is also
discussed by Dr Jardine.) See also ' Med. Times and Gaz./
Oct. 20, 1877, p. 480.— Mather, T., " Eilarice found in the

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PARASITES OF ANIMALS

Intestines of a Dog," ' Veterinarian/ vol. xvi, p. 434, 1843.—
McGonnell, "On D. conjunctum," ' Lancet/ March 30, 1878,
p. 476. — Megnin, P. " On Myobia," 'Abhandl. ueber eine neue
Gruppe von Acariden (u. s. w.)/ in ' Revue f. Thierheilkunde/
Oct., 1878, p. 149. — Idem, " Sur un parasite des oreilles chez
le furet/J (Eec. de Med. Yet./ Oct., 1878.— Idem (avec 0.
Leblanc), " Note sur le Cysticercus celluloses developpe chez le
chien," ' Bullet, de la Soc. cent. vet. de Paris/ 1873. — Idem,
" Sur le Strongylus gig as/' Bullet, de la Societe Bntora. de
France, No. 3, 1879.— Melnihow, N., "On Tcenia cucumerma,
in 'Arch. f. Naturg./ 1869, p. 62.— Miller, K, 'Six Cases of
Strongyle in the Kidney of Putorius vison ' (quoted by Davaine
from Museum of Boston, U.S.). — Whines (see Cobbold). —
Molin, 'Nuovi myzelmintha/ s. 34, 1859. — Idem, 'Una monog.
del gen. Physaloptera (P. terdentata)/ 1860, s. 17. — Idem, 'Una
monog. del gen. Spiroptera (S. subcequalis) / 1860, s. 13. —
Nettleship, E., " Notes on the Rearing of Tcenia echinococcus in
the Dog from Hydatids, with some observations on the anatomy
of the adult worm," 'Proc. Roy. Soc./ 1866, No. 86, p. 224,
with figs. — Noseda, B., " Six Strongyles in the Kidney of the
Agouai"a-gouazura or Puma," in Don F. de Azara's ' Qua-
druples du Paraguay/ Paris, 1801 (see also Azara, Bibl.
No. 28). — Osborne, T. C, "Worms found in the Heart and
Blood-vessels of a Dog (with symptoms of hydrophobia),"
'Western Med. Journ./ rep. in 'Bost. Med. and Surg. Journ./
vol. xxxvii, p. 448, 1847-48. — Osier, W., "Verminous Bron-
chitis in Dogs (from Strongyles)," 'Veterinarian/ June, 1877,
p. 387. — Owen, " On the Anatomy of Linguatala tanioides,"
'Trans. Zool. Soc./ vol. i, 1835, and in ' Lond. Med. Gaz./
1835. — Idem, ''Anatomical description of two species of Entozoa,
from the Stomach of a Tiger, one of which forms a new genus,
Gnathostoma," ' Proc. Zool. Soc./ part iv, 1836. — Parana (and
Grassi), "On a new Species of Dochmius (D. balsami)/' 'Reale
Institute Lombardo di Scienze e Lettere, Rendiconti/ vol. x,
fasc. vi, 1877. — Baynold, T. M., "Tapeworm in the Pointer and
Spaniel," 'Veterinarian/ vol. xiv, p. 694, 1 841 — Roman n,
" Epizooty amongst Cats from Tcenia crassicollis," ' Giornale di
med. vet. practica/ Aoiit, 1877, and in ' Journ. de Med. Vet.,*
Avril, 1878.— Srhn/>j>rrt, M., "Mechanical Obstruction of the
Heart (of a Dog) by Entozoa, causing Death," 'New Orleans
Med. News and Hosp. Gaz./ Jan., 1858, also ' Bost. Med. and
Surg. Journ./ vol. lvii, 1857-58, and in 'Med.-Chir. Rev./

CAUNIVOKA

313

1858.— Stirling, " On the Changes produced in the Lungs by
the Embryos of Ohdanus tricnspis," 'Quart. Journ. Micr. Sci./
1877, p. 145. — Van Beneden, " On the Transmigration of the
Entozoa" (in answer to MM. Pouchet and Verrier), from the
French, by Busk, in 'Quart. Journ. Micr. Sci./ 1862. — Idem,
" Eesearches on the Intestinal Worms/' from Yan der Hoeven's
abstr. in the ' Nederlandsch Tijdschrift voor Geneeskunde/ in
' Ann. Nat. Hist./ vol. iii, 3rd ser., 1859.— Idem, "New Obs.
on the Development of the Intestinal Worms/' from ' Compt.
Eend./ in ' Ann. Nat. Hist./ vol. xiii, 2nd ser., 1 854.— Idem,
" On Filaroides mustelarum," ' Mem. sur les vers intest./ 1858,
p. 267.— Welch, "On Filaria immitis," ' Lancet/ March 8, 1873 ;
also in ' Month. Micr. Journ./ Oct., 1873, p. 157 (with three
plates). — Wilson, W., "A Parasitic Worm infesting the Air
Sinuses of the Weasel," 'Midland Naturalist/ May, 1878.—
Wright, " On Worms found in the Pulmonary Artery of a Dog/'
'Lancet/ 1845, and in ' Veterinarian/ vol. xviii, p. 52, 1845. —
Youatt, " On Tumours (containing Entozoa) in the Stomach of a
young Tiger/' ' Veterinarian/ vol. x, p. 619, 1837 ; see also
Owen on ' Gnathostoma.' — Idem, " Worms (Ascaris marginata)
in the Nose of a Dog/' 'Veterinarian/ vol. v, p. 337, 1832.

PART V (Pinnipedia).

Following the order of classification adopted in my descrip-
tion of the Mammalia in the ' Museum of Natural History/ I
proceed to speak of the internal parasites of the seals (Phocidce)
and walruses (Trichecida) . From their piscivorous habits one
would naturally expect the seals to be largely infested with
entozoa, and yet, though sufficiently victimised, they are not
liable to entertain so great a variety of helminths as the fishes
themselves on which they feed.

The flukes observed in Phoca vitulina are Distoma acanthoides
and Amphistoma truncatum, the latter occurring also in P.
grcenlandica. In another seal (P. barbata) we have D. tenuicolle.
The nematodes are more numerous. The best-known is the
maw-worm {Ascaris osculata), which seems to be always present
in full-grown seals of every kind. In the years 1862-64 I
conducted a series of experiments with the eggs of this worm.
1 reared embryos both in salt and fresh water, but the adminis-
tration of the young worms to various animals led to no result.

314

PARASITES OF ANIMALS

However, I succeeded in watching the growth of the embryos
until they had acquired well-marked digestive organs and
a length of ^' , their size when emerging from
i \ the egg-shell in the water having been about
11 \| tto" onlj- The large strongyle {Eustrongylus

S yigas) has been found in various organs of the
common seal. Of more interest are the Filarice
found in the heart of seals, which in many re-
spects resemble those obtained from the same
situation in dogs. Professors Joly, Leidy, and
myself, have each described a species, but ap-
parently our descriptions all refer to one aud
the same parasite. It has also been seen by
Camill Heller. The close correspondency in
size and other characters of Leidy's Filana
spirocauda and my Filaria hebetata leaves little
doubt as to their identity. As the worms were
both originally noticed by Leidy and Joly in
1858, I cannot pronounce upon the question
no. w.-Mcaris oscn- of priority of discovery. By Joly the worm
was called F. cordis phoca. In Leidy's and
0n6lnaL in my own specimens the males were four inches

long, and the females six inches ; they extended up to 8" in
some of the American examples. The worms found by Prof.
Joly were all females. Professor Millen Coughtrey, who furnished
me with the seal's heart, stated that it was obtained from a male
hoodcap (Stemmatopus cristatus), a rare visitant of our Bi'itish
coasts. This seal was captured on the Cheshire side of the
Mersey river. Leidy and Joly obtained their specimens from
Phoca vitulina. In the common seal have also been found
Ligula crispa, Schistocephalus dimorphus, and Fchinorhynchus
strumosus. In other seals a not uncommon tapeworm of the
Bothriocephalous type is that called Dibothrium hians by
Diesing. To Prof. Krabbe I am indebted for a specimen of
Bothriocephalus fasciatus taken from Phoca hispida. There is a
nematode of frequent occurrence in P. hispida and P. gropnlan-
dica. This is the Ophiostoma dispar of Eudolphi. In addition
to the above I can only add that P. barbata is infested by
Liorhynchus gracilescens, occupying the stomach, and by a tape-
worm, Tetrabothrium anthocephalum, which is found in the lower
part of the large intestine.

Bibliography (No. 46). — Cobbold, " Description of F. hebetata,"

RODENTIA

315

in 'Notes on Entozoa/ part i, sp. 3, 'Proc. Zool. Soc./ Nov.
18th, 1873, p. 741— Idem, " On Ascaris osculata," in * Report of
Experiments respecting the development and migrations of the
Entozoa ' Brit. Assoc. Trans./ 1864, p. 114.— Heller, G., in
< Schrift der zool.-botan. Gesellsch., ' Wien, 1858, s. 83.—
" On a new Species of Haematozoon of the genus Filaria,
observed in the heart of a seal ; " from < Compt. Rend. Acad.
Sci./ 1856, p. 403, in ' Ann. Nat. Hist./ vol. i, 3rd ser., 1858 ;
also abstr. in the < Year Book/ 1859.— Leidy, J., (E. spirocauda)
in 'Proc. Philad. Acad./ 1858, p. 112.

PART VI (Rodentia).

Though very numerous, the parasites of this order are
chiefly interesting as embracing those of the hares and
rabbits, moles, mice, rats, squirrels, and beavers. Some slight
notice, however, will be given of the entozoa of each of the
eleven families into which the order may be divided.

The squirrels (Sciuridce) are liable to be infested by the
common liver fluke (F. hepatica), and also, it is said, by a
cysticercus (0. tenuicollis). I have never encountered this
bladder worm, but in 1864 I described some polycephalous
hydatids (Ccenuri) which I obtained from the viscera of an
American squirrel. I think the host was of the same species
(Sciurus vulpinus) as that from which Mr Chapman has since
obtained an example of Echinorhynchus (E. moniliformis).
This worm also infests the hamster. A very small female
round worm, probably a strongyle, was described by Rudolphi
as Ascaris acutissima. It infests the caecum of the common
squirrel, in which host a species of tapeworm is tolerably
frequent (Tcenia dendritica) . The common European marmot
is infested by T. pectinata, so abundant in hares and rabbits.
I have also noticed it as occurring in the Canadian porcupine
(Hystrix dorsata). The dormice (My oxides) are not much
troubled with parasites, at least I have not encountered any
in our common Myoxus avellanarius. In M. glis, however, a
tapeworm, and at least one species of strongyle (S. gracilis), have
been observed. Dujardin described very fully another strongyle
(8. Icevis) from M. nitela, from the long-tailed field-mouse (j\fus
sylvatica), and from Arvicola suhterraneus. The other species
are Trichosoma myoxi nitelce, and Ophiostoma cristatnm from

316

PARASITES OF ANIMALS

Myoxus dry as, and M. muscardinus. The jerboas (Dipodidce) ,
in common with the hamster and several species of true mice,
are apt to be infested by Ascaris tetraptera ; and a small
nernatoid, apparently immature, was noticed by Otto in the
intestines and in the abdominal walls and cavity of Dipus
tetradactylus. Mice, properly so called, are largely infested,
as is also the hamster (Cricetus vulgaris), which I include in the
Muridoe. In addition to the parasites already mentioned, the
hamster is infested by Taenia straminea. Along with examples
of this tapeworm I have received from Dr Murie some acepha-
locysts found in a hamster which died at the Zoological Gardens.

Flukes exist in the long-tailed field-mouse (Distoma vitta and
D. recurvum), but I have not seen any in our common mice and
rats. However, Dujardin desci'ibes a distome (D. speculator)
in the brown rat (Mus decumanus). One of the tapeworms
observed in the mouse (M. musculus) is Taenia pusilla, also found
in the rat (M. rattus) and long-tailed field-mouse. The house-
mouse likewise harbors T. microstoma and T. leptocephala ; and
an immature cestode has also been seen in the abdomen, probably
a species of Ligula. Various species of rat also harbor T.
diminuta. In regard to the round worms one of the most
common species is Ascaris oxyura. This not only occurs in
rats and mice, but also in voles, water-rats, and many other
rodents. The rodents' whipworm (Trichocephaliis nodosus) is
yet more common in the lemmings, rats, voles, and mice ;
another species (T. unguiculatus) , taking its place in hares and
rabbits, and yet another (T. affinis) in the porcupine. Another
nematoid, very common in mice, is Spiroptera obtusa, occupying
the stomach. I have seen a mouse with its abdomen so dis-
tended by their presence that the animal could scarcely run
along the pathway where it was killed by being trod upon.
According to Marchi, the young of this entozoon dwell in the
fat surrounding the alimentary canal of the larva of an insect
(Tcnebrio molitor). When noticing the parasites of the cat I
referred to Leuckart's interesting discovery of the relations
subsisting between the adult Olulanus tricuspis, found in the
stomach walls of that feline, and the immature encysted worms,
found not only as wanderers in the cat itself but also in the
muscles of mice. The olulanised mouse is thus an intermediate
host. Eats and mice also play the part of intermediary bearers
in the case of two other species of entozoa, namely, Trichina
qpvraMs and Taenia crassicollis, the tamioid scolex or larval

RODENT r A

317

condition of the cat's tapeworm being familiarly known as
Cysticercus fasciolaris. This sexually-immature tapeworm infests
many other rodents, especially the voles (Arvicolidai) . In
regard to Trichina it must not be forgotten that their presence
in rats is not uncommon in some parts of Europe; and this
circumstance may explain the recurrence of trichinosis (first in
hogs and then in man) in certain outlying districts. Only in
this way can the Cumberland outbreak in this country be
accounted for. Here I cannot dwell upon the subject, but in
this connection I may observe that Bakody has in a very con-
vincing manner described a new variety or species of Trichina,
found by him infesting the walls of the stomach and intestine
of rats. In the first instance he detected the worm in associa-
tion with the ordinary T. spiralis, but afterwards separately.
He also obtained it in fowls. The species should be called
Trichina BaTcodyii. Possibly the nematodes observed by Colin
in 1863 also refer to this worm. They occupied tubercles in
the liver of a rat. In regard to the beavers (Castoridce) it
appears that they harbor many species of round worms, and
also several flukes, but they do not appear to have been very
much studied. In Morgan's work on the American beaver
there is a notice in which it is stated that Dr Ely found a very
fine filamentous worm 40"' in length. This does not seem to
correspond with Ascaris castoris (Rud.). He also speaks of
large numbers of a slender white worm, 3" to 5" in length,
found in the peritoneal cavity, and referable to the genus
Filaria. This cannot be confounded with Trichocephalus
castori (Rud.). Moreover, he describes a strongyle (Sclero-
stoma) as infesting the colon, and especially the caecum. These
all appear to be new to science. The Fasciola hepatica is
occasionally found in the liver, but the most common helminth
of beavers is Amphistoma subtriquetrum. Specimens of this
worm may be seen in the British and Hunterian Museums.
As regards the porcupines (Hystricida) I have already men-
tioned the occurrence of a tapeworm in the common species.
The larval Pentastoma denticulatum has been found by Otto
attached to the surface of the lungs, and Redi, about two cen-
turies back, noticed small nematodes lodged in tubercles of the
oesophagus. The late C. M. Diesing obtained Trichocephalus
affinis from the intestines. So far as I am aware, little or
nothing has been said respecting the helminths of the Octu-
dontidce, Chinchillidce, and Cavida. Like other European

318

PARASITES OP ANIMALS

investigators I have dissected guinea pigs (Gavia apercea) with-
out finding any parasites ; but in Brazil a small species of
ascaris (A. uncinata) was found by Natterer in this animal and
also in the paca (Coelogenys paca) The agoutis (Dasyprocta)
harbor Trichocephalus gracilis.

The entozoa of the duplicidentate rodents (Leporidai) acquire
importance from the fact of their abundance and from the
intimate relation which some of them bear to parasites infest-
ing the dog and other animals. Thus, the two commonest
kinds of fluke infesting cattle (Fasc. hepatica and Dist. lanceo-
latum) also attack hares and rabbits ; the former parasite often
producing the rot disease, which is almost as fatal to the rodents
as it is to the ruminants. Mutual infection occasionally results
from this circumstance by the distribution of germs. All
experiment-conducting helminthologists have reared Tcenia
serrata from the Gysticercus pisiformis ; nevertheless, several
English Manuals of Zoology persist in propagating the old
error of Yon Siebold, who supposed he had reared this tape-
worm by the administration of Ccenuri. So far as I am aware,
no feeding experiments have been conducted with the Coenuri
of rabbits (G. cuniculi). These bladderworms infest the soft
parts of the body, often producing tumours having a very
unsightly appearance. For details I must refer to the papers
quoted below. The Norfolk warreners call the infested hosts
" bladdery rabbits." Though apparently most abundant in the
eastern counties of England, these diseased rabbits are by no
means confined to that quarter. Through Mr Alston's help I
have received specimens of Goenurus cuniculi from Ayrshire,
Scotland. Probably this form of Coenurus occurs wherever
rabbits live. In Italy a case is recorded by Perroncito from
the abdominal cavity of a rabbit (coniglio). Every experi-
menter is more or less familiar with the cestode larvae (G. pisi-
formis) found wandering in the abdominal cavity. These were
regarded as flukes by Kuhn (Monostoma leporis). I need
hardly remark that the developmental and structural changes
undergone by these Cysticerci during their residence within
the rabbit have been exhaustively followed out and treated of
by Leuckart. Without dwelling on this subject, I must in
justice add that in this relation the special labors of Kiichen-
meister, Van Beneden, Haubner, Wagener, Roll, Eschricht,
and Moller played no inconspicuous part. My own efforts in
1857, and subsequently, wore not unattended with success. It

RODENTIA

319

therefore seems to ine, without prejudice to the recent experi-
ences of De Sylvestre and others, that further experiments in
this immediate connection are unnecessary. As regards the
nematodes of leporine rodents, probably the most important is
Strongylus commutatus. This parasite, like its husk-produc-
ing congeners, infesting calves and lambs, occasionally sweeps
off great numbers of hares. Such an epizooty occurred in
Thuringia in 1864. The most frequent intestinal parasite of
rodents is probably Oxyuris ambigua, but Strong, retortarformis is
tolerably abundant in the hare, and Trichocephalus unguiculatus
is liable to occur in all leporines. I know nothing of the so-called
Strong, strigosus of rabbits, but Bellingham found it in Ireland.
Olfers and Natterer obtained a small ascaris (A. veligera) from
Lepus braziliensis ; but I cannot help thinking that the large
measle (Gysticercus macrocystis) described by Diesing as three
inches in length, and obtained from the same rodent, must
either have been Goenurus cuniculi or else another form of poly-
cephalous hydatid.

In reference to the ectozoa of rodents it may be said that
they are very numerous. Acari infest rats and mice, and espe-
cially leporines. Thus, in the mouse are found Sarcoptes
notoedre, Bourguignon, var. muris, Megnin, Sarc. musculinus,
Koch, and Myobia musculi, Claparede. It is not very gene-
rally known that wild rabbits are apt to be attacked by the
common autumnal spider (Leptus autumnalis), whence, as once
happened with myself, they may be transferred to the human
body. The ears of tame rabbits are sometimes covered with
acari, which are easily destroyed by the cautious application of a
mixture of carbolic acid and olive oil (one of acid to six of the
oil). Rodents also harbor fleas. At a meeting of the Ento-
mological Society in 1875 Mr Yernall showed living specimens
from the ears of a rabbit, and Messrs Cole and W. A. Lewis
stated that they had obtained fleas from the hedgehog and
European marmot respectively.

Bibliogeaphy (No. 47). — Benedcn (see Van Beneden below). —
Cajpelle, J.} Extr. from a letter, in which the author states that he
had " found worms of the teenia kind in the liver of sixteen out
of eighteen rats," ' Med. Commentaries/ vol. xix, p. 139, 1794;
see also ' Trans. Coll. Phys. of Philad./ vol. i, part ii, p. 60,
1793. — Chapman, H. G., " Echinorhynchus in Squirrel/' ' Proc.
Acad. Philad./ 1874, p. 7Q.—Gobbold, " Note on Ccenurus (from
a squirrel)/' ' Proc. Linn. Soc./ May 5, 1864.— Idem, " On the

320

FARAS1TES OF ANIMALS

occurrence of Tania pectinatain the Porcupine (Ilystrix dorsata),"
in a letter to Dr. Lawson in the ' Canadian Naturalist and
Geologist/ 1862.— Idem, ' On T. serrata/ &c. (see Bibl. No. 45).
— Colin, "On the presence of a Nematode Worm in certain
Tubei-cles of the Liver of a Rat," from ' Rec. de Med. Vet./ in
' Edin. Vet. Rev./ Oct., 1863.—LeucJcart, 'Die Blasenband-
wurrner (u. s. w.)/ 1858 (contains numerous details and figs,
in ref. to Cysticercus pisiformis and T. serrata, &c). — Marchi,
P., 'Mem. della R. Accad. d. Sci. di Torino/ xxv. — Peacock,
" Remarks on the Liver of a Mouse with Cysts containing
Cysticerci," ' Lancet ' and 'Trans. Path, Soc./ 1855. — Perroncito,
E., " Sopra un caso di Ccenurus (in the abdominal cavity of a
rabbit)," « Giornale Med. Veter./ 1876.— Siebold (see Von
Siebold, below). — Sylvestri, De, "Experiments with C. pisi-
formis," 'II. Med. Veterinario/ 1871. — Van Beneden (see Bibl.
No. 45). — Idem, "On Sciurus glacialis and its Parasites," from
'Bull, de 1'Acad. de Belgique/ in 'Ann. Nat. Hist./ vol. xiii,
1854.— Verrall, in ' Proc. Ent. Soc. Lond./ Feb. 15, 1875, p. 3.
■ — Von Siebold, ' Ueber die Band- und Blasenwiirmer/ Leipsig,
1854, and Huxley's edit, for Syd. Soc, 1857. — Idem, "Experi-
ments on the Transformation of the Cystoid Worms into
Taenias," from 'Ann. des Sci. Nat./ in 'Ann. Nat. Hist./
vol. x, 1852. — Idem, " Helminthology," trans, by Busk and
pub. in 'Ray Soc. Rep. on Zool./ 1813-44, p. 446, London,
1847. — Idem, "On the Transformation of Cysticercus pisiformis
into Tania serrata," from ' Zeitsch. f. w. Zool./ in ' Quart.
Journ. Micr. Sci./ 1854.

Part VII (Edentata).

The entozoa of the edentulate mammals are not very
numerous. So far as I am aware only one species has been
described from the scaly ant-eaters (Manidce). This is the
small and probably immature ascaris noticed by Whitefield in
the walls of the stomach of the badgareit or short-tailed pan-
golin (Manis pentadactyla). Amongst the true ant-eaters
(Myrmecophagida.1) a single round worm has also been observed,
but not adequately described. I allude to Marcgrav's "find" in
the little ant-eater (Myrmecophaga didacfyla). I observe that
Rudolphi distinctly refers to this edentate as the tamandua.
Diesing does the same. The ant-eaters are much infested by a

EDENTATA

321

Fig. 60. — Tail of the
male Ascaris retusa.
Enlarged. Original.

thorn-headed worm (Echinorhynchus echinodiscus). On the 1st
November, 1875, I received from Prof. Flower a jar labelled as
follows : " Entozoon found attached to intestine of tamandua
ant-eater." The parasite was procured from the society's
gardens on August 12th, 1871. Natterer originally' obtained
this worm from Myrmecophaga jubata and if. bivittata. Creplin
described it from a M. didactyla from Surinam ('Wiegmann's
Archiv/ 1849). I presume that M. tamandua answers to the
M. bivittata of Geoff roy, as well as to the tridactyle and tetra-
dactyle species of Linnasus. The parasite in question was a
female, measuring exactly 10 inches long, and
had its proboscis firmly anchored within the
gut. The armadillos (Dasypidce) entertain a
variety of nematodes. In 1858 I obtained
several examples of Ascaris retusa, from the
rectum of a poyou or weasel-headed armadillo
(Dasypus sexcinctus) . The worm was first pro-
cured by Natterer from the black armadillo
(D. peba), which host also harbors Penta-
stoma subcylindricum. According to the " finds"
of Natterer and the subsequent descriptions
by Diesing, the two most common helminths of the Brazilian
armadillos are Aspidocephalus scoleciformis and Trichocephalus
subspiralis. As regards the sloths (Bradypidce) it would
seem that they are particularly liable to entertain round
worms. The Ai (Bradypus tridactylus) is infested by Stron-
gylus leptocephalus, Spiroptera gracilis, Sp. anterohelicina, and
Sp. brachystoma; whilst the unau (Choloepus didactylus) harbors
the last-named species and also Sp.. spiralis. All these worms
have boon described by Molin, and, with the exception of the
two first named, were new to science when he wrote his well-
known monograph on the genus. They were collected by
Natterer. All the species infest either the stomach or intes-
tines, with the exception of Sp. spiralis. This singular worm,
like the closely allied Sp>. helicina, infesting the feet of birds,
has the habit of coiling itself amongst the tendons of the digits
of the hind limbs more especially.

Bibliography (No. 48). — Cobbold, "On some new Forms of
Entozoa," 'Linn. Trans./ vol. xxii, p. 365, 1859.— Idem, "List
of Entozoa," &c, < Proc. Zool. Soc./ March 26, 1861.— Idem,
" Notes on Entozoa," part iii, 'Proc. Zool. Soc./ Feb. 1, 1876,
p. 202. — Marcgrav} in his ' Historia rerum nat. Brasil./ 1648,

322

PARASITES OF ANIMALS

p. 226, and in ' Rudolphi's Synopsis/ p. 186. — Molw, "Una
Monografia del gen. Spiroptera," 'Aus dem Sitzungsb. d.
m.-nat. CI. d. k. Akad. d. Wissensch./ Bd. xxxviii, 1859, s. 911,
Wien, I860.— WUtefieU, in ' Edin. New. Phil. Journ./ edited by
Jamieson, 1829, p. 58.

Part YIII (Ruminantia) .

In the matter of parasites this order of mammalian animals
stands second in importance. An entire volume of the dimen-
sions of the present would barely do justice to the subject.
Although in the article " Ruminantia 31 in ' Todd's Cyclo-
paedia/ and in my popular treatise on the mammalia, I have
described the oxen (Bovida) and sheep (JEgoscerida) as separate
families, I shall here speak of their entozoa together ; and, at
the same time, I shall introduce occasional reference to the
helminths of the antelopes and gnoos (Antilopidce) , also of the
giraffes (Oamelopardida), the deer tribe (Oervidce), the camels,
and the llamas (Oamelidce). The parasites of the last family,
however, will necessarily stand somewhat apart.

Almost all ruminants harbor the liver fluke (Fasciola hepa-
tica). This worm has been found in every variety of the
common ox and zebu (Bos taurus, var. Indicus), in the sheep,
goat, and argali (Oct's aries, Oapra hircus, and G. argali), in
the antelopes and gazelle (A. dorcas), in red-deer, roe, and
fallow (Oervus elaphus, G. capreolus, and 0. dama), and in the
two-humped camel (Gamelus badrianus). A closely-allied but
much larger species of fluke (F. gigantea) infests the giraffe
(Gamelopardalis) . All these animals are more or less liable to
suffer from the "rot'''' which is produced by these flukes. Into
the history of the affection the space at my command does
not permit me to enter, but as regards the development of the
common fluke I believe the following conclusions to be tolerably
well founded. I had long entertained the opinion that our
common Planorbis plays the role of intermediate bearer, and
this view has at length received confirmation.

1. The liver fluke, in its sexually-mature state (Fasc. hepatic"),
gives rise to the disease commonly called rot; this affection
being also locally termed coathe (Dorsetshire, Devon), iles (Corn-
wall), and bane (Somersetshire). In France it is known as
the Gachexie aqueuse, and more popularly as pourritwre. In

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323

Germany the epidemic disease is called egelseuclie, and in a
more limited sense either die Faule or die Leberhrankheit.

2. The rot is especially prevalent during the spring of the
year, at which time the fluke itself and innumerable multitudes
of the free eggs are constantly escaping from the alimentary
canal of the bearer. The
germs are thus ordinarily
transferred to open pasture-
grounds along with the fasces
of the bearer.

3. As it has been shown by
dissections that the liver of
a single sheep may harbor
several hundred flukes, and
as, also, a single adult fluke
is capable of throwing off
several thousand eggs, it is
certain that any rot-affected
flock is capable of distributing
millions of fluke germs.

4. Such flukes as have
escaped the host per anum
do not exhibit active powers
of locomotion. Their slight
contractile movements, how-
ever, serve the purpose of
concealing them in the grass,
and probably aid in the
further expulsion of eggs,
which pass from the oviduct
in single file.

5. After the death of the
escaped flukes the further
dispersion of the eggs is faci-
litated by the subsequent
decomposition of the parent
worm, and also by its dis-
integration, partly occasioned
by the attacks of insects. It has been calculated that the
uterus of a full-grown fluke may contain upwards of forty
thousand eggs.

6. By the agency of winds, rains, insects, the feet of cattle

Fio. 01.— Fasciola hepatica. Enlarged. After
Blanchard,

324

PARASITES OF ANIMALS

dogs, rabbits, and other animals, as well as by man himself, the
freed ova are dispersed and carried to considerable distances ;
and thus it is that a considerable proportion of them ultimately
find their way into ponds, ditches, canals, pools of all kinds,
lakes, and running streams.

7. At the time of their expulsion the eggs exhibit a finely
segmented condition of the yolk. The egg-contents continue
to develop whilst outside the parent's body, the granular matrix
finally becoming transformed into a ciliated embryo, which
when set free follows the habit of infusorial animalcules in
general by swimming rapidly in the water. The escape of the
embryo is effected at the anterior pole of the egg-shell, which
is furnished with a lid that opens in consequence of the action
of prolonged immersion, aided by the vigorous movements of
the contained embryo.

8. The ciliated, free-swimming embryo, at the. time of its
birth, exhibits the figure of an inverted cone, its anterior extre-
mity, which is broad and somewhat flattened, supporting a
central proboscis-like papilla. A small pigment spot placed
dorsally, and having the form of a cross, is supposed to be a
rudimentary organ of vision. After the lapse of a few days the
cilia fall off, the embryo then assuming the character of creeping
larvse (planulse).

9. Notwithstanding its abridged locomotive powers the non-
ciliated larvae sooner or later gain access to the body of an
intermediary bearer, within or upon whose tissues it becomes
transformed into a kind of sac or sporocyst. In this condition
the larva is capable of developing, agamogenetically, other
larvas in its interior. The sporocysts are highly organised,
forming redia. According to Willemoes-Suhm, the redia of
Fasciola hejpatica lives on the body of Planorbis marginata.
This organised nurse, which is about a line in length, is the
Cercaria cystoplwra of Wagener. The progeny of this redia
consists of armed Cercarise, which after a time quit the nurse
to pass an independent existence in the water.

10. In the cases of some species of fluke there is reason to
believe that before the Cercaria} gain access to their final or
definitive host they re-enter the bodies of the mollusks. This
they accomplish by means of a boring apparatus, aud having
previously cast off their tails they encyst themselves beneath
the surface of the skin. In this new situation they develop
into the so-called pupa, which is at length passively transferred

KUMINANTIA

325

with the fodder, or drink, to the digestive organs of the host.
In the case of Fasc. hepatica, as probably obtains also with
many other flukes, I think there can be no doubt that the Cercarias
pass directly into the bodies of ruminating animals. The cir-
cumstance that flukes of this species have been found beneath
the human skin shows how considerable are the boring powers
of the armed Cercarise.

In regard to the possibilities of fluke development, that will
be best understood by glancing at the constitution of the zoolo-
gical individual. The sum total of the products of a single
germ may be tabulated as follows : —

Zoological individual (Fasciola hepatica).

a. Ovum in all stages,

b. Ciliated free-swimming embryo,

c. Nurse, germ-sac, sporocyst (redia),} Second "biotoine."

d. Active, migrating, tailed larva (cercaria),

e. Encysted, resting larva (pupa),

f. Sexually-mature fluke (fasciola).
This is a fair representation of the life-phases of the fluke.

The life-phases are rarely less numerous or complicated than
here indicated, but Pagenstecher's researches tend to prove that
under certain climatal conditions the number of larval forms
may vary considerably. In other words, the fluke individual
does not comprise any definite number of " zooids," although
the kinds of zooids are limited. I recognise three (C biotomes."
The first includes only one temporary, independent life-phase,
this is the ciliated animalcule, which I call a " protozooid."
The second " biotome " may comprise only a solitary simple
sporocyst or germ-sac (deuterozooid), but an almost indefinite
multiplication of new and independent germ- sacs, as well as
other more highly organised " nurse formations," may also be
developed from the primary sporocyst (secondary and tertiary
"deuterozooid"). The third "biotome" embraces a large but
variable number of " tritozooids " (cercaria), an equal number,
whatever that may be, of " tetartozooids " (pupee), and, there-
fore, also, a similar number of " pemptozooids " (flukes).

Practically, other curious results arise out of the foregoing con-
si derations. For example, a single sheep may harbor 1000 flukes.
Each fluke will develop 10,000 to 40,000 eggs. Each egg
may give rise to 370 zooids. It thus appears that, if all the
conditions were favorable, a single fluke might originate between

\ First "biotome."

1 Third
| " biotome."

320

PAEASITES OK ANIMALS

three and four millions of individualised life-forms, whilst the
solitary sheep itself would, under the same circumstances, be the
means of causing the production of at least 3,000,000,000 fluke
zooids ! Happily, no such results as this can possibly occur in
nature, since interfering agencies reduce the favorable conditions.
However, the balance of parasitic forms from all sources is
usually sufficient to destroy thousands of sheep annually. The
virulence of rot-epizooty is entirely due to the presence of con-
ditions favoring the development of fluke larvae.

As regards the injurious action of this parasite on animals,
it is well known that in particular years, in England alone,
hundreds, and even thousands, of sheep have been destroyed in a
single season. A writer in the ' Edinburgh Veterinary Review '
for 1861 states that in the season of 1830-31 the estimated
deaths of sheep from rot was between one and two millions.
This would, of course, represent a money loss of something like
four million pounds sterling. As affording additional striking
instances of the disastrous effects of rot, I may cite the statements
of Davaine. Thus : — " In the neighbourhood of Aries alone,
during the year 1812, no less than 300,000 sheep perished, and
at Nimes and Montpellier 90,000. In the inner departments,
during the epidemic of the years 1853-54, many cattle-breeders
lost a fourth, a third, and even three fourths of their flocks."
In like manner our English authority, Prof. Simonds, fur-
nished a variety of painful cases. Thus, on the estate of Mr
Cramp, of the Isle of Thanet, the rot epidemic of 1824 " swept
away £3000 worth of his sheep in less than three months, com-
pelling him to give up his farm." Scoi'es of cases are on
record where our English farmers have individually lost three,
four, five, six, seven, and even eight hundred sheep in a single
season ; and many agriculturists have thus become completely
ruined.

Remarkable periodic outbreaks of this disease are recorded by
Simonds as occurring in England in the successive years of 1809,
'16, '24, '30, '53, and '60 ; whilst, for France, Davaine mentions
1809, '12, '10, '17, '20, '29, '30, '53, and '54, as the most remark-
able years. It would be interesting to know how far these out-
breaks tally with the similar outbreaks which have occurred in
Holland, Germany, and other European districts. The disease
was prevalent during four separate years in France and England
at one and the same time. This, indeed, is no more than we
would naturally expect, considering that the extent of the

RUMINANTIA

327

development of the larval forms must, in a great measure, be
dependent upon atmospheric conditions. A warm and moist
season would alike prove beneficial to the development of the
larvEe and their intermediate molluscan hosts. Their numbers
would also multiply enormously; for, as already remarked, the
degree of non- sexual production of trematode larvae within their
sporocysts is materially affected by climatic changes. On the
other hand, a fine, dry, open season will tend to check the
growth and wanderings of the larvae, and thus render the flocks
comparatively secure.

Considerations like these sufficiently explain many of the
crude theories which were early propagated concerning the
causes of this disease, and in particular, the very generally pre-
valent notion that water, and water alone, was the true source
of the disease. Intelligent cattle-breeders and agriculturists
have all along observed that the rot was particularly virulent
after long-continued wet weather, and more especially so when
there had been a succession of wet seasons. They have likewise
noticed that flocks grazing in low pastures and marshy districts
were much more liable to invasion than sheep which pastured
on higher and drier grounds, but noteworthy exceptions
occurred in the case of flocks feeding in the salt-water marshes
of our eastern shores. The latter circumstance appears to have
suggested the common practice of mixing salt with the food of
sheep and cattle, both as a preventive and curative agent ; and
there can be little doubt that this remedy has always been attended
with more or less satisfactory results. The intelligible explana-
tion of the good effected by this mode of treatment we shall
find to be intimately associated with a correct understanding of
the genetic relations of the entozoon, for it is certain that the
larvae of Fasciola hepatica exist in the bodies of fresh-water
snails. As already hinted from Willemoes-Suhm,s observations,
it is not improbable that the larvae are confined to gasteropod
mollusks belonging to the genus Planorbis.

The symptoms produced by rot are very striking. When the
disease has far advanced it is easy to know a rotten sheep, not
only by its very look, but still more convincingly, as I have myself
tested, by slightly pressing the hand over the region of the
loins. In this region the diseased animal is particularly weak,
and the pressure thus applied instantly causes it to wince. At
the same time the hand feels a peculiar sensation very unlike
that communicated by the spine of a sound animal. In bad

328

PARASITES OF ANIMALS

cases the back becomes hollow, and there is a corresponding
pendulous condition of the abdomen. The spinal columns ulti-
mately stick out prominently, forming the so-called " razor -
back/' As Professor Simonds has well observed, in an earlier
stage of the disease, " an examination of the eye will readily
assist in determining the nature of the malady. If the lids are
everted it will be found that the vessels of the conjunctiva are
turgid with pale or yellowish colored blood, the whole part
presenting a peculiar moist or watery appearance. Later on,
the same vessels become blanched and scarcely recognisable."
The skin also becomes harsh and dry, losing its natural tint,
and the wool is at length rendered brittle, either becoming very
easily detached or falling off spontaneously.

The first thing noticeable in dissecting a rotten sheep is the
wasted and watery condition of all the tissues. There is a
total absence of that firm, fresh, carneous look which so dis-
tinctively characterises the flesh in a state of health. Not only
is the rigidity and firm consistency of the muscles altogether
wanting, but these structures have lost that deep reddish color
which normally exists. When the abdominal cavity is opened
a more or less abundant, clear, limpid, or yellowish fluid will
make its escape, and the entire visceral contents will, at the
same time, display a remarkably blanched aspect. These
pathological changes are also shared by the important organ
especially affected, namely, the liver. This gland has lost its
general plumpness, smoothness, and rich, reddish-brown color,
and has become irregularly knotted and uneven both at the
surface and the margins, its coloring being either a dirty
chocolate brown, more or less strongly pronounced at different
parts, or it has a peculiar yellowish tint, which in places is very
pale and conspicuous. To the feel it is hard and brawny, and
when incised by the scalpel, yields a tough and, in places, a
very gritty sensation. On opening the gall-ducts a dark, thick,
grumous, biliary secretion oozes slowly out, together with
several distomes, which, if not dead, slowly curve upon them-
selves, and roll up like a slip of heated parchment. On further
slitting open the biliary passages, they are found distended
irregularly at various points, and in certain situations many
flukes are massed together, having caused the ducts to form
large sacs, in which the parasites are snugly ensconced. The
walls of the ducts are also much thickened in places, and hard-
ened by a deposit of coarse calcareous grains on their inner

RUMINANTI A

329

surface. Mr Simonds says, that the "coats of the ductus
hepaticus, as also of the ductus communis choledicus, are not
unfrequently so thick as to be upwards of ten times their
normal substance, and, likewise, so hard as to approach the
nature of cartilage." Eespecting their numbers, the greatest
variation exists. The presence of a few flukes in the liver is
totally insufficient to cause death ; consequently, when a sheep
dies from rot, or is killed at a time when the disease has
seriously impoverished the animal, then we are sure to find the
organ occupied by many dozen, many score, or even several
hundred flukes. Thus from a single liver Bidloo obtained 800,
Leuwenhoeck about 900, and Dupuy upwards of 1000 speci-
mens. Even the occurrence of large numbers only destroys the
animal by slow degrees, and, possibly, without producing much
physical suffering, excepting, perhaps, in the later stages.
Associated with the above-described appearances, one also not
unfrequently finds a few flukes in the intestinal canal, whilst a
still more interesting pathological feature is seen in the fact
that the bile contained in the liver ducts is loaded with flukes'
eggs. In some cases there cannot be less than tens or even
hundreds of thousands. Not a few may also be found in the in-
testinal canal and in the excrement about to be voided. Occasion-
ally dead specimens become surrounded by inspissated bile, and
gritty particles deposited in the liver ducts, thus forming the
nuclei of gall-stones. Mr Simonds mentions a remarkable
instance, " where the concretion was as large as an ordinary
hen's egg, and when broken up was found to contain about a
dozen dead flukes. It was lying in a pouch-like cavity of one
of the biliary ducts.'"

In respect of treatment we all know that " prevention is
better than cure." Moisture being essential to the growth and
development of the fluke-larvse, it is clear that sheep cannot be
infected so long as they remain on high and dry grounds, and
even in low pastures they can scarcely take the disease so long
as they are folded, and fed on hay, turnips, and fodder pro-
cured from drier situations. When once the malady has
become fairly developed, internal remedies are of little avail, at
least, in view of producing a thorough cure. Palliative treat-
ment may undoubtedly do good, especially in cases where the
disease is not very strongly pronounced. The most important
thing is the transference of the rot-affected animals to dry
ground and good shelter, supplying them, at the same time,

-330

PARASITES OF ANIMALS

with a liberal quantity of manger food, such as beans, peas, and
other leguminous seeds. The fodder, of whatever kind, should
be frequently changed, and many other hygienic measures
adopted, all tending to promote the appetite and general health
of the animal. An admixture of salines is a matter of essential
importance, especially in cases where the disease is not far
advanced. The beneficial effect of salt is one of those few
points on which nearly all parties are agreed, and its preserva-
tive influence in the case of sheep fed upon salt-water marsh-
land has been previously explained. In regard, however, to
the legion of remedies which have from time to time been pro-
posed, all I need here say is, that most of them when fairly
tested have been found to fail ignominiously. Every year we
hear of the adoption, often with enthusiasm, of new so-called
specifics, or of ancient medicines whose employment had long
fallen into disuse. Thus, for example, in the April number of
the ' Journal des Veterinaires du Midi 1 for 1 860, we find M.
Raynaud strongly recommending soot, in doses of from one to
three spoonfuls, to be followed up by the administration of a
grain of lupin for tonic purposes. In like manner, we received
from France wonderful accounts of the medicinal virtues of a
certain foetid oleaginous compound, the value of which was put
to a fair test by our distinguished veterinarian, Professor
Simonds. Having with infinite care and trouble undertaken a
series of experiments with the remedy in question, Mr Simonds
writes in the c Scottish Farmer and Horticulturist ' to the effect
that, as a result of his inquiries, he fears " we must conclude that
this supposed cure of rot in sheep has proved quite ineffective
for good." The last new " cure " announced is by Mr Robert
Fletcher (' Journ. Nat. Agric. Soc. of Victoria/ Dec, 1878).

The examination of rotten sheep is not altogether free
from danger. Professor Simonds tells us that in August,
1854, " a person of intemperate habits, following the occupa-
tion of a country butcher, was employed in skinning and
dressing a number of rotten sheep on the premises of a farmer
in the county of Norfolk. The sheep were necessarily opened
when warm, and while he was so engaged he complained
greatly of the sickening smell. The same evening he was
attacked with choleraic disease, and two days afterwards was a
corpse." This case is highly instructive and, when taken in
connection with the well-known fact that animals affected with
the disease putrefy very rapidly, clearly points to the neces-

It UM IN ANT I A

331

sity of removing slaughter-houses far away from densely popu-
lated localities.

Notwithstanding the above statement, there is little or no
danger to be apprehended from the consumption of the flesh of
rot-affected animals. On this vexed question -we have the
strong testimony of the late Dr Rowe, of Australia, who, after
leaving the medical profession, became a large and successful
stockowner, and devoted himself especially to this question.
Dr Rowe, writing from the Groulburn district, said : — The mere
presence of flukes in the viscera of an animal is no proof that it
is unfit for human food. For inspectors of slaughter-houses to
adopt such a test of wholesome food would be the greatest
mistake. It would afford no protection to the public against
unhealthy food, would increase the price of animals, and be
ruinous to our farmers and graziers. If the consumption of
flukey beef and mutton were prejudicial to the health of man,
there would be very few people alive in this part of the colony;
for, to my certain knowledge, they have had no other animal
food to live upon for the last twenty-five years, yet for physical
ability I believe they may be favorably compared with the
inhabitants of any other part of Australia." Speaking of his
own experiences, Dr Rowe avers that he found the common
liver fluke in sheep, cattle, goats, opossums, kangaroos, geese,
ducks, and other creatures, but he had never encountered it in
men, dogs, or pigs. On the whole I think we may agree with
Dr Rowe, in regarding the consumption of the flesh of rot-
affected animals as free from danger provided only the meat, be
well or even moderately well cooked. It must be borne in
mind, however, that an essential objection to its consumption
lies in the fact that the watery and otherwise chemically
deteriorated flesh is comparatively innutritious. It must also
be noted that the meat-supply from fluke-affected animals, as
usually sold in the markets, is chiefly derived from animals
which have only entered the early stage of the disorder, that is,
long before the watery and wasted condition of the muscles
has fairly set in.

Respecting the other trematodes I have to observe that
Distoma lanceolatum not only infests the liver ducts of cattle
and sheep, but also the deer tribe. Its larvee are likewise
supposed to reside in Planorbis marginatus. Still more common
and widespread amongst ruminants is the Amphistoma conicum,
occupying the paunch. It has been found in the ox, sheep,

332

PARASITES OF ANIMALS

musk-ox, elk, roe, fallow, red-deer, goat, and doreas-antelope ;
also in Gervus campestris, G. nambi, G. rvfus, and G. simplicornis.
Prof. Garrod has also recently shown me examples from the
sambu deer of India (0. Aristotelis) . Diesing'g A. lunatum,
infesting Gervus dichotomus, is inadmissible. Two other species
of Amphistome (A. explanatum, A. crumeniferum) are said to
infest the zebu ; and I have described another (A. tuberculatum)
from the intestines of Indian cattle. An aberrant amphis-
tomatoid entozoon (Gyrocotyle rugosa) has been found in a Cape
antelope (A. pygarga). Of more interest, however, is the cir-
cumstance that Dr Sonsino has discovered a species of Bilharzia
(B. bovis) in Egyptian cattle and in sheep. The eggs of this
species are distinctive, being fusiform and narrowed towards
either pole.

Comparatively few tapeworms are found in ruminants. Cattle
are infested by Taenia expansa and T. denticulata, the former of
these two species being also more or less prevalent in sheep,
antelopes, and deer. Other alleged species (Taenia fimbriata
and T. caprae) appear to me more than doubtful. Unquestion-
ably the common Taenia expansa is capable of giving rise to
severe epizooty among lambs. The privately communicated
evidence of Professors Brown and Axe, and published evidence
supplied by Messrs Cox and Robertson on this head, are con-
clusive. Mr George Rugg has also (in a letter to Prof. Simonds,
dated Dec. 4th, 1878) communicated the particulars of an out-
break in which " large numbers of lambs perished rapidly" from
tapeworms in the intestines, the parasites varying from one to
five or six feet in length. This tapeworm (T. expansa) is also
very prevalent in Germany. Ruminants, however, both at home
and abroad, suffer much more severely from bladder- worms.
Of these, Echinococcus veterinorum, Cysticercus tenuicollis, and
Goenurus cerebralis, are not only shared alike by all varieties of
cattle, sheep, and goats, but they also infest the deer tribe,
antelopes, the giraffe, and even camels. In 1859 I obtained
the slender-necked hydatid from a spring-bok (Gazella). Besides
these larval cestodes, cattle are very liable to harbor measles
(Cysticercus bovis), whilst sheep also entertain an armed Cysti-
cercus (G. ovis). I cannot again dwell at any length upon the
source of these immature helminths, but I may remark upon
the extreme frequency of measles in Indian cattle. This is
explained by the careless habits of the people. They not only
consume veal and beef in an imperfectly cooked state, but when

RUM INANTIA

suffering from tapeworm no precautions are taken to prevent cattle
from having access to the expelled proglottides of Tcenia medio-
canellata. The subject has already been dealt with in the first part
of this work, and also in my 'Manual/ quoted in the bibliography.
The mutton measle is described under the heading of Tcenia
tenella. In like manner I must refer to the ' Manual' for a detailed
account of the gid hydatid (Coenurus cerebralis) . How many kinds
of Ccenuri exist it is impossible to say, but I am of opinion that
the various polycephalous bladder-worms found by Rose, Baillet,
and Alston in rabbits, by myself in a lemur and in a squirrel,
and by Engelmeyer in the liver of a cat, are referable to tape-
worms specifically distinct from the Tcenia coenurus of the dog.

It was in 1833 that Mr C. B. Eose, formerly of Swaffham,
Norfolk, discovered an undoubted example of polycephalous
hydatid in the rabbit, the parasite in question bearing a very
close resemblance to Goenurus cerebralis. As the accuracy of
Rose's determination respecting the characters of the hydatid
has been called in question, I again invite attention to the
original description as recorded in the ' London Medical Gazette 1
for November 9th, 1833. At page 206, vol. xiii, of that perio-
dical, after describing the common Coenurus cerebralis of the
sheep, Rose writes : — " This {i.e. G. cerebralis) is the only
species of Coenurus noticed by authors, but I have met with
another. It infests the rabbit, and I have found it situated
between the muscles of the loins. It is also met with in the
neck and back. This hydatid grows rapidly, and multiplies
prodigiously, and being seated near the surface it soon projects,
and sometimes forms a tumour of considerable magnitude.
When the warrener meets with a rabbit thus affected, he punc-
tures the tumour, squeezes out the fluid, and sends the animal
to market with its brethren. I possess a specimen of this
species in a pregnant state. The earliest visible state of gesta-
tion is a minute spot, more transparent than the surrounding
coats of the parent; this enlarges till it projects from the
parietes of the maternal vesicle. It continues to enlarge until
it becomes a perfect hydatid, attached by a slender peduncle
only ; even whilst small, other young are seen sprouting from
it, and so on in a series of three or four. My specimen
exhibits them in every stage of growth, from a minute point co
a vesicle the size of a hen's egg. As I can see no difference
in structure between this hydatid and the last-mentioned (i.e.
Coenurus cerebralis), I am unwilling to consider it a different

334

PARASITES OF ANIMALS

species, for surely a varying locality ought not to constitute a
specific character."

The observations of Rose did not escape the well-known
Dutch author, Numan. In a foot-note to his memoir, entitled
" Over den veelkop-blaasworm der Hersenen," he makes the
following observations : — " Rose observes that he has found
Ccenurus in bladdery rabbits (blaaszieke konijnen) in the skin,
and in the cellular tissues of the trunk and extremities. The
veterinary surgeon, Engelmeyer, of Burgau, says he has also
found the G ceriums (Veelkop) in the liver of a cat {' Thierarzt-
liche Wochenschrift van 1850/ s. 192). These observations
differ thus far from those of other writers, according to whom
the Ccenurus is only found in the brain and spinal marrow.
However, it is not impossible in particular cases that some
parasites may have strayed from their ordinary dwelling-places."
Numan seems to have been not a little puzzled to account for
these discrepancies, and he was altogether undecided regarding
the mode of propagation of Coenuri and Cysticerci. This will
be gathered from the following passage, which I quote in the
original :

" Ik moet het onbeslist laten, of de grond begin sels, waaruit
de wormen uit de blaas ontspruiten, als wezenlijke of als zooge-
naamde kiemen (gemma) zijn te houden, waaromtrent de gevoe-
lens der voornamste Natuuronderzoekers, die zich met de
nasporing der blaaswormen hebben onledig gehouden, nog
uiteenloopen. Gulliver, door Rose (a. p. pag. 231) aangehaald,
houdt ze voor eijeren, in den Gysticercus tenuicollis, en Groodsir,
mede aldaar genoemd, spreekt ook van ova bij den Gcenurus
cerebralis ; doch de laatstgenoemde en Busk houden ze voor
gemma. Hier wordt voots gewezen op Owen en de meeste
onderzoekers van den tegenwoordigen tijd, die het daarvoor
houden, dat alle hydatiden zich alleen door gemma reproduceren.
Rose merkt voorts aan, dat, hetzij men de geboorte dezer inge-
wandswormen toekenne aan eijeren of kiemen (gemma), dit om
het even is, wat hunne verspreiding (dissemination) betreft,
daar zij ingesloten zijn, waardoor de wijze, hoe zij naar buiten
komen en verspried worden, tot dusver een gesloten boek is."

The idea of Numan that these are strayed forms of Ccenurus
cerebralis is not convincing. It must not be forgotten, however,
as Leuckart and Numan have both reminded us, that Eichler
discovered an hydatid about the size of a goose egg in the sub-
cutaneous tissue of a sheep. This bladder-worm supported

RUM1NANTIA

335

nearly two thousand heads. In regard to true hydatids or
acephalocysts in ruminants, on which subject I have already
dwelt at much length, I may again observe that the Hunterian
Museum contains some remarkable examples. In 1854 I
obtained Cysticerci from a giraffe, and I have reason to believe
that similar bladder-worms infest antelopes and deer.

The nematodes of the ruminants are both numerous in, and
destructive to, their bearers, those infesting the lungs being
productive of a parasitic bronchitis termed husk or hoose. In
cattle the lung-worm (Strongylus micrurus) is particularly fatal
to calves, whilst 8. filaria attacks sheep, and especially lambs.
A larger but less common lung strongyle ($. rufescens) is some-
times found associated with the latter. In 1875 I conducted
experiments with the view of finding the intermediate hosts of
S. micrurus, and I arrived at the conclusion that the larva3 of
this parasite are passively transferred to the digestive organs of
earth-worms. The growth and metamorphoses which I witnessed
in strongyloid larvae taken from earth-worms (into which I had
previously introduced embryos) were remarkably rapid, and
accompanied by ecdysis. The facts were as follows. -About the
middle of October, 1875, I received from Messrs Farrow, of
Durham, a fresh and characteristic specimen of diseased lungs,
in which the bronchi were swarming with Filariee.

In reference to the case itself, Mr George Farrow afterwards
informed me by letter that the calf was one of a herd of seven,
whose ages respectively varied from four to six months. At
the time of his writing (October 20th) the remaining six animals
were progressing favorably towards recovery — a result which
Mr Farrow attributes to the employment of inhalations of tur-
pentine and savin, combined with the internal adminstration of
tonics. In regard to this plan of treatment, and in reference
to the source of infection, he adds : — " I should have preferred
trying the inhalations of chlorine gas, but as the patients were
so very young and in poor condition, I deemed it advisable to
try a milder course of treatment.

" The history of the case is brief. The cattle are on a very
dry and well-drained farm, but during the summer there was
a great scarcity of water, and they were supplied from a stagnant
pool which eventually became dry. This, in my opinion, is
where the disease originated."

Mr George Farrow's opinion is probably correct, being in
harmony with the most recent results of scientific research as

336

PARASITES OF ANIMALS

made known more particularly by Leuckart. But the facts
thus conveyed do not explain the whole truth ; or, rather, they
convey it only in a very incomplete manner. Professor Leuckart' s
experiments were made with several species such as Strongylus
armatus of the horse, 8. rufescens, S. hypostomus, and S.filaria
of the sheep, and S. commutatus of the hare. Still, as regards
the strongyles, partial as the results have thus far appeared,
there cannot be a doubt that his successes with several allied
nematode species form a key by which we may yet unlock and
expose to view the entire life-history of that specially obnoxious
form under consideration, namely, Strongylus micrurus. To
summarise the whole matter in a few words, Leuckart supposes
that all these strongyloids require a change of hosts before they
can take up their final abode in the sexually-mature state. This
he infers especially because their respective embryos display
characters very similar to those exhibited by Olulanus. He
believes that either small mollusks or insects and their larva3
play the role of intermediary bearer. His experiments with the
embryos of Strongylus filaria prove that these larvae can be kept
alive for Several weeks in moist earth, and that whilst so
conditioned they undergo a first change of skin within a period
varying from eight to fourteen days. Experiments on sheep,
made with these moulting larvse, led only to negative results.
Unless the following facts be accepted, the scientific position
remains pretty much where Leuckart left it.

On the 22nd of October, 1875, at 1 p.m., I placed the entire
egg-contents of the uterus of a Strongylus micrurus on a glass
slide hollowed out in the centre. Probably something like ten
thousand ova were thus brought under observation, yet only
three were noticed as freed from their shells, probably as the
result of accidental rupture. Two of these displayed lively
movements. In round numbers the ova gave a measurement
of of an inch in length by ~ of an inch in breadth, whilst
the free embryos measured about ^ of an inch long, and less
than j^o of an inch in thickness. The integument of the embryo
displayed neither markings of any kind nor any double contour.
The contents of the worm were granular throughout, these
granules being crowded in the centre of the body, but scarcely
visible towards the head and tail, where for a considerable
space (fully fa") the worm was perfectly transparent. No trace
of any sexual organs or their outlets was visible. An examina-
tion of numerous eggs and free embryos obtained from near the

EUMINANTIA

337

primary bronchial bifurcations (of Mr Farrow's specimen)
yielded the same microscopic results, the only thing worthy
of remark being that the embryos from the mucus seemed much
more lively than those which, as I supposed, had accidentally
escaped their shells.

At 1.30 p.m. I placed some free embryos in two watch-glasses,
one containing water and the other saliva, and placed them before
the fire. Being called away professionally I found on my
return at 3 p.m. that evaporation to dryness had occurred in
the interval. All my attempts to resuscitate the embryos by
moisture proved unavailing, a result which, though negative,
proves how little capable these embryonic creatures are of
enduring desiccation. If these facts be confirmed, their prac-
tical significance is not without value in relation to the choice
of dry pasturage grounds for the rearing of young cattle. I
may add that whilst half an hour's immersion of the dried
embryos failed to restore any sign of life, the previous
warmth and moisture had caused many more embryos to
escape their shells during the time they were placed before the
fire.

At 4 p.m. I passed some very rich mould through muslin.
Some of this finely sifted earth I placed in a watch-glass, adding
a little water to moisten it, and also numerous eggs and free
embryos. In a wine-glass and also in a small jar I placed some
coarse earth with water added to make thin mud, and to both
of these I added, not only eggs and embryos, but also portions
of the reproductive organs of the adult female worms.

On the 23rd of October, at 2 p.m., I examined the contents
of these vessels. All the embryos in the vessels containing the
coarse earth were dead, but several were found alive in the
watch-glass containing the fine moist mould. Structually these
latter had undergone no perceptible change beyond a somewhat
closer aggregation of the somatic granules.

Although the embryos in the coarse wet mud had perished,
the eggs with unhatched embryos appeared to have retained
their vitality. Of this fact, indeed, I subsequently obtained
abundant proof ; and I also satisfied myself that the death of
the embryos had not resulted either from the coarseness of the
earth or from excessive moisture, but from the presence of
numerous shreds of the uterine tubes which I had somewhat
carelessly added to the vessels. Previous experiments, con-
ducted many years back, had indeed taught me that few if any

338

PARASITES OP ANIMALS

nematoid larv89 can resist the fatal action of putrid matter,
however slight the putrescence.

Having removed the offending shreds, I next placed a quan-
tity of living ova together in the earthenware jar, and allowed
the earth-contents to become much drier by evaporation before
the fire. I also left others in a watch-glass, which was placed
under a bell-jar enclosing several ferns.

On the 25th of October I removed particles of the moist earth,
altogether weighing about two grains, and, on submitting them
to microscopic examination, had the satisfaction to observe about
a dozen living embryos, some of which exhibited very lively
movements. There was not the slightest indication of putridity ;
nevertheless, I noticed several shreds of the adult worms whose
presence had been accidentally overlooked, and, curiously enough,
all the embryos subsequently removed from the immediate
neighbourhood of these decomposing shreds of tissue were
almost motionless and apparently in a moribund condition.
On examining the contents of the watch-glass placed under the
fern shade, I noticed several points of interest. First of all
the earth contained strongyle embryos, such as I had seen
before. Secondly, the surface of the mould was being traversed
by three or four briskly-moving Thysanurida, hunting about
with all that restless activity which Sir John Lubbock has so
well described. Thirdly, in marked contrast to the behaviour
of these I noticed several slow-moving Acariclce, apparently also
employed in searching for food. And lastly, while thus engaged,
the surface of the mould in the centre of the deep watch-glass
was suddenly upheaved, by which I was at once made aware of
the presence of another most welcome and unexpected intruder.
In short, an earth-worm had crept from the dry mould in which
the ferns were growing, and had taken up its temporary abode in
the soft moist experimental-earth contained in the watch-glass.
"When contracted, this Lumbricus terrestris was barely an inch
in length. On placing it under the half-inch objective glass, I
noticed a single embryonic strongyle adhering to the skin, but
not firmly, and evidently only in an accidental way, so to speak.
It was clear to me that it possessed neither the intention nor the
power to penetrate the chitinous integument of the earth-worm.

Having in the next place removed the Lumbricus with a pair
of forceps, and having washed it under a current of water, I
snipped off the lower end of the body, and allowed some of the
intestinal contents to escape on a clean glass slide for separate

EUMINANTIA

339

microscopic examination. Immediately, to my satisfaction, I
found that the faecal contents displayed a large quantity
of my strongyle ova, enclosing still living embryos, and in
addition several free embryos presenting characters which
declared that they were from the same source. Clearly they
had been ingested by the earth-worm along with its ordinary
food. One or two of the embryos were conspicuously larger
than their fellows, but the structural changes they had under-
gone were not so marked as to lead me for a single moment to
associate them with any of the various sexually- mature worms
which have been described as normally infesting the earth-worm.
I had no doubt whatever that such slight structural changes as
were now discernible had resulted from growth and develop-
ment consequent upon this accidental admission into the body
of the intermediate bearer which might or might not prove to
be its legitimate territory. It will be seen that subsequent
observations tended to affirm the truth of this view. I made a
careful examination of one of these larvae, whose active move-
ments were such as to render the process exceedingly tedious.
The earth-worm itself (or rather its unequal halves) was placed
in a fresh watch-glass containing ordinary mould. The larvae
or embryos obtained from the earth-worm now measured about
i of an inch in length, their heads exhibiting a short and
simple chitinous buccal tube, whilst their tails were somewhat
more pointed and bent upward. The somatic granules were
more crowded, rendering the position of the intestinal tract
more marked, though, as yet, the differentiation gave no indi-
cation of the formation of a distinct intestinal wall. There
was no perceptible increase of thickness of the body of
the embryos. The results thus far naturally encouraged
me to procure some fresh earth-worms for experimental
purposes.

On the 26th of October I found that the halves of the earth-
worm were alive, and I left them undisturbed in rather dry
mould, freshly added. To a watch-glass containing newly
sifted earth and embryos I added a fresh garden-worm, which
was rather sluggish from the cold ; and in the original jar I
placed another smaller and very active earth-worm obtained the
same morning. Finding the soil in the jar congenial, this
lumbricus soon buried itself. Another and larger earth-worm
subsequently added refused to follow this example. It was
therefore removed from the jar. Believing the fine and arti-

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PARASITES OF ANIMALS

ficially prepared soil to be still much too moist, I caused further
evaporation ; and I afterwards found that the thicker the mud
the more suitable it proved as a residence for embryonic nema-
todes and earth-worms alike.

On the 27th I found the small earth-worms in the jar bur-
rowing freely and throwing up faecal casts. From one of my
watch-glasses the worm had escaped, its place being occupied
in the meantime by an actively crawling Julus. I put a second
/ idiis, obtained from the mould in the fern jar, to form a com-
panion (in view of other experiments), and I also added a fresh
earth-worm, covering all by another inverted watch-glass, which
I thought would prevent their escape.

In the next place I examined the halves of my original
experimental earth-worm. They were scarcely capable of
motion, but retained a certain amount of vitality. The tail
was the more active half, and unfortunately it was soon after-
wards lost. Carefully washing the superior half, and transfer-
ring its contents to a glass slide, I immediately detected under
the microscope a large number of embryos. They were in a
state of marked activity, the largest having increased to about
of an inch in length, whilst their structure had become cor-
respondingly advanced. Here, again, there was no room for
doubt as to their source, especially as they individually dis-
played different degrees of organisation, all answering to one
and the same embryonal type. I now observed a distinct
oesophagus, the rest of the intestinal tract being still more con-
spicuous than heretofore, though, as yet, no true cells marked
the limitation of the stomach and chylous intestine.

After an hour's immersion in cold water some of the larvae
became much less active, whilst others were motionless, so that
I feared all were about to perish. In the hope of keeping a
few of them alive I now added to the slide some finely sifted
grains of mould, placing the slide under a small bell jar which
protected some of my ferns. The remains of the moribund
earth-worm were also covered with mould.

Other larvae, derived from the earth-worm, were placed on
the moist pinnae of a living fern-frond which supported small
drops of water, for by this process I hoped in some mea-
sure to imitate the dew which naturally condenses on the grass
and fodder of our low-lying fields. At 3.15 p.m. of the same
day (27th) I also examined a fresh worm pellet from the
jar, and found it to contain living strongyle embryos, which as

RUM1NANTIA

341

heretofore had not exhibited the slightest advance either in
respect of size or structure.

At noon on the 28th I again sought for the larger larvae,
first of all on the slide covered with fine earth, and afterwards
within the remains of the upper half of the original earth-worm.
On the slide I could detect none, but within the intestine of
the worm there were still two living larvae left, whose characters
corresponded precisely with the largest that I had previously
obtained from the same source only the day before. They had
undergone, however, no further change in structure, and their
measurements remained precisely the same.

At 12.30 p.m. I snipped off two or three of the terminal fern-
fronds on which I had placed a few advanced larvae. On
examination under the half-inch objective I immediately de-
tected one of the larvae cruising about most actively. On
adding a drop of water it soon rushed across the field of the
microscope, its movements being thoroughly eel-like. The size
of this larva had so much increased that it was now visible to
the naked eye, measuring, indeed, as much as ^" of an inch
from head to tail. Moreover, its organisation had advanced in
a marked degree. Thus, the digestive organs were better
defined, and on one side of them there appeared a regularly
arranged congeries of cellules, forming the commencement of
the reproductive organs. As yet, however, I could not pro-
nounce as to the sex.

At 1.45 p.m. I again examined a few grains of earth from
the jar, when I at once noticed five or six active embryos whose
structure failed to show the slightest advance upon that origi-
nally described. It was evident that the jar contained thou-
sands of them ; and since no ova were found, it became pro-
bable that all their embryonic contents had escaped to swell the
number of free larvae, leaving their very delicate envelopes to
perish. I think I had hit upon the most suitable degree of
moisture favorable to this result.

In the next place I sought for the earth-worm that had been
placed in the infested soil between two watch-glasses. It had
escaped. This obliged me to transfer the mould to a rather
wide-mouthed and open phial, in which four more fresh lumbrici
were placed. I feared the closing of the bottle would be
detrimental.

Later in the day I selected an earth-worm which had not
been exposed to strongyle infection, but which was in a mori-

342

PAHAS1TES OP ANIMALS

bnnd condition. In the intestine there were several free
neinatoids and also several psorosperuis of the genus Monocystis,
so well illustrated by E. Ray Lankester. As to the nematoids,
which were filariform, they neither corresponded in size nor
structure with my strongyle embryos.

At 1 p.m. on the 29th I renewed my examination of the
larva removed from the fern-pinnule. It showed a further
stage of growth, the male character of the reproductive organs
having become apparent. The now tolerably well-formed vas
deferens had pushed the chylous intestine on one side, whilst a
series of caudal rays, five on either side, supported two narrow
membranous wings, which represented the lateral lobes of the
hood of the adult strongyle.

At 1.30 p.m. I submitted the intestinal contents of four fresh
earth-worms removed from my garden to microscopic examina-
tion, but no nematoids were found in any one of them.

About 2 p.m. I removed another large and active strongyle
larva that had been reared on another fern-pinnule. It was
of the same size as that previously described, but was in the
act of changing its skin, It was then put aside along with the
other worm under the glass shade.

At 3 p.m. I intended to have examined one or more of the
earth-worms placed in the open-mouthed phial, but all had
escaped and buried themselves in the fern-mould out of reach.

At noon on the 30th I renewed my examination of the two
large larvee whose developmental changes I had been instru-
mental in producing from the time of their escape from the
egg-coverings. I saw no reason to doubt that the sequence of
changes thus far noticed referred to the species of parasite
under consideration. Both larvae were active, but the moult-
ing one had now completed its ecdysis. Its sexual distinctive-
ness had become yet more pronounced by the formation of
two rather short and stout spicules, the point of the tail dis-
playing a very minute awl-shaped projection. The lateral
membranes had not visibly increased in size. One of these
larvas, the first under observation, now perished from the injuries
sustained during inspection.

Again, and later in the day, I sought to clear up any doubts
that might still suggest themselves respecting the source of
these larvse, by once more submitting the intestinal contents of
two fresh and uninfected earth-worms to careful scrutiny. In
the first worm no parasite could be found, and in the secoud

EUMINANTIA

343

only one minute nematoid ; its organisation, which was sexually
incomplete, neither corresponded with my strongyle embryos,
nor, so far as I could judge, with Goeze's Ascaris minutissima
microscopica (the Anguillula lumbrici of Diesing and others),
nor with Dujardin's Dicelis filaria. It was a very long and
narrow creature, but I lost it whilst attempting to secure an
accurate measurement. I should say it was about —th of an
inch in length, and not more than ^^th. in breadth. I made a
rough outline sketch of it.

In view of further observations I now placed five more
earth-worms in the jar containing strongyle embryos, and I
also placed six others in the phial which contained coarser
mould, and only a comparatively small number of the original
strongyle embryos. The phial was closed with a cork and
half buried in the fern-mould of one of my larger Wardian
fern-pans. Before this transfer was made I again took an
opportunity of ascertaining by microscopic evidence that the
embryos lodged in the coarse and fine mould had none of them
made the slightest advance in organisation. The worms placed
in the jar immediately proceeded to bury themselves.

At noon on the 1st of November I sought to get further
results from the only large free larva which now remained to me
(for the fern-pinnules on which the larvaa were originally placed
had dried up and no third specimen could be discovered).
Structurally the larva presented no advance. It there-
fore appeared to me necessary to place it under new con-
ditions in view of exciting further progress towards sexual
maturity and adult growth. To transfer it to the bron-
chus of a living calf would, of course, have been the
crucial experiment, but the hopelessness of getting any satis-
factory result from this solitary transfer deterred me from the
attempt. On a larger scale, with many larvae, a positive issue
would of course prove decisive. Accordingly, the only thing I
could do, in partial imitation of nature, was to try and induce
some further changes by placing the larva in human saliva,
kept warm artificially. As a first step I immersed the creature
in a little of the secretion added to the glass slide, when it
immediately displayed very lively movements, such as could
only be fitly described as frantic. This encouraged me to
replace the slide under one of the fern shades without applying
any additional heat. I then left it.

At 12.30 p.m. I selected three of the eleven worms lodged in

344

PARASITES OF ANIMALS

the infested earth, namely, two from the jar and one from the
closed phial, and made a microscopic examination of their
respective intestinal contents. In one of the worms from the
jar I found several embryos clearly referable to my strongyles,
their structure showing scarcely any advance upon that exhi-
bited by the embryos in the mould itself. The weather was
now excessively cold and the larva) were motionless ; neverthe-
less, the application of warmth showed that they were by no
means dead. The faecal matter obtained from the worm that
had lived in the phial displayed an immense number of
infusoriaa {Bacteria) which rushed about rapidly over the field of
the microscope. No other signs of life were detected.

On the 2nd of November I found my solitary strongyle larva
alive, but its movements, though active, were by no means so
active as on the previous day. No fresh structural changes
had occurred.

At noon on the 3rd the larva at first lay almost motionless
in the now thick and ropy saliva ; nevertheless, on applying a
thin glass cover its movements became tolerably vigorous.
During its quiescent state I succeeded in getting a good view
of the caudal rays and other imperfectly developed organs> of
which I retain figures.

Having now satisfied myself that other new conditions were
necessary to enable the larva to arrive at sexual maturity, I
sought to transfer it to a glass tube filled with fresh saliva. This
transfer was a matter of difficulty. After passing the thick
ropy saliva into the tube, I examined the slide and found that
the larva was gone. I concluded it was in the tube, which,
in order to keep the contents warm, I subsequently carried
about concealed in my under- clothing during the day and placed
in my bed during the night. This increase of temperature,
however, caused decomposition of the saliva ; so when next day
I diligently sought for my experimental nematode it was no-
where to be found. Thus terminated my observations on the
first set of embryos, which had enjoyed their temporary sojourn
in the intestinal tract of the earth-worm, and which had certainly
afterwards undergone a series of marked structural and mor-
phological changes, accompanied with ecdysis.

The weather had now been for several days exceedingly cold,
but on the 4th a favorable change set in, which led me to hope
that I might be able to verify the facts above recorded.
Accordingly, as a new point of departure, I re-examined the

la MINANTIA

:; 15

fine mould, and at once found my embryos in a high state of
activity. The mould, however, appearing too moist for the
earth-worms, I permitted further evaporation before closing the
jar with a glass cover. Four days subsequently I examined
the intestinal contents of two of the earth-worms. In one of
these, an inch in length, no parasite of any kind could be de-
tected ; but in the other, which was beyond three inches in
length, there were numerous Opalina besides several strongyle
embryos, the latter presenting characters not visibly in advance
of those still living in the mould. All of them were motionless,
as if they had not got over the shock produced by previous
cold. Moreover, the weather had again become cold, and thus,
when I again inspected my experimental embryos living in the
jar, I also found them motionless, so different from their
behaviour on the 4th. However, since a further result with
the earth-worm embryos appeared possible, I placed some of the
faecal matter, already ascertained to contain a few of them, on
the fronds of a thoroughly moist and dew-covered Asplenium
bulbiferum. This plant was in a fern-pan which had the advan-
tage of considerable fire- warmth during the day. When, how-
ever, on the 15th of November, I examined the faecal earth
removed from several of the pinnules, I failed to find any of
the embryos. Possibly they had wandered, for the entire
frond was covered with dew-drops, which was not the case with
the fern that I had previously experimented on with such satis-
factory results. At all events, whether they had wandered or
had perished, their apparent absence in no way affects my pre-
vious record ; and the more so since only a few had been observed
in the faecal matter. The smallness of the number found in the
earth-worm was also readily accounted for. Thus, when at 1.30
p.m. on the 15th I made a diligent search for embryos in several
grains of the fine mould, not a single young strongyle could be
detected. Possibly the frost of the previous night had killed
them. The earth-worms were still alive and in good condition.

In conclusion, I may observe that every experimenter with
helminths is well aware how unfavorable the winter season is
for this kind of research. If a repetition of this inquiry in the
spring or summer should confirm these results, it will prove a
clear and substantial addition to our knowledge of the develop-
ment of the strongyles. Meanwhile, I think that the data
above given render it highly probable that the larvae of the
hoose-producing strongyle {8. micrurus) are passively trans-

346

PARASITES OF ANIMALS

forred to the bodies of setigerous annelids, which are thus called
upon to act as intermediate hosts. If this be so, it is further
certain that important structural changes with ecdysis follow
after their escape from the earth-worms or other annelids,
moisture, dew, or water being essential to the penultimate
stage of growth. Final passive transference, either with
fresh fodder from swampy grounds, or, it may be, from pond
water, ultimately enables them to acquire their definite sexual
form, size, and other adult characteristics.

According to Megnin it is not the Strongylns filaria, but a
hitherto unknown and totally distinct species (Strongylus minu-
tissimus) which occasions pneumonia in Algerian sheep. In
England the parasitic bronchitis affecting sheep is generally
called the "lamb disease." This is unfortunate, because many
other parasites prove destructive to lambs. One of the most
injurious species is Strongylus contortus, infesting the true
stomach, whilst 8. hypostomus, occupying the small intestines,
is almost equally obnoxious to the ovine bearer. By Leuckart
and others this last-named worm is retained in Dujardin's genus
Dochmius, in which genus another species occurs (D. cemuus).
This worm is quite distinct, but not readily distinguishable by
the naked eye alone. It occasionally occupies the upper part
of the colon, as well as the lower end of the small intestine.
A rarer intestinal worm in lambs is the Strongylus filicollis.
Several other strongyles infest the ox (8. radiatus, S. inflatus,
8. gigas), goat (8. venulosus), and stag (8. ventricosus).

As showing the extraordinary prevalence and destructiveness
of entozoa in certain countries, I will adduce an instance in
which my opinion was requested and given some five years
since. My informant stated the case somewhat in the following
manner : — On a farm in New South Wales, and lying about 200
miles to the north-west of Sydney, on the Trafalgar tributary
of the Macquarie river, out of a flock of about 8000 sheep no less
than 1200 have perished. In many instances post-mortem
examinations were made, worms appearing in all cases to be the
cause of death. There were four kinds of parasites present.
The most numerous were red and white, "marked like a
barber's pole." These occurred chiefly in the fourth stomach
and commencement of the duodenum, but some were found
throughout the entire length of the small intestine. A second set
comprised small black worms, resembling needles, scattered only
in the lumen of the intestines. The third set were tapeworms,

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347

each being several fathoms in length. The fourth set was made
up of white threadworms, individually measuring two inches in
length. These occupied the bronchial tubes, and where charac-
terised by my informant as " the most deadly of all." Without
the aid of specimens I at once recognised these brief diagnostic
characters as severally referring to Strongylus contortus, Boch-
mius hypostomus, Tcsnia expansa, and Strongylus filaria.

What the inquirer desired at my hands was " full informa-
tion respecting the general principles to be carried out in view
of the prevention of this parasitic disease, regard being had to
the difficulty of finding any food but pasture, to the number of
animals to be treated, and to the not unfavorable circumstance
that the run is divided by fencing to a great extent." I was
also requested to explain the best modes of treatment, being at
the same time informed that turpentine drenchings had already
been employed with only "partially effective" results. I was
also expected to give numerous and varied formulae, to be tried
in succession, supposing the first should fail. Of course, it should
have been known that I neither prescribe medicines nor accept
fees in respect of animal patients ; but, as in this instance
my opinion was permitted to assume the form of a " written
scientific report," I was pleased to have an opportunity of com-
menting freely and fully on the significance of the facts sub-
mitted. My advice took the form of a long report, which might
here be usefully given in extenso were it not somewhat of the
nature of a private and privileged communication. I have no
doubt that the stockowner would be pleased that I should
utilise his remarkable " case " for the benefit of agriculturists
and others ; but it is for him to publish the " opinion " as it
stands, should he think fit to do so.

Practical men, on reading the few foregoing particulars, will
perceive that one of the principal obstacles to success in cases
of this kind lies in the circumstance that artificial food can only
be procured with difficulty. Where the source of the disease
is associated with the pasture- supply, any treatment, however
effectual for a time, can only be followed by partially satis-
factory results.

The destructive powers of any one of the above-mentioned
parasites being sufficient to produce a fatal lamb-disease, it is
clear that when two or more of these particular species attack
their victim in considerable numbers, the ovine-bearer has
little chance of recovery. The intestinal strongyles, by means

348

PARASITES OF ANIMALS

of their oral armature, behaving as veritable leeches, will, if not
expelled in good time, produce a rapidly fatal anaemia, precisely
in the same way as the human Anchylostomum of the tropics.

The worst of dealing with this sheep -parasite is that it will
not succumb to ordinary doses of salines like the stomach stron-
gyle ; moreover, the little leech-like wounds will probably
bleed after the parasites have been compelled to abandon their
hold. Prevention is better than cure. Accordingly, I sought
to explain the origin of these creatures, and in what possible
ways the germs of the various species could be destroyed, or at
least limited in numbers.

As to the drugs and inhalations to be employed, it would
be difficult to advise any more effective than those commonly
in vogue, the great thing being to effect changes of pasture
and ground, to look to the purity of the water-supply, and to
supply the best kinds of nourishment after active treatment.
The diseased animals should, from the very first, be separated
from their companions, because the amount of germ distribution
is thereby greatly lessened. They should be at once drenched
or treated by inhalation (as the parasitic nature of the attack
requires), and the enclosure in which the animals have been
temporarily housed should be thoroughly scoured with boiling-
hot water impregnated with salt.

The nomenclature of the parasitic diseases of animals is
excessively vague. Thus, apropos to the case above recorded, I
may mention that an American veterinary practitioner appeared
to be much shocked that I should have had the temerity to speak
of four distinct kinds of lamb- disease. It is in this way that
practical men often commit serious mistakes by rolling together
disorders that are totally distinct. If it were true that epizooty
in lambs is exclusively due to Strongylus filaria, then profes-
sionals might aptly speak of the parasitic bronchitis of young
sheep as lamb-disease ; but we now know that several other
helminths prove terribly fatal to lambs, occasioning death in
totally different ways. In one set of cases the animals are
asphyxiated ; in another set they become fatally ansemic ; and
in a third set they perish from the severity of nervous reflex
irritations. Lastly, it may be remarked that, in view of the
successful management of the parasitic disorders of animals,
the veterinary practitioner must necessarily be guided by the
same general principles as the physician. For myself, I may
say that I have hitherto designedly withheld mauy practical hints

BDMINANTIA

349

which a long experience with human patients suggested, not
wishing to appear to dictate to those who are constantly seeing
animals. However, since (contrary to my own wishes) it has
happened that both professional men and agriculturists have
not only invited me to give opinions, but have, at various times,
asked me to prescribe, it seems there can have been no impro-
priety in publishing my views on this subject. Certainly I
have had no professional motives to serve.

Of the few non-strongyloid nematodes, one of the commonest
is Trichocejphalus ajjinis. I have obtained this worm from the
giraffe, and the parasite may be said to infest all ruminating
animals, not excluding even the camels and llamas. As before
remarked, the whipworm has been known to produce severe
symptoms in man, and it occasions " scour " in the sheep. The
eyes of cattle are occasionally infested by Filaria lacrymalis
and F. papillosa. The last named is the common eye-worm
of the horse. On Feb. 27th, 1875, Dr Edward L. Moss, of
H.M.S. " Alert," brought me three examples of a nematode
which I referred to Filaria terebra. Dr Moss obtained these
parasites in 1874, during the time that he had charge of the
Naval Hospital at Bsquimalt, Vancouver's Island. • They occu-
pied the abdominal cavity of the black-tailed deer (Cervus
columbianus) . The worms were mostly found lying amongst
the coils of the small intestine. They were not attached to the
peritoneal membrane. Dr Moss had shot seventeen deer in all,
the males and females being in about equal proportion ; never-
theless, not one of the bucks showed any trace of the presence
of these entozoa. This absence of parasites in the male deer
is noteworthy. Hitherto the worm appears to have been
observed in the red deer (0. elaphus), and by Natterer in three
species of American roe {0. rufus, G. simplicicornis, and 0.
nambi). Two of the worms measured each about 2\" in length,
the third exceeding 3". They displayed in profile two promi-
nent oral papilla?. Probably there were four of these processes,
such as Dujardin described in his Filaria cervina, which,
according to Diesing, is a synonym. They all possessed
spirally twisted tails.

Amongst the arachnidan parasites of ruminants having
entozoal habits are Pentastoma denticulatum and P. constrictum.
The former larval worm is excessively common in cattle, sheep,
deer, and antelopes. According to Khind, the adult worm
(P. tamioides) also infests the sheep. The P. constrictum has

350

PARASITES OF ANIMALS

hitherto only been found in the giraffe. On the 1 0th February,
1859, I obtained numerous examples (P. denticulatum) from a
bubale (Antilope bubalis) which died at the Zoological Society's
Gardens. The greater number occupied the surface of the
lungs and intestines ; some few, however, were enclosed in
cysts beneath the pleura. In the spring of 1860 I also pro-
cured several specimens from the abdomen of a cape guevi
(Oephalopus pygmceus).

The ectozoa of ruminants have received much attention, but
I can merely indicate the known forms. Following Megnin's
classification we have three well-marked varieties of the acarine
genus Sarcoptes ($. scabiei, var. ovis, var. caprae, and var.
cameli), two varieties of Psoroptes (P. longirostris, var. bovis
and ovis), and Ohorioptes spathiferus. This last is the true
mange mite of the ox (or Symbiotes bovis of Gerlach). A
variety of the follicle mite infests the sheep (Demodex follicu-
lorum, var. ovis). Numerous species of tick (Ixodida) have
been more or less fully described. Of these we have the Cara-
partos of the Portuguese (Ixodes bovis), attacking cattle ; the I.
reduvius, attacking sheep ; the I. plumbeus, said to attack
lambs ; the I. albipictus and I. unipictus, found on the moose-
deer. Probably this species also attacks cattle. A most
horrible arachnidan is found on camels. I allude to Galeodes
araneoides belonging to the Solpugida. This parasite will bite
severely any person who attempts to dislodge it from the bearer.
Turning to the insects, we find ruminants liable to be annoyed
alike by flies (Diptera), fleas (Aphaniptera) , and lice (Hemiptera).
Various species of four different families of flies are apt to prove
troublesome. Of the CEstridce, attacking the ox, we have Hypo-
derma bovis, whose larvae form tumours or warbles on the back ;
also H. lineata, Dermatobia noxialis, and Cephenomyia bovis
(mihi). The larvae of the latter reside at the root of the tongue
and adjacent parts. In the sheep we have CEstrus ovis, (E. pur-
pureus, and Hypoderma lineata. Various species also attack
goats and antelopes. Dr Kirk presented me with specimens of
CEstrus from the frontal sinuses of a harte-beest or caaraa, and
they have also been obtained from the sassabe, the saiga or coins,
from the gnoo, and from the brindled gnoo, kokoon or gorgon.
Mr Charles Danford presented me with several bots from an
ibex. One or more species of Hypoderma have likewise been
removed from the gazelle and other antelopes. The deer tribe
are much attacked by bots. In the red deer we have Hyp. actaon

RUMINANTIA

351

and H. diana, a species also infesting the elk. The throat-grubs
are Oeph. rufibarbis and Pharyngomyia picta ; another species,
also occurring in the fallow deer, O&ph. ulrichii, infests the elk,
and G. stimulator the roe, the last-named deer being also infested
by Hyp. diana. A throat- fly infests the reindeer, which is also
frequently attacked by Hyp. tarandi. Specimens of the latter
worm have been obtained by Dr Murie at the Zoological Gardens.
The Hunterian Museum also contains these and other species of
bots, presented by myself in Mr Andrew Murray's name. A
subcutaneous bot has been found in the musk-deer. A throat-
bot (G. maculata) infests the dromedary.

In regard to the so-called free dipterous parasites and other
noxious insects that attack ruminants, their name is legion.
One of the worst is the tsetse (Glossina morsitans), immortalised
by Livingstone. Of the Muscida we have the ox- fly (Musca
bovina), the sheep-fly (M. casar), and the executioner (M. car-
nifex). Of the Tabanidee we have T bovinus and T. autumnalis,
Chrysops ccecutiens, and the allied Asilus crabroniformis (Asi-
lidte). Amongst the specially noxious insects must also be
placed Stomoxys calcitrans and Rhagio columbaschensis. This
fly proves fearfully destructive to cattle in Hungary and Servia.
Lastly, I can only further mention the common Melophagus
ovinus. This is nothing more than a gigantic louse, which
from long use agriculturists and veterinarians persist in calling
the sheep-tick. It belongs to the Hippoboscidce, the members
of which family only attack quadrupeds and birds. As regards
the lice (Anoplura), I have to mention Heematopinus vituli of the
calf, H. eurysternus of cattle, and H. stenopsis of the goat ; also
Trichodectes scalaris, T. spheerocephalus, and T. caprce. These
infest the ox, sheep, and goat, respectively.

For some account of the protozoal parasites (Psorospermice,
&c.) infesting the flesh of ruminants I must refer the reader to
Book I, Section IV, Part VI of this treatise.

Bibliography (No. 49). — (Anonymous), "On the Hydatid in
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pp. 156-7, 1863. — (Idem), " Tapeworm Epizooty in Notting-

352

PARASITES OF ANIMALS

hamshire," ' Brit. Med. Journ./ 1858. — (Idem), " Prevalence of
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1862. — Idem, " The Whipworm of Ruminants," ibid., Dec,

1863. — Idem, " Parasite-larvae," ibid., March, 1863. — Idem, "On
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Nematoid Worms," 'Veterinarian/ Oct., 1876. — Idem, "Record
of preliminary Experiments with the Eggs and Embryos of the
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in 'Entozoa/ pp. 145-183; also in 'Manual/ and in the chap,
on " Parasitic Diseases " contributed to ' Williams' Principles
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searches," 'Veterinarian/ Dec, 1877. — Idem, ' Amphistomes of
the Ox/ see Bibl. No. 51. — Cooper, J., "Three Cases of
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vol. xxviii, p. 446, 1855. — Creplin, fig. of Amphist. crumeni-
ferum, in ' Wicgmann's Archiv/ 1847, tab. ii, s. 30.— Crisp, E.,

RUMINANT! A

353

« On the ' Lamb-disease/ of which Parasites in the Lungs are
generally the Cause or Consequence/' repr. from ' Journ. of
Bath and West of England Soc./ in July to October Nos. of
'Edin. Vet. Rev./ 1863— Idem, "Note on Str. filaria," ' Proc.
Zool. Soc./ 1856.— Danford (see Cobbold, Bibl. No. 52).— De
Beck, " On the Draconcule (Strong, filaria aut veinulosus ?) of
Lambs/' from 'Ann. de Logelin/ in ' Veterinarian/ vol. v,
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from ' Mem. de la Soc. Vet. du Calvados/ in ' Veterinarian/
vol. ix, p. 115, 1836. — Dupuy, "An Hydatid in the Lumbar
portion of the Spinal Marrow of a Lamb, aged eighteen
months," from ' Journ. Theorique et Prat./ in ' Veterinarian/
vol. iv, p. 285, 1831. — Engelmeyer, ' Thierarztliche Wochen-
schrift/ 1850, p. 191. — Findeisen, "Ech. in der Lunge," 'Repert.
f. Thierheilkund/ 1875, s. 48.— Fry, J., "Worms in the Trachea
(of cattle)," 'The Hippiatrist/ vol. iii, p. 5, 1830. — Furstenberg,
"On Pent, tanioides of the Sheep," 'Edin. Vet. Rev./ 1863.
— Gamgee, J., " On Parasitic Diseases," extr. from his ' Rep.
to the Privy Council/ ' Edin. Vet. Rev./ Dec, 1863. — Idem,
" On Cattle Diseases (including those produced by Entozoa)/'
from letters in the 'Times' of Oct. 22, Nov. 10 and 13, &c. ;
in 'Edin. Vet. Rev./ Dec, 1863. — Idem, "On Diseased Meat
(especially in relation to Trichina)," ' Pop. Sci. Rev./ Jan.,
1864. — Idem, " On Sturdy in Sheep (with figs, from Van
Beneden, Leuckart, and Cobbold)," ' Edin. Vet. Rev./ vol. i,
p. 440, 1859. — Idem, " Gleanings from the Researches of
Eschricht, Haubner, A. Thomson, &c, respecting the Origin
and Development of the Entozoa," ' Veterinarian/ 1855. —
Giacomini (see Bibl. No. 13). — Gulliver, "On the Structure of
the Entozoa belonging to the genus Cysticercus," 'Med.-Chir.
Trans./ ' Lond. Med. Gaz./ and 'Lancet/ 1840-41. — Idem,
" Notes on the Ova of Dist. hepaticum, and on certain Cor-
puscles obtained from the genus Cysticercus," 'Proc. Zool.
Soc./ March, 1840, and in 'Ann. Nat. Hist./ vol. vi, 1841;
also 'Month. Journ. Med. Sci./ vol. ii, 1842, and 'Micros.
Journ. and Struct. Rec./ p. 95, 1842. — Hewlett (see Bibl. No.
13). — Holmes, J., "Filaria in the Bronchi of a Calf," 'Vet.
Rec./ vol. i, p. 125, 1845. — Hunter, J., " Ileum of a Ruminant
containing Acephalocyst Hydatids /' see description of prepara-
tion No. 863 in the ' Catalogue of Mus. Roy. Coll. Surg. Lond.'
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Humerus of an Ox" (ibid., prep. No. 864). — Idem, "On

354

PARASITES OF ANIMALS

Hydatids of the Sheep," in supp. to his paper on ' Human
Hydatids/ in ' Trans, of Soc. for the Improvement, &c./ vol. i,
1793, p. 34. — Huxley, " On the Anatomy and Development of
Echinococcus (from a Zebra)/' ' Proc. Zool. Soc./ and 'Ann.
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of the Cornish graziers)/' ' Veterinarian/ vol. iv, p. 573, 1831.
— King, E., " On the Propagation of Eot (by means of the eggs
of Fasc. hepatica) in Sheep/' ' Veterinarian/ vol. ix, p. 95, 1836.
— Krabbe, " Husdyrenes Indvoldsorme," ' Tidsskrift for Vet.'
(See also my notice of the memoir in ' Lond. Med. Eec' for
1872 ; repr. in ' Veterinarian/ May, 1873.) — Kiichenmeister (see
Bibl. No. 13). — Leaver, T., " Cases of the Husk (from worms)
in Cattle," ' Veterinarian/ vol. ii, p. 355, 1829. — Lepper,
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Bibl. No. 13). — Lord, J., " On some of the Parasites principally
affecting Ruminants," 'Trans. Vet. Med. Assoc./ 1842—43. —
Masse (see Bibl. No. 13). — Mayer, T., "On Hoose in Cattle
(from Filaria)," 'Veterinarian/ vol. xiii, p. 227, 1840.— M'Call,
J., "On Sturdy in Sheep," ibid., vol. xxx, p. 267, 1857.—
Me'gnin, P., " Le Str. minutissimus," ' Bullet, de la Soc. Cen-
trale Vet./ in 'Eec. de Med. Vet./ July, 1878, and in 'Ann.
de Med. Vet./ Oct., 1878, p. b63.—Moorcroft, "Brain Hydatids,"
f Med. Facts and Observ./ 1792.— Morton, W. J. T., " On the
Entozoa affecting Domesticated Animals, and particularly on
Fasc. hepatica or Liver Fluke in Sheep," ' Veterinarian/ vol. xii,
p. 735, 1839.— Mosler (see Bibl. No. 13).— Murie, J., "On the
occurrence of CEstrus tarandi in a Eeindeer in the Zoological
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a Leech (Trocheta) found in the Viscera of a Molluscan Deer
(Cervus nioluccensis, Miiller)," ibid., 1865. — Numan, "Over den
Veelkop-blaasworm der Hersenen " (this beautifully illustrated
memoir, in the ' Trans, of the Dutch Soc. of Sciences/ supplies
an elaborate bibliography of continental writings on Ocenwrus
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p. 225 et seq.— Oliver (see Bibl. No. 13).— Padley, G., 'On
Entozoa from a Sheep / see Sandie. — Parsons, " On Diarrhoea
in Lambs (with bronchial worms)," 'Veterinarian/ 1855, p. 685.
—Patellani, "Sturdy in Cattle," from ' Munchen Jahresbe-
richt/ in ' Veterinarian/ vol. xxx, p. 81, 1857.— Pellizzari (see
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Bibl. No. 13).— Ralph, T. 8., " On the Parasitic Nature of

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355

Pleuropneumonia," &c, two papers in ' Austr. Med. Journ./
1865.— Ranle," Pulmonary Entozoic Disease of Sheep/' 'Path.
Soc. Trans./ 1858; see also 'Veterinarian/ vol. xxx, p. 708,
1857.— .Raynaud, " A Word on the Cachexia, or Rot in Rumi-
nants/'' trans, from the ' Journ. des Vet./ by W. Ernes, in
'Veterinarian/ vol. xxxiii, p. 488, 1859.— .Bead, R., "Destruc-
tion of Strongylus and Filaria in the Bronchial Passages of
Calves, through nasal inhalation of ether, chloroform, oil of
turpentine, or rectified oil of amber," t Veterinarian/ vol. xxi,
p. 604, 1848.— Reck (see De Reck).— Reed, R., "Congenital
Hydatids in a Lamb," 'Veterinarian/ vol. viii, p. 551, 1835. —
Rhind, " Description of a species of Worm (Pentastoma) found
in the Frontal Sinus of a Sheep," 'Farrier and Naturalist/
vol. iii, p. 277, 1830, and ' Lancet/ 1829.— Robertson, " Remarks
on Taenia in Lambs," ' Rep. of Scottish Med. Vet. Soc./ in
'Veterinarian/ 1875, p. 80. — Rochard (see Bibl. No. 13). — Rose,
C. B., " On Ccenurus and Acephalocysts," ' Lond. Med. Gaz./
vol. xxiv, p. 525, 1844. — Idem, " On the Anat. and Physiol, of
the Cysticercus tenuicollis," ' Roy. Med.-Chir. Soc. Trans.,' and
'Lancet/ 1848. — Idem, "On the Vesicular Entozoa, and par-
ticularly Hydatids," 'Lond. Med. Gaz./ vol. xiii, p. 204, 1833-
34. — San die (with Padley), " On Entozoa in the Lungs of a
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1875, s. 295. — Shenton, "Worms from the Stomach of a Cow,"
'Veterinarian/ 1844, p. 487. — Siedamagrotzhy , "Hydatids in
the Liver of a Cow," ' Bericht iib. das Veterinaweseu im Kon.
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Worms in the Stomach (abomasum), being remarks on Mr,
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Idem, ' The Rot in Sheep, its nature, cause, treatment, and
prevention/ London, 1862. — Idem, "On Filarige in the Bronchi
of Calves," 'Trans. Vet. Med. Assoc./ 1843, p. 517.— Idem,
" On Strongylus in the Bladder and Intestines," ibid., 1843. —
Idem, " On Hydatids of the Liver of a Sheep (Mr Scruby's
case)," ibid., p. 331. — Idem, " On Disease of the Mesenteric
Artery from Strongyli within the Vessel," ' Path. Soc. Trans./
1854. — Idem, "Lecture on the Nature and Causes of the Disease
known as Rot in Sheep," vol. xxxiv, p. 274, 1861. — Speuce, G.
W., " On GEstrus of the Ox," ' Edin. Med. Journ./ 1858, and
'Edin. Vet. Rev./ vol. i, p. 400.— fitt Gyr (see Bibl. No. 13).—
Stoddart, J., " Case of Hydatids in the Liver (of a Cow),"

350

PARASITES OF ANIMALS

'Veterinarian/ vol. xi, p. 637, 1838. — Sutton (Lecture),
f Gardiner's Chronicle/ June 29, 1872.— Sylvester, F. B., "Cases
of Parasites infesting the Brains and Intestines of Lambs/'
' Vet.Eec./ vol. ii, p. 40, 1846. — Thudichum (see Bibl. No. 13).
— Idem, " Echinococci from the Sheep's Lungs," ' Rep. Med.
Soc. Lond./ in ' Assoc. Med. Journ./ 1856, p. 195. — Tommasi
(see Bilb. No. 13). — Van Beneden, " On the Development of
Coenurus," from ' Comp. Rend./ in 'Ann. Nat. Hist./ vol. xiv,
1854. — Watson, K. W., " Experiment for the Cure of the gidd
(Coenurus) in a Sheep/' 'Lond. Med. Repos./ 1815. — Willemoes-
Sithm,va. ' Sieb. and Koll. Zeitschr./ Bd. xxv, s. 176. — Wilson,
JE., " On the Anatomy of Trichocephalus affinis," ' Vet. Rec./
1846. — Wymann, "Note on Filaria in the Bronchi of a Sheep,"
see Anon, on " Entozoa and Parasites," in f Amer. Journ.
Sci./ vol. xxxix, p. 183, 1840. — Youatt, " On Hydatids in the
Sheep (symptoms, prevention, treatment, &c.)," ' Veterinarian/
vol. ii, p. 519, 1834. — Idem, " On the peculiar Bronchitis in
young Cattle, accompanied by Worms in the Bronchial Passages,"
part of lecture, in e Lancet/ 1832. — Idem, "Hydatids in the
Brain of a St Domingo Goat," 'Veterinarian/ vol. ix, p. 443,
1836. — Idem, " Bronchitis from Worms (in Oattle)," ibid.,
vol. vi, p. 177, 1833.— Idem, "Hydatids in the Brain (in Cattle),"
ibid., vol. vii. — Tvart, " Brain Hydatids," ibid., 1828, p. 19. —
Zahn, " Lungenwiirmer beim Reh," ' CEsterr. Vieteljahrschr. f.
w. Vet./ 1875, s. 125.— Zum (see Bibl. No. 13).— For further
references to the literature of Hydatids in Animals, see Biblio-
graphy No 20, o, in the first half of: this work.

Paet IX (Solidungula).

It will naturally be expected that I should give a full
account of the parasites of the solipedal, solidungulate, or
equine mammals. As regards the horse I regret that I cannot
meet this expectation in so complete a manner as the subject
deserves ; nevertheless, with the aid of an extended biblio-
graphy the summary here offered will be found to be tolerably
exhaustive. At all events I think I may say that no similar
record has hitherto been attempted.

The liver fluke {Fasciola hepatica), though not very frequent
in the horse, is not uncommon in the ass. In dissecting-room
subjects at the Royal Veterinary College it is often encountered.

SOLIDDNGULA

357

In France it was originally found in the horse by Daubenton.
As I learn from Sonsino, Dr. Abbate Bey recently recorded a
similar find at Cairo. In solipeds generally the liver fluke
appears to be almost harmless, for, notwithstanding the fright-
ful ravages produced by rot amongst a variety of animals
besides sheep, we have no evidence of the destruction of horses
from this cause. In the German outbreak of 1663-65 multi-
tudes of cattle and deer perished, and in the French outbreak
of 1829-30 five thousand horned beasts succumbed in the
arrondissement of Montmedy alone. In neither of these
epizootics were the solipeds affected. More importance attaches
itself to the study of the amphistomatoid flukes. These
parasites, though in a scientific sense only recently discovered
in equine bearers, have been long known to the natives of
India. They appear to be capable of producing serious intes-
tinal irritation. I have described two forms (Amphistoma
Gollinsii and A. Coll., var. Stanleyi), which infest the colon.
The specimens sent to Prof. Simonds from India by Mr Stanley,
Y.S., were much larger than those sent to me from Simla by
Mr Collins, V.S., some ten years later (1875). As in all other
amphistomes obtained from the intestines of elephants and
cattle, the worms, when fresh, were of a bright brick-red color.
By the natives of India these parasites are called Masuri;
but no description of the worms had been published prior to
the account which I gave of the contributions forwarded by
Major-General Hawkes, Mr Collins, and Mr Stanley.

I shall have occasion to speak of the elephant's Masuri
further on ; but in the meantime I must remark that the
generally received notion as to the parasitic cause of the earth-
eating propensities of various animals seems to have some
foundation in fact. Not alone from Major-General Hawkes in
Madras, from Mr Folkard in Ceylon, and from various other
trustworthy sources, have I been informed of this habit on the
part of Indian horses, but Dr Rowe told me that Australian
horses, and even sheep, infested with stomach-worms, are in the
constant habit of consuming large quantities of sand. From
all the facts that have come before me, I am inclined to think
that gastric or intestinal irritation, however brought about
may induce the habit in question, parasites being only one of
the many sources of irritation giving rise to symptoms of colic
in solipeds and pachyderms alike. At all events the African
elephants at the London Zoological Society's Managerie, as

358

PARASITES OF ANIMALS

repeatedly witnessed by myself, are in the habit of swallowing
large quantities of soft mud during the summer months, but
no traces of masuri have as yet been detected in their
fasces.

When by letter I informed Major- General Hawkes of an inte-
resting find by Mr Collins of about a thousand Amphistomes in
the colon of a horse that had died at Simla, the announcement
called forth a reply which is sufficiently instructive to be quoted.
Writing from Secunderabad in July, 1875, he says, respecting
this " find : " — " Your statement has incidentally thrown light
upon a subject which has puzzled many of us in this country.
It occasionally happens that a horse, on being opened after
death, is found to have accumulated in his intestines large
quantities of sand and gravel. In a recent case this accumula-
tion amounted to 14^ lbs. Until recently it was always held
that this gravel or sand could only be introduced with the
animal's food. All grain in this country is trodden out by
bullocks on an earthen floor, and the grain undoubtedly contains
a proportion of sand and gravel derived from this source.
Although this ought to be carefully washed out before it is given
to the horse, still, owing to the carelessness of the native horse-
keepers, this cleaning is, I expect, often omitted. In the daily
' feed ' of eight or ten pounds of grain given to each horse the
utmost quantity of sand or gravel that could be found admixed
therewith would not probably exceed two or three ounces ;
consequently it would take from 77 to 116 days to accumulate
so large a quantity as 14| lbs. Now, the advocates of the theory
of the gradual accumulation of sand in this way have never been
able to explain why the grain, grass, hay, and other ingesta
should pass in the ordinary way through the intestines, whilst
this sand or gravel remains behind. One can understand the
possibility of such substances as wool, hair, or similar matters
concreting in the alimentary canal, though I believe they are
usually found in the stomach, and not in the intestines ; but
how a most incohesive substance like sand can possibly accumu-
late in the gradual way required by their theory I have never
heard even plausibly explained. On the other hand, the fact
that horses are often excessively addicted to eating earth is
well known; and if my memory serves me correctly, it was
found necessary, about twenty years ago, to remove the mud-
walls of the pickets surrounding some of the horses of a
mounted corps in this presidency in consequence of this habit.

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359

Now, given the fact that the amphistoma has been found in the
horse (as your specimens prove), may we not fairly suppose it
possible that the animal resorts to the same mode of ridding
himself of this parasite as does the elephant ? and also, would
it not in a much more natural manner account for the large
quantity of gravel or sand found in the intestines than does the
theory of gradual accumulation ? Reasoning from analogy, as
in the case of the elephant, this eating of earth in the horse
would be an instinctive effort on the part of the " host " to rid
himself of the parasite. This self -taken remedy is doubtless in
many cases quite effectual, though unnoticed. The fatal cases
are probably those in which the horse has either overdone the
remedy or where the
system was too de-
bilitated to carry off a
quantity of sand or
gravel that would
otherwise have safely
passed through the
intestines of a horse
in more robust health.
The actual fact must,
of course, be verified
by careful investiga-
tion/'

Closely allied to
the Masuri is an
amphistome which I
originally named Gas-
trodiscus Sonsinoii,
but which should be
altered as opposite
(Fig. 62). It exceeds

in length and in
breadth (16 mm. long
by 10 broad). Its dis-
covery by Dr Sonsino
was one of the results
of his examination of
sixteen carcases of soli-
peds that died during
the Egyptian plague of 1876. Specimens having been forwarded

-tr.

Fio. 02 —Gastroiliscus Sonsinonis. a, Month ; i, caudal sucker
and posterior mesial cleft; c, left lip of the gnstric disk; </,
anterior mesial cleft; e, e, gastric suekerlets ; /, reproductive'
papilla. Enlarged. Original.

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PARASITES OF ANIMALS

to Panceri, Von Siebold, Leuckart, and myself, most of us at
once agreed that the worm was new to science. Pointing to
the genera, Notocotylus and Aspidocotylus, I explained its close
affinity to the latter more particularly. Whilst Notocotylus has
fifty supplementary suckers on its back, Aspidocotylus has nearly
two hundred small ventral suckers seated on a convex disk.
In Gastrodiscus a still larger number of suckerlets are placed
in the deep concavity of a large gastric disk formed by the
outstretched and inrolled margins of the body of the parasite.
Zoologically speaking, the odd thing about this singular worm
lies in the circumstance that its nearest fluke-relation, so to speak
(Aspidocotylus mutdblis), dwells in a spiny-finned fish (Cata-
phractus) ; and this fish itself forms an aberrant genus of the
family to which it belongs (Triglidae) . From what has been
said it will be seen that our Gastrodiscus must not be confounded
with Gotylegaster cochleariform (or with its synonym Aspidogaster
cochleariformis) , to which parasite Von Siebold was, I believe,
induced to refer it. Like most of the true amphistomes, the
worm in question infests the intestines. Although discovered by
Sonsino at Zagazig in plague-affected corpses, there is no reason
to suppose that this helminth was in any way etiologically
connected with the Egyptian epizooty.

The tapeworms of the horse are of great interest practically.
Excluding Sander's Tcenia zebra, which was doubtless T. plicata,
at least five species have been described, but they may probably
be all reduced to two distinct forms and their varieties. Whilst
Tania plicata acquires a length of three feet, the strobile of
T. perfoliata never exceeds five inches. The lobes at the base of
the head in the latter are distinctive. The former is usually
confined to the small intestine, but the perfoliate worm often
occupies the cascum and colon in great numbers. As regards
T. mamillana, I may say that neither Gurlt's descriptions nor his
figures are convincing. The worm is, I believe, identical with
T. perfoliata. In like manner, after going into the matter
with some care, I am accustomed to speak of Megnin's T. inerme
as T. perfoliata, var. Megnini, and of JBaillet's T. innome as T.
perfoliata, var. Bailletii. I have examined great numbers of
equine tapeworms, but whether my determinations on this point
are correct or not, the case recorded by Megnin is of remark-
able interest. Clinically, indeed, it is not entirely unique, since
a somewhat similar case has been recorded by Mr Poulton.
In Megnin's equine patient the autopsy revealed the presence

SOLIDUNGULA

:;c,i

of 200 bots, 153 lumbricoids, upwards of 400 oxyurides,
and several thousand palisade worms, besides numerous tape-
worms. In Mr Poulton's patient large quantities of tape-
worms were found in the duodenum (and in large sacs of the
walls of other sections of the small intestine), and also myriads
of the little four-spined strongyle, in addition to about a score
of palisade worms. Both Megnin' s and Poulton's patients died
suddenly j but the great interest attaching to Megnin' s case
arises from the boldness of manner in which the French savant
interprets the phenomena of the intestinal sacculation in relation
to the development of the tapeworms. M. Megnin assumes
that the sacs are due to the formation of polycephalous or
coenuroid scolices. Without contradicting Megnin' s ingenious
interpretation of the phenomena in question, I may say that the
difficulty I have in accepting his view arises from the circum-
stance of the rarity of the occurrence of these sacs. In Poul-
ton's case of Taenia perfoliate the sacs were present, and they
were productive of similar results ; but in the scores of other
recorded cases of sudden death from the same species of tape-
worm (as published by Mr Eees Lloyd, and myself), the pre-
sence of such sac-formations is not once mentioned. To be
sure, their presence may have been overlooked, but this is
scarcely likely, seeing the great care taken by Mr Lloyd in
conducting the autopsies. I cannot dwell upon the subject at
greater length. The presence of so many sexually-immature
strobiles, combined with the existence of the intestinal wall
sacs, certainly does seem to point to the existence of coenuroid
bladder- worms, but until the existence of the polycephalous
scolex be actually demonstrated one must be cautious in con-
cluding " that the horse nourishes at the same time the strobila
and scolex of the unarmed tapeworm." Practically, we now
know for certain that not only are tapeworms capable of pro-
ducing a fatal issue in isolated cases, such as those recorded
by Megnin and Poulton, but that they may also be productive
of disastrous epizooty, as proved by Mr Lloyd in the case of
Welsh mountain ponies.

In this connection I may perhaps be pardoned for saying
that this discovery in 1875 was one of the practical results
directly issuing from the publication of my ' Manual' in
1874. The attention of the veterinary profession having
been called to the subject of parasitic epizooty, Mr Lloyd
was the first to make search for helminths amongst some

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PARASITES OF ANIMALS

few of the carcases of the hundred and more equine animals that
perished in South Wales. Two totally distinct epizootics pre-
vailed. In the Beacons district tapeworms alone were the
cause of death, whilst in the Deangunid district scores of animals
perished from strongyles. In another district a hundred
animals perished from tapeworms. These parasites I identified
as examples of Strongylus tetracanthus and Tania perfoliata.
Taking all the helminthological facts together we have made a
great advance both in hippopathology and equine epidemio-
logy; and, as I observed at the time, the scepticism which not
unnaturally still exists (in reference to entozoa as a frequent
cause of death amongst animals, both wild and domesticated)
will sooner or later be dispersed by that wider attention to the
subject which our labors have invoked.

In relation to equine disease the facts brought forward
are too important to be dismissed in a single paragraph.
As two distinct kinds of parasitic epizooty were discovered,
the circumstances connected with their separate detection must
be noticed at greater length. Further on, I shall again deal
with the helminthiasis due to strongyles. It was on the 17th
of April, 1874, that I received from Mr Lloyd, of Dowlais,
Glamorganshire, a communication calling my attention to a
fatal epizootic affecting ponies. He supposed the outbreak to
be due to parasites. On the following day I also received a
parcel containing portions of the lower intestines, which had
been removed from one of the diseased animals. The victim in
question, a pony mare, had died on or about the 12th of April,
at Llangunider, Breconshire. Mr Lloyd states in his letter
that he " presumes " that the pony's death was caused " by the
presence of small worms," examples of which he now for-
warded for the purposes of identification and investigation. He
also sent some equine tapeworms. Mr Lloyd had already
inferred that his small worms were " strongyles and in regard
to the tapeworms he says : — " This species of parasite has caused,
or is supposed to have caused, the death of at least one hundred
mountain ponies." The investigation being immediately pro-
ceeded with, I may so far anticipate my record of the results
obtained as to state at once that the facts observed by me con-
firmed Mr Lloyd's suspicions — proving, beyond a doubt, that
the pony above mentioned had succumbed to injuries inflicted
by myriads of minute strongyles. Not only did I find the
fascal matter of the colon loaded with mature strongyles, but

SOLIDUNGULA

003

the walls of the intestine were also occupied with encysted and
immature forms of the same nematode species. To such an
extent had infection taken place, that I was enabled to count
no less than thirty-nine strongyles within the space of the one
fourth of a square inch. All parts of the sections of the colon
under examination were almost equally invaded ; so that, taking
the average, I am clearly within the mark in saying that every
square inch of the gut yielded at least one hundred parasites.
The walls of the entire colon must therefore have been occupied
by tens of thousands of these creatures, to say nothing of the
scarcely less numerous examples lying free or lodged within the
faecal contents of the bowel.

In a second and more extended communication, sent in reply
to inquiries as to the cestodes, Mr Lloyd (whose letter I have
abridged) writes : — " I regret that I cannot give you very full
particulars respecting the tapeworms. During the last twelve
months mountain ponies grazing on the lower districts of
Breconshire, which comprise some of the highest mountains in
South Wales, have been dying in great numbers, from what the
farmers indefinitely term inflammation. From what I have seen
and heard, it appears that there are three causes of death, the
tapeworm, the small worms (which I presumed were a kind of
strongyle), and catarrhal disorders, such as have been common
among horses of late. By far the greater number of deaths
(from what I can glean) have been caused by the parasites. In
the Tstradfellte or Penderin districts there has been no investi-
gation, although the disease has reigned there for a longer
period, about eighteen months, with (from what I have heard)
a larger number of deaths than elsewhere ; so I shall let these
remain for the present, as I have not had an opportunity to see
or hear anything authentic about them. In the Talybont
district the cause appears to be the small worms (like those I
sent). The owner of the animals said that a month ago, when
he went to look after his ponies, they were appearing quite well,
and looked as well as he could expect them at this season,
but he was astonished to find some of them a fortnight ago
looking very lean and wasting, and he thought that the weather
was the cause of it, yet resolved to see them of tener ; the next
time he saw them one was dead, and knowing of the loss in the
neighbourhood, and fearing he would be a sufferer, he sought
aid, applying to me. When I arrived two days following two
more were dead, and they presented an emaciated appearance.

364

PARASITES OF ANIMALS'

The post-mortem examination revealed a healthy condition of
the whole of the intestines, save slight thickening of some parts
of the colon and rectum, which contained, enclosed in the
mucous membrane, in cysts or minute sacs, worms coiled upon
themselves. Each cyst, containing one worm, was best seen
by transmitted light. The colon was nearly full of faecal
matter, which contained thousands of parasites scarcely visible.
The largest were very few in number, not exceeding an
inch in length and barely one sixteenth in diameter at the
middle portion. They somewhat tapered at both ends. The
caecum was half full of fluid fasces, containing no visible worms ;
the rectum, with fasces of natural consistence, the examination
of which reavealed only two or three evident worms ; so that
the examination of fasces of living animals giving results like
this would not assist the diagnosis, unless suspected. Small
intestines — these latter contained about a dozen bots, which
were nearly free, but had pierced to the muscular coat. I
should think they had participated in the disease. The right
lung had been slightly congested ; nothing else abnormal to be
seen. Possibly congestion of lungs would arise from the
distress when pained with worms, for the pony was found on its
back with its head in a thicket — it had not appeared to have
struggled — with its teeth firmly closed.

" Respecting the animals affected with Tasnia, it is remark-
able that, as a rule, they are in fair condition. The average
time they appear to be troubled with the worms is two months,
and the symptoms observed have been many. They are at
first seen to be unable to keep up with the other ponies, ex-
tending the head and turning the upper lip up, rubbing the
quarters, staring coat, suddenly appearing distracted, seizing
turf in mouthfuls when being griped or pained, others running
away as fast as they can go, or rolling and kicking on the
ground for five or ten minutes, then walking away as if nothing
had happened, if coming down a slope quickly almost sure to
fall headlong, easier caught, not unfrequently coughing, groaning
noise, appetite good, and, what is peculiar in some of them, lame-
ness of one of the hind limbs, mostly the near hind leg, with
slight knuckling over at fetlock.

" Post-mortem appearance. — Abdominal viscera normal, save
rectum, which is in some places slightly congested ; colon
nearly full of fasces, no worms ; cascum, in which worms are
alone found, is nearly full of fascal matter of thicker consistence

SOLI DUNG ULA

365

than usual, and nearly half made up of worms j stomach half
full of partly digested food j heart and lungs healthy ; Schnei-
derian membrane injected; mucous membrane of trachea and
part of larger bronchiae of a more or less livid colour (which
may be owing to asphyxia) j corner of tongue bitten off ; mouth
very close.

" Several animals were found at times lying dead together.

" Of the Tcenia as many as three or four ponies, which some
hours previous had been seen grazing unaffected, were found
dead on the same spot ; and this to my own knowledge, one
farmer having lost ten.

" Of the small worms I have been told by a farmer that in
his district one of his neighbours had lost twelve ponies."

As I had partly misunderstood my informant's original
statement, Mr Lloyd, in a third communication, repeated the
evidence, emphatically reminding me that " the ponies affected
with tapeworms are in a district six or seven miles distant from
those affected with strongyles. Those troubled with tapeworms
are in good condition, as a rule, up to death ; they are noticed
to be troubled generally for two months previous to death,
and may be seen at one hour grazing and apparently well, and
dead or dying the next hour. As many as four have been
found dead at the same spot. In this (the Beacons) district
the tapeworms alone have been found and not a single strongyle.
In the Deangunid district strongyles only have been found, such
as I sent you. The ponies have been noticed ailing for three
or four weeks, becoming rapidly emaciated and dying from
exhaustion. In tapeworm-affected animals the caecum is nearly
half full of these parasites. The animals thus affected are
on the red sandstone formation, whilst those affected with
strongyles occur on the limestone formation — the latter affording
the drier situation."

Being on the teaching staff of the Royal Veterinary College
I was particularly glad to have the authority of an experienced
veterinary practitioner to testify to the injuriousness of Tcenia
perfoliata in the horse. Over and over again I had pointed
out to the members of my class the desirability of examining
the faeces of solipeds where obscure symptoms of intestinal irrita-
tion existed. Not only so ; at the request of friends I wrote out
prescriptions suitable for equine patients suffering from tapeworm.
I felt the more indebted to Mr Lloyd, inasmuch as his practical
views served to strengthen the propositions I had advanced in

3G6

PARASITES OF ANIMALS

connection with internal parasites as a frequent cause of epizo-
otics. My views were criticised at the time with a vigour and
warmth well worthy of those who are afraid of advancing
epidemiological science too rapidly ; but it seems that so far
from my having overstepped the bounds of moderation in this
matter I had, in reality, been too cautious. Certainly it can
now no longer be said that " the symptoms created by tape-
worms in the horse are of little or no consequence." Here,
therefore, I repeat, we have made a clear and rapid advance in
our knowledge of helminthic disease ; and from the impulse
thus given to hippopathology it is only reasonable to look for
still further advances in veterinary medicine. By-and-by, the
scepticism which not unnaturally exists in reference to entozoa
as a frequent cause of death amongst animals, will be dispersed
by even yet clearer enunciations regarding the important part
these parasites play in the destruction of our most valuable
creatures.

Apart from the question incidentally raised by Megnin as to
their origin and mode of development, the presence of larval
cestodes in horses cannot be passed over. The common hydatid
(Echinococcus veterinorum) , though not of frequent occurrence,
is occasionally productive of fatal consequences. Very inter-
esting cases are recorded by Messrs Henderson and Kirkman,
aided by the valuable comments of Professor Varnell. Mr
Hutchinson observed an hydatid in a horse's eye, and Mr Yin-
cent noticed lameness, as resulting from hydatids. But one of
the most interesting cases of hydatids in solipeds is that
described by Professor Huxley, from a zebra that died at the
Zoological Gardens in 1852. As stated in Huxley's elaborate
memoir (freely quoted in the first part of this work), the liver
was found to be " one mass of cysts, varying in size from a
child's head downwards." The zebra's death was purely acci-
dental, as it broke its neck while at play in the paddock. The
long bladder-worm of the horse (Cysticercus fistularis) is entirely
unknown to me, and, as before suggested, may be a mere
variety of the C. tenuicollis of ruminants. An authentic in-
stance of the occurrence of the gid hydatid (Ooenurus cerebralis)
in the horse is recorded by Gurlt. Lastly, in relation to the
question of food, it is worthy of remark that whilst beef, veal,
pork, and even mutton, are apt to be measled, the muscle -flesh
of horses is not liable to be infested by Cysticerci. This is a
fact in favor of hippophagy.

SOLID UNGULA

367

The nematodes of solipeds are very numerous, and first in
importance must be placed the palisade worm (Strongylus
(t r mat us). This worm was known to Ruysch (1721). The old
naturalists recognised two varieties {major and minor). These
we now know to be merely the final stages of growth of one
and the same entozoon ; and in both stages the worm inflicts
severe injury upon the bearer, chiefly, however, whilst wander-
ing through the tissues. The palisade worm has acquired no-
toriety principally on'account of its causing verminous aneurism,
nevertheless, this pathological change is not, in itself, the most
disastrous evil produced by the worm. In the adult state the
female reaches a length of two inches, whilst the male rarely
exceeds an inch and a half. The posterior ray of the caudal
membrane or hood of the male is three- cleft. In both sexes
the head is armed with numerous, closely- set, upright denticles,
presenting the appearance of the teeth of a circular saw or
trephine. The eggs are elliptical and somewhat constricted at
the centre, their contents forming embryos after expulsion from
both parent-worm and host. The larvee are rhabditiform,
changing their skin, in moist earth, in about three weeks, at
which time they part with their long tails. According to
Leuckart, they pass into the body of an intermediate bearer
before entering the stomach of the definitive or equine host.
From the alimentary canal they pass to the blood-vessels,
causing aneurism, and thence they seek to regain the intestinal
canal, where they arrive at sexual maturity. It is during
their migratory efforts that they give rise to dangerous sym-
ptoms in the bearer, not unfrequently causing the death of
young animals, especially yearlings. In the adult state the
worm is also dangerous to the bearer, as it produces severe
wounds by anchoring to the mucous membrane of the gut.

The proofs we possess as to the frequency of abdominal, espe-
cially mesenteric, aneurism from this source are overwhelming.
Prof. Bruchmiiller estimated the percentage of aneurismal
horses, six years old and upwards, at 91 per cent., and it is a
matter of common observation in veterinary dissecting rooms
that verminous aneurism is rarely or never absent in the ass.
Professors Dick, Simonds, Pritchard, Williams, and many other
English and Scotch veterinarians of eminence, have all borne
testimony of this kind, and, for myself, I may say that one of
the earliest pathological appearances with which I became
familiar, some thirty years back, was that presented by niesen-

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PARASITES OF ANIMALS

teric arterial disease of the ass. In relation to fatal colics in
the horse the study of verminous aneurism is of the highest
moment. On this subject Prof. Friedberger has published some
valuable lectures, in which, amongst other points, he incidentally
remarks upon the comparative freedom of military horses from
aneurism as compared with ordinary laboring horses. This
arises partly from the fact that the latter are not cared for to
the same extent, dietetically and otherwise ; and, moreover,
cavalry horses are, as a rule, younger than ordinary working
animals. Whilst Friedberger, in his suggestive brochure, does
ample justice to the writings of his colleague, Dr Bollinger, it
may be said, in like manner, that he does not fail to recognise
his great indebtedness to the researches of Leuckart. So
practically important, however, do I deem Bollinger's summary
of the whole subject in relation to the hippopathological aspects
of parasitism, that I feel it desirable to record his conclusions
at full length. No professional man having any pretensions to
a knowledge of the veterinary art — or, for that matter, to
parasitism in relation to sanitation — should remain uninformed
on this subject. Dr Bollinger's results are thus stated :

1. The worm aneurism of the visceral arteries of the horse,
existing in 90 to 94 per cent, of adult horses, has a general
correspondence with the aneurisma verum mixtum of man. It
is, however, distinguishable from the same by its seat, cause,
character of its walls, contents, and mode of termination. The
worm-aneurism arises from a parasitism of the palisade worm
(Strongylus armatus), owing to an inflammatory affection of the
arterial walls which it causes, and which one may describe as
a recurrent traumatic endo-arteritis. This holds good for all
the visceral arteries, with the exception of the abdominal aorta,
in which an aneurism may arise from local increase of pressure.

2. The formation and further development of the aneurism is
also favored by the narrowing of the arterial calibre, which
is caused by the inflammatory swelling of its walls, and also by
the contemporaneous formation of a thrombus (clot), this latter
still further supporting and exciting the inflammation of the
inner coat.

3. Whilst the causes above mentioned (and of these more
particularly the continued presence of the palisade worms and
the plugging of the smaller arteries by thrombi) favor the
growth of the worm-aneurism, the small size of the same, not-
withstanding the years it has existed, is explained by the con-

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369

siderable hypertrophy of the muscular layer, by the tough fibrous
capsule formed in many cases by the connective tissue of the
mesentery, and by the adhesion of the intestines to the perpen-
dicular and free-lying anterior mesenteric artery ; in particular
this last-named circumstance does not allow of any very consider-
able shortening of the mesentei'ic artery, which would necessarily
be accompanied by considerable dilatation of the arterial tube.

4. The favorite seat of the worm-aneurism is the trunk of the
anterior mesenteric artery, directly at its origin from the abdo-
minal aorta. Most frequently that part of the arterial trunk is
dilated from which the arteria ilea, csecales, and colica inferior
(arteria ileo-cceco-colica) arise, less frequently the arteria colica
superior at its origin, and the arteries of the caecum and colon
in their course in the meso-csecum and meso-colon. The
verminous aneurism also occurs in the cceliac artery (Bauch-
schlagader), in the posterior mesenteric artery (Gekros-arterie),
in the renal artery, and in the abdominal aorta. A horse is not
unfrequently afflicted with several aneurisms of this kind at one
and the same time. Thus in one case (described by Bollinger)
there were six of these aneurisms affecting the abdominal aorta
and its branches in the same horse. The verminous aneurism
may occur from the sixth month of life onwards, and with
increasing age ; the number of horses free from such aneurisms
becomes continually smaller.

5. The size of the aneurism varies between that of a pea and
that of a man's head. The dilatation is, as a rule, equal on all
sides, the form being usually thumb-shaped or bottle-shaped,
passing into that of a cone or long oval figure. This general
configuration is principally due to the free and moveable situa-
tion of the anterior mesenteric artery.

6. In contrast to aneurisms in man, the walls of the worm-
aneurism of the horse are almost without exception indurated.
In addition to the mesenteric connective tissue, all the arterial
coats, and especially the tunica media, generally take part in this
induration. The hypertrophy of the media, which stands
unique in respect of what is known of arterial disease, forms a
compensatory action of the arterial wall, analogous to the
muscular hypertrophy of the heart in valvular disease. This
change in the media points to the fact that in the development
of aneurism in man the early disturbance of the nutritive pro-
cess in the tunica media is not a less essential factor than the
degeneration of the tunica intima. '

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PARASITES OF ANIMALS

The changes in the intima are the least constant. They
present all stages of progressive and retrogressive metamor-
phosis, from simple induration to ulceration and calcification.
In the walls of the verminous aneurism one not unfrequently
finds all the pathological changes exhibited by atheroma in man.
Calcification is a common form of the retrograde process, and,
in very rare cases, may pass on to the formation of true bone.

7. In addition to the palisade worms, one almost constantly
finds a parietal thrombus contained in the aneurism. It covers
the inner wall either partially or completely, being in the latter
case perforated for arterial offshoots. This clot may occlude
the artery, and it is not unfrequently continued into the arterial
branches (peripherally) or into the aorta (centrally). Amongst
the various changes that the clot undergoes, organisation of its
outermost layer and softening are the most frequent. The
constant occurrence of this clot is due to the presence of the
worms, to the inflammation, ulcerative and regressive affection
of the intima, and to the dilatation of the arterial tube.

8. The palisade worms are seldom absent from aneurisms of
the horse. Their not being present is merely an accidental
circumstance. On the average, nine palisade worms go to a
verminous aneurism, and eleven in the horse. The highest
number of worms found in one horse reached 121. Not unfre-
quently, also, palisade worms, or their coverings in the form of
larval skins, are found in the aneurismal walls. The immigra-
tion and emigration of the palisade worms out of the intestine
into the aneurism, and the reverse, take place probably, as a
rule, within the arterial circulation. The path of the worm
does not appear to be always the same, inasmuch as they
can also wander through the peritoneal cavity. The worms
found in the aneurismal walls are probably mostly only strayed
specimens.

9. From a comparative pathologico-anatomical point of view,
the developmental history of the aneurysma verminosum proves
that a circumscribed endo-arteritis can determine the formation
of an aneurism.

10. Like the worm-aneurism itself, atheroma of the abdo-
minal arteries arises from a circumscribed acute and subacute
endo-arteritis. The histological changes in the secondary
atheroma of horses are perfectly analogous to those of the
spontaneous atheroma of man. Idiopathic atheroma, as seen
in man, does not occur any more in the horse than in the other

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371

domestic animals. Atheroma in the horse is always secondary.
To be sure, one observes an idiopathic chronic endo-arteritis in
many abdominal arteries of the horse, which, however, never
exhibits indications of atheromatous degeneration.

11. In consequence of its position the worm-aneurism of
horses is not open to physical examination, and on that account
cannot be diagnosed by physical signs ; moreover, it offers no
characteristic symptoms. Its termination by rupture is
extremely rare, the aneurisms of the abdominal aorta being
more disposed to rupture than those of the anterior mesenteric
artery. Of eighteen cases of known perforation, fifteen opened
into the peritoneal cavity, and three into the bowel. The
dangerous symptoms of the worm-aneurism are exclusively due
to embolism and thrombosis of the affected artery, arising from
the parietal clot. The latter becomes especially dangerous
through its increasing size and the softening which often
accompanies it. The absorption and shrinking of this parietal
clot, be it organised or not, is materially assisted by the high
pressure to which it is exposed.

12. The very marked symptoms of vascular obstruction — the
sero-haemorrhagic intestinal infarct — in embolism and thrombosis
of the mesentric arteries are easily explained by paralysis of
the muscular coat of the intestine, by the absence or paucity of
valves in the portal vein, by the readiness with which meteor-
ismus (or flatus) arises, especially in herbivora, and by the loose
consistence of the intestinal walls or villi.

13. The occlusion of the intestinal arteries, especially that
arising suddenly, always has for its result a partial or complete
paralysis of the portion of bowel which they supply. The
palsy of the intestine causes the forward movement of the
intestinal contents to cease, a stoppage of the faeces, a hin-
drance to the discharge of faeces and gas, and also that exceed-
ingly dangerous formation of gas (within the intestinal tract)
which in the herbivora is so abnormal, both quantitatively and
qualitatively.

14. In embolism and thrombosis of the mesenteric arteries
the symptoms during life are entirely identical with those
observed in the so-called colic of horses, as has been determined
by numerous observations. The partial paralysis of the bowel,
which is brought on by the embolism and thrombosis of the
mesenteric arteries, forms in great part the chief and leading
feature of the series of symptoms known as the " colic " of

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PARASITES OP ANIMALS

horses. The palsy of the bowel which arises in this way may
explain also the frequent ruptures of the digestive canal and
the greater number of its changes in position. The latter are
specially favored by the structure of the abdominal viscera in
the horse.

15. The old changes which one finds in the peripheral
branches of the anterior mesenteric artery, in the form of
expired and partly absorbed embolic and thrombolic processes
(pigmentation, arterial and venous thrombi), particularly in
connection with those arteries which are seats of the aneurism,
decisively prove that the large majority of colics resulting in
recovery, so far as they do not depend upon known injuries,
are caused by paralysis of the bowel from embolism and
thrombosis. The sudden occurrence, course, and result of these
kinds of colics also testify to their embolic origin.

16. The cedematous, inflammatory, and hemorrhagic pro-
cesses that one often finds described as the cause of death in colic,
almost exclusively depend on thrombosis and embolism of the
mesenteric arteries, the cases forming about 40 to 50 per cent,
bf all fatal colics.

17. The rapid course in fatal colics, as well as the prepon-
derating symptoms of dyspnoea in cases of recovery, is finally
due to the abnormal development of gas in the alimentary
canal. In addition to the diminution of the respiratory surface
by the lofty position of the diaphragm, a direct gas-poisoning
(carbonic acid and sulphuretted hydrogen) probably contributes
to the intensity of the symptoms and the rapid course by
diffusion of the abnormally developed gas out of the intestinal
canal into the blood.

18. The variety of the anatomical derangements caused by
embolism and thrombosis of the intestinal arteries is faithfully
mirrored by the variety of the clinical symptoms and the
different degrees in the intensity and course of the colic.

19. Amongst every 100 horses afflicted with internal disease,
40 are ill with colic. Among any hundred deceased horses 40
have perished from colic, and among 100 colic patients 87
recover and 13 die. The figures prove that neither amongst
the epizootic nor sporadic diseases of horses is there any other
affection which occurs so frequently, or claims anything like so
many victims. Like the frequency of the worm-aneurism, the
amount of disease and mortality increases with advancing age.
The etiology of the colic of horses finds in the thrombosis aud

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373

embolism of the mesenteric arteries, with the consequent
paralysis of the bowel, an all-sufficient explanation, whilst
the causes of colic hitherto accepted were for the most part
insufficient.

20. In a great number of cases the thrombus of the worm-
aneurism is continued past the mouth of the anterior mesenteric
artery, into the lumen of the aorta, and, as such, is the exclusive
cause of the embolisms of the pelvic and crural arteries which bring
about the intermittent hobblings (the author says " intermit-
terenden Hinken," not " Hahnentritten," the usual equivalent
term for stringhalt). Considering the excessive frequency of
the thrombus being continued into the aorta, it becomes highly
probable that a great part of the diseases and lameness of the
posterior extremities ('* Hiift und Kreuzlahme, unsichtbarer
Spath, &c," which may be rendered " sciatic and hip or spinal
lameness, obscure spavin, &c") are due to occlusion of the arteries.

21. Owing to the fibrous thickening of the connective tissue
of the root of the anterior mesenteric round the aneurism, and g
to the considerable size of the latter, disturbances of the inner-
vation of the intestine, (as well as) hindrances to the passage of
the chyle, and irregularities in the portal circulation may be
created, which may well lie at the root of many chronic dis-
turbances of digestion in horses.

22. Considering the great losses and heavy social disad-
vantages that are occasioned by the colic of horses to the
horse-breeder, to agriculture, and to the general welfare, it is
of the highest importance to discover means which should
prevent the introduction of the embryos with the food, and, as
a consequence, the migration of the palisade worms into the
mesenteric arteries of, the horse.

I wish it to be distinctly understood that the above summary
is translated from Bollinger (Die Kolik ; s. 257). Instructive
cases have been recorded both at home and abroad. Prof.
Varnell has remarked that (t foals and yearlings suffer more
from parasites in the paddocks than they do on adjoining farms
where only a few animals are bred." This is explained by
the relatively greater amount of egg-dispersion proceeding from
the infected brood-mares. It is quite evident that the lives of
many valuable animals are annually sacrificed by the neglect of
hygienic arrangements. The palisade worm is chiefly destruc-
tive to young animals, and as Mr. Percivall has well remarked,
these parasites are " commonly the cause of lingering and hidden

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PAKA8ITES OF ANIMALS

disease, terminating in death," without any suspicion on the part
of the practitioner as to the nature of the malady. Instructive
cases of this form of helminthiasis are given by Messrs Littler,
Wyer, Harris, Meyrick, Litt, Percivall, Tindal, Walters,
Brett, Aitken, Mead, Clancy, Baird, Mercer, Wright, Seaman,
Hepburn, and others.

Second only in clinical importance is the little four-spined
strongyle (S. tetr acanthus). The sexes, often seen united, are
nearly of equal size, the largest females reaching nearly §".
They infest the ceecum and colon, and have been found in all
varieties of the horse, ass, and mule. The worm occurs in
immense numbers and is a true blood-sucker. Its presence
occasions severe colic and other violent symptoms, often proving
fatal to the bearer. As already announced, in connection with
my account of the tapeworms of the horse, this little worm
may produce a virulent epidemic (epizooty). In the sexually -

immature state the worm occu-
pies the walls of the large intes-
tine, where it gives rise to con-
gestion, ecchymosis, inflamma-
tion, and the formation of pus
deposits. The species is readily
recognised by its bright red
color, by the four conical spines
surrounding the mouth, by the

Fig. 63— Larvffi of Strongylus tetracantlius. a, two neck-bristleS, and by the

from the walls of the intestine ^natural size) ; _ pi

b, the same (enlarged) ; *, an injury ; c, younger long three-lobed hood of the
specimen (in sM); d, the same (enlarged). Ori- .

giuai. male, the posterior three-cleft

ray having a rudimentary or fourth branch attached to its
outer edge. In some specimens sent to me by Mr Whitney, I
found this supplementary process fully twice as long as
Schneider has represented it.

From the earliest times this entozoon has been confounded
with the palisade worm. Rudolphi and several of his successors,
and also in recent times Ercolani and Colin, regarded this worm
as the progeny of Strongylus armatus. During my earlier
examinations I likewise fell into the error of describing the
immature worm as representing a new species. The parasites
described by me as Trichonemes {T. arcuata) were identical
with those which Prof. Dick had previously described as
" worms at different stages of growth," in his MS. sent to Dr
Knox, 1836. Parasites of this kind were described by Dr

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Knox as "Animals similar to Trichina;" by Diesing as the
" Nematoideum equi caballi f by Mr. Littler as " Extremely
small ascarides," in a letter to Mr Varnell ; by Mr Varnell
himself as " Entozoa in various stages of growth and by
Prof. Williams as " Entozoa from the intestinal walls/' in a
letter to myself, dated March 13th, 1873. In reference,
however, to Mr VarnelPs account of Mr Littler' s specimens I
may observe that the appearances which he at first merely
described as "blood spots," he afterwards characterised as
dark points "containing young worms in various stages of
growth."

As regards the course of development of this worm we have
yet much to learn. Although the worm is a frequent cause of
epizooty in this country it appears to be but little known on
the Continent. Krabbe makes no mention of the helminthiasis
set up by the four-spined strongyle, but he points out that the
young occupy the mucous membrane, in which situation they
lie coiled so as to present to the naked eye the appearance of
little dark spots (Husdyrenes Indvoldsorme, 1872, p. 17, ' Aftryk.
af Tidsskr. for Vet/). However, Leuckart's account of the
appearances presented in a case brought under his notice is
instructive. He writes : — " I have hitherto had only a single
opportunity of examining the strongyle capsules in the intestinal
membrane of the horse. Their presence is limited to the caecum
and colon, but they are so abundant in this situation that their
numbers may be estimated by many hundreds. It was thus
likewise in the case in question, the investigation of which by
myself was rendered possible through the friendliness of Prof.
Haubner of Dresden. The capsules were of oval form, and
glimmered through the mucous membrane as opaque spots,
mostly from one to three millimetres in size. In several of
these capsules nothing was found beyond a greasy mass of a
brownish color, which might readily be taken for a tuberculous
substance ; but the greater number of them contained a coiled
worm, from three to six millimetres long, their breadth being
0-15 to 0-26 mm. (which is ^' to . They exhibited a highly
colored stout intestine, and a thick-walled oral capsule of
0*022 mm. in depth and 0*025 mm. in breadth. On the dorsal side
two three-cornered chitinous lamellae arise from the shallow
floor of the small oral capsule. The cuticle, notwithstanding
its firm structure, was still destitute of annulations. The tail
(0-15 to 0*18 mm. in length) was strongly marked off from the

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PARASITES OF ANIMALS

rest of the body, being of a slender cylindrical form with a
rounded-off extremity. The development of the sexual apparatus
had not yet commenced. Notwithstanding the great differences
of size presented by the body, the structure of all examples was
exactly the same to the minutest particular, without exception.
Also the smallest specimens, which scarcely measured one milli-
metre, found in capsules of 0*3 mm. in diameter, were distinguish-
able only by the absence of the oral cup, whose position was
represented by a slender and thickened chitinous cylinder, as
obtains in the earliest parasitic juvenile condition of Dochmius
trig ono ceplialus. The transformation to the form presenting an
oral cup occurs through a moulting, which is accomplished
already in examples of 1*5 mm. in diameter. Later, also, the
worms cast their skins in their capsules, without, however,
changing the oral cup. In regard to the final purpose of this
metamorphosis, my investigations have left me entirely in the
lurch ; nevertheless, I do not entertain the smallest doubt that
the worms which I have here described are the larval forms of
Strongylus tetr acanthus."

From numerous examinations I have satisfied myself that
the worms after escaping the walls of the intestine — and
they may often be observed in the very act of passing —
re-enter the lumen of the bowel to undergo another change
of skin prior to acquiring the adult state. This they accom-
plish by rolling themselves within the faecal matter of the
horse's intestine. The best examples I have seen of this
phenomenon occurred in a case for the clinical particulars of

which I am indebted to Mr Cawthron.
Most interesting was it to notice these
immature worms, each coiled within a
sort of cocoon, which Mr Cawthron
termed a cyst. All the forty little
cocoons more or less resembled pills,
the bright red color of their contained
worms strongly contrasting with the
fig 64.-iS%sts or pellets con- dark color of the cocoons. They con-
1SSLSL X-imenlTS listed of compressed debris, which under
ie,eiadBan(i4taii sh^)oi X^wornu the microscope showed many common
original, forms of vegetable hairs and paren-

chyma, besides raphides and chlorophyll-granules. Internally,
there was a cavity corresponding with the shape of the worm.
In one instance I noticed that the worm had nearly completed

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377

its ecdysis, a portion of the old skin still remaining attached to
the tail.

As already remarked, the evidence respecting the frequency
and destructiveness of this little worm in England is now quite
overwhelming. In a series of papers contributed to the ' Vete-
rinarian ' (too long for full quotation here), I have endea-
voured to do justice to the " finds " and observations of those
members of the veterinary profession who were good enough to
supply me with valuable notes and communications. In parti-
cular must my indebtedness to Mr Rees Lloyd be acknowledged,
for, as previously observed, he it was who first recognised the
parasitic character of the Welsh epizootic outbreaks. In the
Deangunid and Talybont districts these strongyles proved
terribly fatal to mountain ponies. It appears that the owners
of the animals, as soon as they perceived anything amiss,
at once disposed of them by sale, evidently anticipating fatal
results sooner or later. The facts connected with some of the
isolated cases brought under Mr Lloyd's care are especially
interesting, as showing the virulence of the symptoms set up.
Thus on the 9th of Feb., 1875, some time after I had identified
the species from specimens he had sent me, Mr Lloyd writes
as follows : — " The last case I had was one which had been
sold in this way, and which had suffered now and then from
colicky pains for the space of about two months. The animal
had been drenched with febrifuges and rubbed with stimulating
liniments about the throat. However, I was sent for one
evening, about an hour before the patient's death. I soon
diagnosed the case as parasitic, and at the same time considered
it to be a hopeless one. I remained with it the whole of the
time, of which about forty-five minutes of the most acute
pain was borne by the trembling beast, which was leaping,
rolling, and tossing itself about with astonishing rapidity.
The bulging eyes, gnashing teeth, foaming mouth, and sharp
peculiar hoarseness, were pitiable to behold; when suddenly all
was silent, he quietly rose to his feet, and nipped the grass as
if nought had troubled him. I then trotted him quietly up
a few yards of rising ground in the corner of the field, when
he immediately got down to rise no more. The next day I
examined him, and found myriads of the four-spined strongyles,
a large number being encysted." Mr Rees Lloyd's account of
this case is so graphic that I have reproduced it without
abridgment. Speaking of another patient, a mare, he says,

378

l'AUASITES OF ANIMALS

" she had voided thousands of those parasites, and was in a
frightfully emaciated condition, but beyond a craving appetite
there was little else to be noticed." Notwithstanding the few
diagnostic indications afforded, Mr Lloyd, being led by the
history of the case to suspect worms, at once examined the
fasces, when he discovered thousands of these minute nematodes.
Clinically, these facts ought not to be lost sight of. In addition
to Mr Lloyd's cases I have received valuable particulars of
others at the hands of Prof. Williams, Messrs Cawthron,
A. Clarke, T. Gerrard, D. M. Storrar, and J. W. Whitney.

Practically, it is important to inform the persons most in-
terested that an active " drench " may be serviceable in dislodg-
ing the free intestinal worms, but the administration of purga-
tives must not be persevered in. As in the somewhat parallel
case of Trichinosis in the human subject, the fatality of the
disorder depends not upon the free and mature worms, but upon
the migrating and sexually-immature forms. All attempts by
means of active drugs to poison the entozoa, when once they
have gained access to the tissues (whether actually capsuled or
not), are worse than useless. By all means let the animals have
a dose of aloes in the first instance, followed by warm bran
mashes ; but thereafter let every care be taken to support the
patient's strength. Especially should the exhibition of turpen-
tine be avoided. Without doubt the cause of this, as of other
similar epidemics, is primarily referable to atmospheric conditions
which favor the multiplication of parasites. As the practical
man cannot alter these climatal changes, he must do his best to
check the disorder by removing the victims to new localities ;
or, if the animals must remain in infected districts, by supplying
them with various kinds of artificially prepared fodder, supple-
mented by carefully filtered water. In this way, I believe,
epidemics may be arrested, but they cannot be stamped out
altogether, except by the adoption of measures which would be
alike impracticable and unwarrantable.

Passing to the consideration of other intestinal nematodes, the
next in importance is the large lumbricoid (Ascaris megaloce-
phala) found in all solipeds, including the zebra. Whilst the
male worms rarely exceed seven inches in length, the females
sometimes reach seventeen inches. Science is indebted to
Schneider for setting at rest all doubt as to the specific dis-
tinctness of this worm. The far larger number of caudal papilla?
at once distinguishes it from the lumbricoid of man and the

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379

hog. The equine Ascaris may occur in any part of the alimen-
tary canal, but the small gut forms its proper head-quarters.
The entire course of development of this worm has not been
traced ; nevertheless, Heller found human lumbricoids measur-
ing less than the eighth of an inch. It is not likely that any
intermediate host is necessary for the growth of the larvae, prior
to their access to the definitive host. I have reared the larvae
in impure water and in moist horse-dung, up to the size of ^ of
an inch. They were then furnished with a completely-formed
digestive apparatus. Davaine kept the intra-chorional embryos
alive in water for five or six years. His experiments on rats,
dogs, and on a cow, led to no decisive results ; but it is impor-
tant to know that the eggs of lumbricoids effectually resist
dryness. According to Davaine, however, embryonal develop-
ment is thus arrested (except in Ascaris tetraptera of the
mouse).

Seeing how readily the most ordinary attention to cleanliness
must suffice to prevent lumbricoid helminthism, it is scandalous
that so many severe cases of disease from this source should
ever and anon turn up and be reported. In no properly con-
ducted stable are these large entozoa ever to be seen in any
considerable numbers ; for so long as the water-supply is good
and the fodder clean there is no possibility of infection. A
fertile source of infection, however, results from allowing horses
to drink at foul road-side ponds and from open waters in the
vicinity of stables and paddocks where foals are reared. Into
the clinical bearings of the subject I do not enter, but a host
of interesting records of lumbricoid disease may be found in
veterinary journals, both home and foreign. These have their
counterpart in the very similar cases recorded in the medical
journals, and quoted by me in the 34th bibliography of this
work. From Sonsino's report these worms do not appear very
common in Egypt, but the veterinary inspector, Dr Zunhinett,
had occasionally met with them. From Messrs W. Awde,
J. B. Wolstenholme, and other English veterinary surgeons, I
have received notes of interesting cases, but in this connec-
tion I can only further refer to the published cases of Messrs
Anderson, Boddington, Cartwright, Harrison, Moir, and Wallis.
The French cases, by M. Cambron and by M. Veret, are par-
ticularly instructive. Many of the cases give fatal results.
In one fatal instance a pupil of mine counted over 1200 of
these worms, and in a similar fatal case Mr Lewis reports that

380

PARASITES OF ANIMALS

he found the small intestine literally crammed, some thousands
of worms being huddled together in large masses.

The next nematode of general interest is the pinworm
{Oxyuris curvula). Professional men often confound it with
the palisade worm, and it has even been mistaken for the rat-
tail maggot (Helophilus) . The longest males measure If",
and the females often beyond 4 inches. This worm infests the
colon in great numbers, the species being easily recognised by
its long subulate tail. Like its much smaller congener infest-
ing man, this worm occasions severe local irritation, clusters of
the eggs often accumulating to form yellow inprustations at the

Fig. 65. — Head of Oxyuris curvula. Highly magnified. After Busk.

verge of the anus. Equine pinworms are vegetable feeders,
and, like human Oxyurides, are conveyed to the bearer in a
direct manner. No horse properly looked after can be infested
by these worms. Local washings and stable cleanliness being
secured by an attentive groom, the animals are safe. Prophy-
lactic measures of this kind are all-powerful against infection.
Notwithstanding the ease, however, with which the oxyuris
disorder may both be prevented and cured, we find it prevails
extensively everywhere, alike in mankind and in solipeds. Dr
Sonsino found these parasites abundant in Egypt, some of
the worms reaching a length of nearly five inches (120 mm.).
Mr Emmerson has given an interesting account of the prejudicial
effects of these entozoa in the horses of Singapore.

One of the most remarkable equine parasites is that which I
am in the habit of calling the large-mouthed maw-worm (Spi-
roptera megastoma), in contra-distinction to the small-mouthed
species (S. microstoma). In this country the worm has
attracted little notice, but through the kindness of Mr Spooner

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381

Hart, of Calcutta, and of Mr Percivall, of the 11th Hussars,
stationed at Umballa, I have had abundant opportunities of
examining this entozoon and the singular pathological appear-
ances which it occasions. This parasite was first described by
Rudolphi, who says : — " Spiroptera capitis discreti ore magno
nudo, cauda feminae rectiuscula acuta, mavis simpliciter spirali,
corpusculis rotundis ad basim penis styliformis." The worm
was afterwards observed by Schultze, Chabert, and frequently
also by Andral, but the best accounts of it are those given by
Gurlt, Yalenciennes, and Dujardin. Schneider has likewise
done much to set at rest disputed points. Respecting the
Spiroptere du Gheval, Dujardin, writing in 1844, observes that
" Rudolphi at first studied this helminth from examples found in
great number by Reckleben, at Berlin, in tubercles of the
stomach of two horses. Quite recently, M. Yalenciennes, at
Paris, has found it frequently in tumours, from twenty to forty
millimetres in size, in the stomach of eleven horses out of
twenty-five that he had subjected to this kind of research.
These tumours, lodged between the mucous and muscular layers
of the digestive canal, are perforated by several holes travers-
ing the mucous membrane. They are divided internally by a
number of folds into numerous intercommunicating cavities,
and sometimes filled with solid mucus and very many spirop-
teras. It is from examples collected by M. Yalenciennes that
I have been able to study the parasite."

As regards the description of the worm, it is almost needless
to say that Dujardin' s account is minute and admirable in all
respects. In fact, no naturalist ever exceeded the Rennes savant
in carefulness and accuracy of detail. An interesting point con-
nected with these stomach-worms lies in the circumstance that
Gurlt recognised two varieties, one of which he termed Sp. meg.,
var. major. It remained for Schneider to show that the larger
worms formed an altogether distinct species, which he termed
Filaria microstoma (' Monogr./ 1. c, 1866, s. 98). It was not
unnatural that Rudolphi and his successors should confound
these two forms together, and it is also not a little curious
that the smaller of the two species has the larger mouth.
Practically, veterinarians will probably rest content to know
that whilst the Spiroptera megastoma occupies tumours in the
walls of the stomach, the S. microstoma is always to be found
free in the cavity of that organ. Any helminthologist who
may chance to have read the Ceylon Company's report on the

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PARASITES OF ANIMALS

fatal epidemic affecting the mules of the Mauritius in 1876 can
scarcely fail to have observed that the worm called Ascaris
vermicularis by Mr Bradshaw is none other than our Sp.
megastoma. The description of the tumours as " reticulated "
sufficiently explains their honeycomb- like appearance, but I
think that the expression "alveolar" would better convey their
true pathological character. Mr Spooner Hart compared these
structures, which he terms " abodes/' to mole-hills, ' but there
is no good ground for supposing that the wanderings of the
parasites are in any sense comparable to the burrowings of the
mole. In like manner the expression " nidus," employed by
Mr Bradshaw, though suggestive of their nest-like appearance,
is to some extent misleading, as it implies that the worms form
a nide or brood. Possibly, it may turn out that all the nema-
todes in each tumour have been bred in the spot where they are
found, but hitherto they have only been seen in the adult state.
Earlier stages of growth should be diligently sought for.
Widely dissimilar as the two maw-worms are, it would not
greatly surprise me to learn that Sp. megastoma and S. micro-
stoma are dimorphic conditions of one and the same entozoon.
At all events, Ercolani's determination of the relations subsist-
ing between Ascaris inflexa and A. vesicularis suggests a possible
analogy of this kind. I may mention that the male Bpiroptera
megastoma reaches nearly one third and the female one half
of an inch in length. A constriction separates the head
from the body. The mouth is surrounded by four thick
horny lips, the dorso-ventral pair being the larger. The tail
of the male is spirally twisted, and furnished with lateral
bands supported by three or four ribs. It carries two curved
spicules of unequal size. There are five pairs of caudal
papillae, the tail being bluntly pointed in both sexes. The
vulva of the female is placed about \" below the head. The
eggs are linear or very narrow, and furnished with thick shells.
According to Sonsino, who found Sp. megastoma in five out of
sixteen Egyptian horses, the verminiferous growths are usually
seated near the pyloric end of the stomach, as many as four
tumours occurring at one time. Neither Sonsino nor any other
observers already quoted appear to think that these morbid
changes in any way interfere with the healthy performance of the
gastric functions. However, I am of opinion that at least one
recorded fatal case of parasitism, producing rupture of the
stomach, affords an instance, however rare, of the injurious

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383

action of this entozoon. It is reported under the signature
of " Argus," quoted below.

In this connection I may mention that in 1864 Prof. Axe
observed some small worms, scarcely visible to the naked eye,
in the mucous membrane of the stomach of a donkey, the same
worms being subsequently observed in three other donkeys
brought to the dissecting room of the Royal Veterinary College.
From the examination of a drawing of one of the male worms,
executed by Prof. Simonds, I am led to believe that the
parasites are entirely new to science. The hood being well
marked there can be no doubt as to the strongyloid affinities
of the worm. I therefore propose to call the worm after its
discoverer (Strongylus Axei).

In regard to Sp. microstoma, the males measure up to §",
whilst the females have a long diameter of £" or rather more
(10'"). The small-mouthed maw- worm lives free in the stomach,
and, as Krabbe observes, not unfrequently in very considerable
' numbers. It does not appear to be capable of injuring the host.
One of the most interesting equine nematodes is the eye-
worm. Most veterinary writers speak of it as the Filaria oculi,
but to helminthologists it is better known by the more correct
designation, F. pajpillosa. Though commonly obtained from
the eyeball and its tunics, the worm infests various tissues and
organs of the body, being found in the thorax, abdomen, mem-
branes of the brain, muscles, and cellular tissues. It infests
the ass and mule, and also horned ruminants. The males
attain a length of three inches and the females seven inches.
The head is broad, with a gaping mouth armed with a
ring of chitine and two prominent denticles. There are
also two papillae on the neck near the middle line, besides
sixteen caudal papillae, eight on either side. The tail of the
male is spirally twisted, that of the female only slightly curved.
Notwithstanding the many opportunities afforded of examining
this parasite in the fresh state, very little is known respecting
its origin and course of development. Dr Manson, who found
that the mouth was armed with a five- or six-toothed oral saw,
considers that the eye is not a proper resting place for the
parasite, and that when one wandering worm comes across the
track of another it follows it up from sexual instinct, and thus
several may be found together in one place. The tracks are
readily seen by the naked eye. Dr Sonsino speaks of it as a
" yellow line." This Italian observer found the worm in twelve

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PARASITES OP ANIMALS

out of the sixteen solipeds he examined during the plague.
Each horse showed from two to a dozen worms " in the peri-
toneal cavity, wandering free on the serous lining, without
causing any apparent mischief to the membrane.-" On one
occasion Sonsino found the worm in the liver. From the
similarity of habit there can be little doubt that the cases
of guinea-worm (F. medinensis) recorded by Clarkson and
others, as occurring in the horse, were merely examples of
F. papillosa. I think so all the more because the lamented Fed-
schenko verbally expressed to me his astonishment that I had
in my introductory treatise (p. 387) spoken of the Dracunculus
as an equine parasite. I did so on the authority of others.
To the Rev. Horace Waller I am indebted for specimens of the
eye-worm brought from Assam, and to Mr Spooner Hart for
others sent from India. For examples occurring in England I
am indebted to Mr Haydon Leggett, who, in 1875, sent me
three specimens extracted from the eye of a five-year-old mare.
Mr Steel has also given me an example of F. jpapillosa taken
from the peritoneum of a donkey. Similar cases are constantly
occurring in the practice of veterinarians in Hindostan. Highly
interesting Indian cases are recorded by Kennedy, Molyneux,
Twining, and Breton, and in addition to these I may also par-
ticularise those of Macnamara, C. Percivall, Hickman, Clarkson,
Skeavington, and Jeaffreson. The cases by Lee and Grellier also
deserve attention.

Another species of thread-worm {Filaria lacrymalis) is occa-
sionally found in the horse between the lids and eyeball. It is
a comparatively small and harmless parasite, the males measur-
ing \" in length and the females It also infests the ox.
Both the large and small eye-worms are vivipai*ous, and, not
improbably, both of them are the means of conveying embryonic
Filarise into the circulation. Be this as it may, we owe to Dr
Sonsino the discovery of haematozoa in an Egyptian horse.
The larval worm was provisionally named by him Filaria san-
guinis equi. The microscopic nematodes closely resemble the
larvEe of F. sanguinis hominis, but they are smaller. The horse
from whose blood Dr Sonsino obtained the minute worms was
also found, by post-mortem examination, to have been infested
by Filaria jpapillosa, a circumstance which naturally suggested
a genetic relation between the larval and adult parasites.
Similar, if not the same, microscopic worms had been previously
discovered by Wedl, who/ primarily and independently regarded

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385

them as embryos of F. papillosa. Another curious filaria-like
entozoon is the reticulated threadworm (Onchocerca reticulata).
In England we have no acquaintance with this singular parasite,
but it appears to be tolerably common in Italy. Excellent
figures of it have been given by Diesing. Both males and
females are in the habit of coiling themselves within the muscles,
where they are found invested by a capsule of connective tissue.
When unrolled the sexes are found of equal size, acquiring a
length of The worm has a simple unarmed mouth, its

body being marked by a series of annulations formed of incom-
pletely anastomosing rings. It does not appear to possess any
clinical importance.

In connection with the equine nematodes I need only mention
the lung- worm (Strongylus micrurus). Its importance in relation
to the production of husk or parasitic bronchitis in calves has
already been considered. The worm is rarely productive of
mischief amongst solipeds, nevertheless, in the dissecting-room
subjects at the Royal Veterinary College, the presence of these
parasites in the lungs is frequently noticed. Lastly, it only
remains for me to observe that the renal strongyle (8. gigas) is
occasionally seen in the horse. In 1792 M. Chabert found one
in the left kidney, and similar cases have since either been
witnessed or reported by Rudolphi and Leblanc.

Of the numerous insect parasites and tormentors of solipeds
the gadflies (CEstridce) demand chief attention. For special
description of the forms, Brauer's monograph is the most, and,
in fact, the only reliable authority. Here it is not possible to
give the characters of the various equine species, of which at
least half a dozen are known to science. As remarked by me
in the special chapter contributed to Prof. Williams' well-known
veterinary treatise, the common gad-fly (Gastrophilus equi)
attacks the animal whilst grazing late in the summer, its object
being, not to derive sustenance, but to deposit its eggs. This
it accomplishes by means of a glutinous excretion, causing the
ova to adhere to the hairs. The parts selected are chiefly those
of the shoulder, base of the neck, and inner part of the fore legs,
especially about the knees, for in these situations the horse will
have no difficulty in reaching the ova with its tongue. When
the animal licks those parts of the coat where the eggs have
been placed, the moisture of the tongue, aided by warmth
hatches the ova, and in something less than three weeks from
the time of the deposition of the eggs, the larvas make their

386

rARA SITES OF ANIMALS

escape. As maggots they are next transferred to the mouth,
and ultimately to the stomach along with food and drink. A
great many larvae perish during this passive mode of immigra-
tion, some being dropped from the mouth, and others being
crushed in the fodder during mastication. It has been calcu-
lated that out of the many hundreds of eggs deposited on a
single horse, scarcely one out of fifty of the larvae arrive within
the stomach. Notwithstanding this waste the interior of the
stomach may become completely covered with " bots." Whether
there be few or many, they are anchored in this situation chiefly
by means of two large cephalic hooks. After the bots have
attained perfect growth they voluntarily loosen their hold, and
allow themselves to be carried along the alimentary canal until
they escape with the fasces. Many persons suppose that during
their passage through the intestinal canal they re-attach
themselves to the mucous membrane, thereby occasioning severe
intestinal irritation. This is an error. In all cases they sooner
or later fall to the ground, and when transferred to the soil
they bury themselves beneath the surface, in order to undergo
transformation into the pupa condition. Having remained in
the earth for a period of six or seven weeks they finally emerge
from their pupal-cocoons as perfect dipterous insects. It
thus appears that bots ordinarily pass about eight months of
their lifetime in the digestive organs of the horse.

That they are capable of giving rise to severe disease there
can be no doubt, but it is not often that the disorder is
correctly diagnosed, since it is only by the passage of the
larvae that the practitioner can be made aware of their presence.
Mr J. S. Wood has published a case of tetanus in a mare,
associated with the larvae of (Estrus equi, and Mr J. T. Brewer
has also given a case where the duodenum was perforated by
bots. Mr Goodworth records an instance of pyloric obstruction
from the same cause, and Mr W. Coupe informed me in 1876
that he had a drove of foreign ponies under his care, all of
which suffered irritation from haemorrhoidal bots. He removed
them with a pair of forceps. Although frequently said to do so,
the common bot does not attach itself to the rectum before finally
escaping the host. The larvae of O. hamorrJioidalis normally
reside there. In this situation they seriously inconvenience the
bearer. The bots of O. nasalis are often confounded with those
which ordinarily occupy the stomach of the bearer. The larvae of
G. nasalis commonly reside in the duodenum near the pylorus.

SOLIDUNGULA

387

According to Schwab and Brauer, they rarely occupy the
stomach. As occurs in the common species, this bot passes
away with the faeces, and does not attach itself to the lower
bowel. The bots of Brauer's G. inermis much resemble
those of G. equi, but they are much smaller and attach
themselves to the wall of the small intestine. The bots of
G. pecorum, which dwell in the rectum, are readily recognised
by their peculiar form and scanty spination. They are pointed
in front and truncated posteriorly. An assinine variety of
G. equi has been described by Bilharz, whilst another distinct
species (G. flavipes) attacks the ass and mule. The bot-larvas
of the latter host require recognition and description. A great
variety of other equine bot-flies have been described, but all,
or nearly all, of them are mere synonyms of the above-men-
tioned forms. For the limitation of the species I accept
Brauer's authority, and likewise his nomenclature. A great
deal of nonsense has been written respecting bots. It is a
relief to believe that G. (CEstrus) veterinus, G. ferruginatus,
G. jubarum, G. (OE.) Glarkii, G. salutiferus, G. subjacens, and
many others, are not good species, at least that they are mere
synonyms. In regard to the occurrence of subcutaneous bot-
like maggots in the horse and ass, no doubt need exist on this
point. I am indebted to Mr Percy Gregory for characteristic
specimens taken from the back, neck, and withers of a four-year-
old gelding. They appear to correspond with the Sypoderma
Loiseti of Joly. Similar maggots have been found in the ass by
Herr Erber, but Brauer refers these to H. silenus. Prof. Briick-
miiller published a case where the brain was infested by larvae ;
and Mr Shipley has sent me an example of H. equi, which he
states he removed from the choroid plexus of the brain. In ad-
dition to the cases by Woods, Goodworth, and Brewer, already
quoted, others have been published by Tyndal and Cartwright.

Amongst the numerous other parasitic dipterous larvae one
must notice the rat-tailed maggots (Helophilus) . A genuine
instance of this kind has been brought under my observation,
but the example recorded by Professor Axe was spurious.
Professor Simonds and myself saw this supposed maggot,
which was merely a very stout and pregnant Oxyuris curvula.
Another genuine case was published by Mr. Stanley. This is
quoted by A. Numan in his essay on Ganurus. I have previously
mentioned my having received an Helophilus-larva that had
passed from the human body. One of the most troublesome

PARASITES OF ANIMALS

external parasites is the so-called horse-tick or forest-fly
(Hippobosca equina). They attack the abdomen, flanks, and
inner part of the thighs in great numbers, occasioning great
distress to the bearer. Being of leathery toughness their
bodies are not easily crushed, and they are removed only with
great difficulty. There is an equine disease in Sweden called
Stachra, which is erroneously attributed to injuries produced by
a species of fly-maggot (Lixus) which lives on the fine-leaved
water-drop wort (Phellandrium) . As regards the so-called
free parasites, or rather non-parasitic obnoxious insects, which
torment solipeds, it is impossible even to enumerate them.
The tsetse of South Africa (Glossina morsitans) is terribly
fatal to the horse, but it is said that the mule, ass, and zebra
do not suffer from its bites — an immunity shared by swine,
goats, antelopes, and man himself. Major Vardon's rash
experiment (based on the supposition that horses deprived of
fresh green food would not suffer from the attacks of the fly)
proved fatal to an animal which he purposely exposed on a
much infested hill-top. The horse died ten days after it was
bitten. According to Chapman, the bites of four tsetse flies
are sufficient to kill an ox, but in man the irritation produced
is very slight. Amongst other insects proving troublesome to
solipeds may be mentioned the leg-sticker (Stomoxys calcitrans) ,
the clegg (Hcematopota pluvialis) which is very abundant in the
West Highlands, various species of Tabanidce and Asilidce
(Tabanus autumnalis , T. bovinus, Ghrysops cmcutiens, Asilus
crabroniformis), and also a host of ordinary flies and gnats
(Muscidce and Tifulidce), as, for example, Anthomyia meteorica
and Guhx equinus. In India the bite of a species of Simulia
gives rise to the formation of open sores of the most intractable
character. As regards hemipterous insects it may be said that
many species of lice (Anoplura) produce what is called phthiriasis
or lousiness in the horse, some of them being derived from
poultry. The best known species are Trichodectes equi, T.
scalaris, Hamatopinus equi, B. vituli, H. eurysternus, and the
ass-louse {E. asini). Of the half dozen or more species infesting
the hen (belonging to the genera Goniocotes, Liotheum, &c.)
it is not probable that more than one or, at most, two of them
are concerned in the production of poultry-lousiness in the
horse. As an equine disorder this kind of phthiriasis was first
described by Bouley. Cases in England have been observed
by Messrs. Henderson, Moore, and Woodger. For some

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389

caccount of cases of lousiness due to Hcematopinus I am indebted
to Mr S. Butters. As regards the scab, itch, and mange
insects or mites (Acaridce), three perfectly distinct forms are
known. Adopting M. Megnin's classification they are Scar-
coptes scabiei, var. equi, Psoroptes longirostris, var. equi (being
the Dermatodectes equi of Gerlach), and Ohorioptes spathiferus,
var. equi, which is the Symbiotes equi of Gerlach. All the
species have been beautifully illustrated by M. Megnin, whose
memoir has dispersed many of the clouds of error and mis-
representation which have hitherto surrounded the subject.
Whilst Psoroptes forms the true horse-mite, and attacks various
parts of the body, Ohorioptes confines its attacks to the posterior
regions. Messrs South and Day and myself have verified
some of the facts recorded by Megnin in respect of the struc-
ture and habits of this last species. Another kind of mite
(Glyciphagus hippopodos) is stated to infest the ulcerated feet
of horses. It would appear that no true ticks properly belong
to solipeds ; nevertheless, the common cattle-tick (Ixodes bovis)
occasionally attacks horses. Probably several other species of
Ixodidee, known to infest other animals, behave in the same
way. The Arachnidan called Pentastoma tcenioides, though,
properly belonging to the dog, has on several occasions been
detected in the nasal or frontal sinuses of the horse. Such,
instances are recorded by Chabert and Greve. The largest
example of this singular entozoon seen by myself was obtained
from the same situation, and presented to me by the late
Mr C. B. Rose, whose writings I have frequently quoted in
connection with the Coenuri of rabbits.

Bibliogeaphy (No. 50). — Aitken, J., "Worms in the Sper-
matic Artery of a Colt," 'Veterinarian/ p. 683, 1855. — Ander-
son, J., " A Case of Strangulation of the Ileum (with Lunibrici),"
' Veterinarian/ p. 261, 1859. — (Anonymous), "Rupture of the
Stomach, associated with the existence of Cysts between its
Coats containing Worms/' ' Veterinarian/ March, 1864, p. 151.
— (Anon.), " Extraction of Filaria oculi from the Horse," by
"Miles," 'Veterinarian/ 1864, p. 218.— (Anon.), "Case of
F. oculi in the Horse," 'Veterinarian/ 1864, p. 218. — (Anon.),
" Worms in the Coats of the Stomach of a Horse," by "Argus,"
in the 'Veterinarian/ 1865, p. 151. — Baird, W., "Notice of
Sclerostoma in the Testicle of the Horse," ' Proc. Zool. Soc./
1861. — Blancharcl, " Anoplocephala perfoliate," 'Ann. des Sci.
Nat./ 3rd scr., torn, x, p. 345. — Bollinger, 0., ' Die Kolik der

390

PARASITES OF ANIMALS

Pferde und das Wurmaneurisma der Eingeweidearterien/ Mun-
chen, 1870. (Reviewed by myself in the ' Veterinarian/ Jan.—
April, 1874.) — Bovett, " Existence of FilariaB in a Sinuous
Ulcer of the Withers of a Mare/' '"Veterinarian/ p. 515, 1861.
— Bradshaiv, 0., " Death of Mules from Parasitic Disease in
the Mauritius (with remarks by Dr Cobbold)," ' Veterinarian/
Dec, 1876, p. 837. — Brauer, F., ' Monographic der (Estriden/
Wien, 1863. — Idem, " (Estr. (Hypoderma) Clarkii," in ' Ver-
handl. der zool.-bot. G-essellsch. in Wien/ xxv, p. 75. — Idem,
" On Gephenomyia trompe from the Reindeer," ibid., p. 77. —
Idem, " Hypoderma bonassi from the American Buffalo," ibid.
(all with figs.), 1875. — Breton, " On the Worm found in the
Eye of the Horse," ' Calcutta Med. and Phys. Soc. Trans./
vol. i, 1825, p. SSI.— Brown, D. 8., "The (Estrus equi, or
Horse-bot," the ' Veterinary Journal,' July, 1877, p. 14. —
Briickmuller, " Larvae in the Brain of a Foal," from ' Viert. f iir
wissensch. Vet./ in ' Veterinarian/ p. 82, 1857. — Cartwright,
W. A., " On Strangulation of the Bowels, associated with about
150 Bots, and also some 150 Ascarides, in the Horse," 'Vete-
rinarian,' p. 413, 1833. — Idem, " Case of about 200 'Bots in
the (Esophagus ' of a Horse," ibid., p. 400, 1828. — Chambron,
" On a Parasitic Malady in the Horse," from ' Ann. de Med.
Vet./ in ' Edin. Vet. Rev./ 1861 ; see also (Oambron) ' Vete-
rinarian/ 1860, p. 612. — Clark, B., " Obs. on the genus (Estrus,"
'Linn. Trans./ vol. iii, p. 289, 1797.— Idem, 'An Essay on the
Bots of Horses and other Animals,' London, 1815. — Clarksov,
N. F., " Case of Filaria oculi in the Horse," ' Vet. Rec./ vol. i,
p. 73, 1845. — Idem, " Case of Filaria medinensis in the Horse,"
ibid., 1845. — Oobbold, " On the Diptera (bots)," in a chap, on the
Parasitic Diseases of Animals, in Williams' work (1. c, Bibl.
No. 48). — Idem, "Obs. on rare Parasites from the Horse," 'Vete-
rinarian,' Feb., 1874. — Idem, "Further remarks on rare Parasites
from the Horse," ibid., April, 1874. — Idem, "Fatal Epid. affecting
Ponies," ibid., June, 1874. — Idem, "Remarks on Megnin's
Tapeworm," ibid., Sept., 1874. — Idem, " Report on Parasites
(sent from India by F. F. Collins and Spooner Hart)," ibid.,
Nov., 1874. — Idem, " Epizooty in the Horse, more especially in
relation to the Ravages produced by the Four-spined Strongyle
(S. tetracanthus)," ibid., April, 1875.— Idem, "The Egyptian
Horse Plague in relation to the question of Parasitism," ibid.,
Nov., 1876. — Idem, "Description of the new Equine Fluke
(Gastrodiscus Sonsinoii)," ibid., April, 1877. — Idem, " On

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391

Worm-like Organisms within the Mitral Valve of a Horse/'
ibid., 1877. — Idem, "Entozoa of the Horse and Elephant" (see
Bibl. No. 51).— Idem, " Observations respecting the Large-
monthed Maw-worm of the Horse/' ' Veterinarian/ Jan., 1877.
—Oouchman, T., " Worms in the Kidney of a Colt/' ' Veteri-
narian/ p. 145, 1857— Ooupe, W., "Fatal Parasitism in a
Colt/' .' Veterinarian/ Dec., ]876, p. 844. — Davaine, " Les
Cestoides," in ' Diet. Encycl. des Sci. Med./ p. 591— Dick (see
Knox) . — Dickens, 0., ' Joint-lameness in Colts, associated with
and symptomatic of Lumbricoid Worms/' ' Veterinarian/ p. 601,
1863. — Dan, B., "Remarks on Entozoa of the Horse/' 'Vete-
rinarian/ 1854, p. Uh.—Dupuy (with M. Prince), " Filaria? in
the Great Mesenteric of a Horse," ' Veterinarian/ 1835, p. 570.
— Emmerson, 0., " Prevalence of Entozoa among Horses in the
Island of Singapore," ' Veterinarian/ 1861, p. bH.—Friedberger,
F., ' Die Kolik der Pferde/ Berlin, 1874. (Eeviewed by me in
the ' Veterinarian/ Jan.— April, 1874.) — Fry, " Case of Worms
in the Horse," ' The Hippiatrist/ &c, vol. iii, p. 10, 1830.—
Gamgee, J. (senior), "On Bots," 'Edin. Vet. Rev./ July, 1858.
— Good-worth, 8., " Obstruction of the Pyloric Orifice of the
Stomach by Bots," 'Veterinarian/ p. 410, 1837. — Grellier, J.,
" On the Worm in the Eye (of the Horse)," ' Veterinarian/
p. 18, 1844.— Harlan, B., "Case of a Colt killed by Worms,"
in his 'Med. and Phys. Researches/ p. 554; see also 'Med.-
Chir. Rev./ 1836. — Harris, " A Case of Worms in the Arteries
of a Colt," 'Veterinarian/ p. 307, 1834.— Harrison, J. D., "The
singular effect of Worms in the Stomach of a Mare," 'Vete-
rinarian/ p. 331, 1842. — Hickman, T., " Worm in the Eye of
the Horse," 'Edin. Vet. Rev./ 1864, p. 653.— ffop&mso™, F.,
" Account of a Worm in the Horse's Eye," from ' Trans, of
Amer. Phil. Soc./ in 'Med. Comment./ vol. xi, p. 166, 1784. —
Hutchinson, J., " Hydatid in the Eye of a Horse," ' Path. Soc.
Trans./ and rep. in ' Lancet/ 1857. — Huxley, " On Echinococcus
(from the Zebra)," see Bibl. No. 20, o. — Jeaffreson, W., "Case
of Removal of a Worm from the Eye of an Arab Horse,"
'Lancet/ p. 690, 1836-37, and 'Veterinarian/ p. 471, 1837. —
Kennedy, M., " Account of a Nondescript Worm (Ascaris pel-
lucidus) found in the Eyes of Horses in India," ' Trans. Roy.
Soc. Edin./ vol. xi, p. 107, 1816. — Kirkman, 'Hydatids' (see
Bibl. No. 20, o). — Knox, 'Edin. Med. and Surg. Journ./ 1836.
— Krabbe (1. c, in text). — Lee, 0. A., " On Filaria papillosa in
the Anterior Chamber of the Eye of a Horse (and on Fil arias in

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PARASITES OF ANIMALS

general, &c.)," ' Amer. Journ. Sci. and Art./ vol. xxxix, p. 278,
1840.— Lessona, 0., " On the Bot (or CEstrus) of the Horse,"
from 'Recueil de Med. Vet./ in ' Veterinarian/ p. 156, 1854.
— Leuckart (1. c., in text). — Litt, W., "A Singular Case (of an
immense number of Worms in a Colt)/' ' Veterinarian/ p. 529,
1852. — Macnamara, "On F. papillosa in the Eye of Man and
the Horse," ' Indian Ann. Med. Sci./ 1864. — Marcet (see Bibl.
No. 34).— Mead, J., "A Worm in the Scrotum of a Colt,"
c Veterinarian/ 1 843, p. 648. — Megnin, " Petit Taenia inerme du
Cheval," in ' Bull, de la Soc. Cent, de Med. Vet./ t. vi, 3e serie,
p. 112. — Idem, ' Monog. de la tribu des Sarcoptides/ &c. ; see
also review by myself in the e Veterinarian/ Aug., 1877. — Mercer,
J., " On Entozoal or Worm-aneurism," ' Lond. Med. Gaz./ 1847,
and in part x of " Contrib. to Zool. Path.," in the ' Veterinarian/
p. 33, 1846. — Meyrick, J., " Death of a Colt from Entozoa within
the Abdomen," ' Veterinarian/ 1859, p. 695. — Moir, J., "Rupture
of the Ileum resulting from Worms," ' Veterinarian/ 1857,
p. 265. — Molyneux, R., " On Worm in the Eye of Horses in
India," 'Veterinarian/ 1828, p. 309. — Morgan, A., "Case of
Hydatid in the Brain of a Mare," ' Veterinarian,' p. 396, 1855.
— Numan, A., ' Ueber die Bremsen Larven, im Magen der
Pferde/ 1837. — Idem, "Entozoon (Monostoma settenii) from the
Eye of a Horse," from ' Tidschr. voor naturl. Geschied. en
Physiol./ 1842, in ' Med.-Chir. Rev.,' 1842.— Idem (for remarks
on Oysticercus fistularis), ' Over den veelkop blaasworm' (1. c,
Bibl. No. 49), p. 263.— Peall, T., " A Discourse on ' Worms/ "
at p. 37, in his ' Observations, chiefly practical, on some of the
more common Diseases of the Horse/ pub. at Cork, 1814. —
Percivall, 0., "Worm in the Eye of the Horse (two cases),"
' Veterinarian/ p. 75, 1828. — Percivall, J., "A Case of Ascarides
in the large Intestines of the Horse," ' Veterinarian/ p. 858,
1829. — Poulton, T. J., " Large numbers of Parasites in the
Intestines of a Mare," ' Veterinarian/ 1866, p. 385. — Seaman,
J., " Worms in the Blood-vessels of Horses and Colts," ' Edin.
Vet. Rev./ 1864, p. 520. — Simonds, "On Disease of the
Mesenteric Artery, produced by Strongyli," ' Path. Soc. Trans.,'
1854. — Slceavington, O., " On Worm in the Eye of the Horse
(three cases)," ' Veterinarian/ 1834, p. 196. — Sonsino, P., " On
the Entozoa of the Horse in relation to the late Egyptian
Equine Plague," ' Veterinarian/ Feb. and March, 1877. — Tyndal,
J., " Worms in the Intestines of a Mare," ' Veterinarian/ 1843,
p. 629. — Twining, W., " Obs. on the Filaria or Threadworm

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393

found in the Eyes of Horses in India/' c Calcutta Med. and
Phys. Soc. Trans./ vol. i, p. 345, 1825 ; rep. in ' Veterinarian/
p. 114, 1828. — Valenciennes, " On the Spiroptera mecjastoma of
Gurlt," abstract of a paper from Acad. Sci. of Paris, reported
in ( Lancet/ 1843. — Varnell, " Remarks on Oases of Parasitic
Disease in Horses," 'Veterinarian/ p. 201, 1864. — Veret, "Per-
foration of the small Intestines by Ascarides lumbricoides (in
the Horse)," from ' Rec. de Med. Vet./ in ' Veterinarian/
p. 569, 1837. — Vincent, "Curious case of Incurable Lameness
from Hydatids," ' Veterinarian/ 1848, p. 674 ; see also p. 3,
Hid, — Wallis, " Note on the occurrence of (250) Lumbrici in a
Horse," ' Veterinary Record/ 1849, p. 300.— Walters, B. 0.,.
"Parasites in the Kidneys of a Mare," ' Veterinarian/ 1866,
p. 265. — Woodger, "Hydatid in the Brain of a Horse," ' Vete-
rinarian/ 1863, p. 75. — Woods, J. S., " Tetanus in a Mare,
associated with the Larvse of (Estrus equi within the Stomach and
Duodenum," ( Veterinarian/ 1859, p. 693. — Wright, " Strongyli
in the Scrotum of a Colt," 'Veterinary Record/ 1849, p. 385.
— Yonatt, " Worms between the Tunics of the Stomach," ' Vete-
rinarian,' 1835, p. 571. — Idem, " Tetanus, Worms in the Trachea,
and Dilatation of the Heart, in a Zebra," ibid., p. 504, 1836. —
— Idem, " Worms in the Nasal Cavity of the Horse," ibid.,
p. 329, 1832. — Zangger, "Remarks on Entozoa/' from the
French, ' Veterinarian/ 1855, p. 463.

Part X (Pachydermata)

Concerning the parasites and parasitic diseases of this mis-
cellaneous assemblage of large mammals, I shall first speak of
those of the proboscideans (Elephantidge) . Except by myself,
they have been but little studied, and I am yet waiting for an
opportunity to give further time to their consideration. When
Diesing published his ' Systema ' only two helminths were
referred to the Indian elephant, namely, Ascaris lonchoptera and
an undescribed fluke supposed to be a distome. The whole
subject requires revision, but I think the following species must,
for the present at least, be allowed recognition : — Fas-
ciola Jacksoni (mihi), Amphistoma Ilaivlcesii (mihi), Ascaris lon-
choptera (Diesing), Sclerostoma spinuliferum (Baird), and
Dochmius Sangeri (mihi). Either the Ascaris or the Sclerostoma
is probably identical with Rudolphi's Strongylus elephantis.

At the Norwich Meeting of the British Association, in 1868,

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PARASITES OF ANIMALS

I exhibited two flukes received from J. S. Thacker, V.S., of the
Madras Army. They were handed to me by the late Dr Baird,
and were labelled " Distoma taken from liver of elephant and
forwarded for classification. " I stated at the time that these
entozoa were identical with certain flukes previously obtained
from the duodenum and biliary ducts of an Indian elephant,
and which, though carefully preserved in the Boston Museum,
U.S., had never been properly described. They were only
briefly noticed by Dr Jackson in his ' Descriptive Catalogue ' of
the Museum. In the summer of 1868 fifteen specimens of fluke,
removed from Burmese elephants, had been forwarded to and
received by Professor Huxley from Rangoon, accompanied by a
statement to the effect that they were the cause of an extensive
and fatal disease in Burmah. Through the kindness of Prof.
Huxley I was allowed to make use of his specimens for the
purpose of comparison and identification, and thus it became
evident that our specimens were of the same species. It was
also evident that the species could be none other than that
represented by the Boston specimens. Further examination
having made it clear that the organisation of these flukes departed
from the ordinary distome type, I named the parasite Fasciola
Jacksoni, at the same time offering the following description
(' Entozoa/ Supp., 1869, p. 80) : — Body armed throughout with
minute spines, orbicular, usually folded at either end towards
the ventral aspect, thus presenting a concavo-convex form ; oral
sucker terminal, with reproductive papillae about midway between
it and the ventral acetabulum ; intromittent organ \" in length ;
digestive apparatus with two main zigzag-shaped canals, giving
off alternating branches at the angles thus formed, the ultimate
ceecal ramifications occupying the whole extent of the body ;
length, when unrolled, from \" to f", breadth J" to
Now, if reference be made to the appendix of the late C. M.
Diesing's ' Systema Helminthum/ it will be found that Jackson's
statement had not escaped that helminthologist's notice, though,
not having seen any specimens, he was not unnaturally led to
place the species amongst the distomes proper. In Diesing's
subsequently published ' Eevision der Myzelminthen/ the species
is formally characterised as the Distomum elephantis of Jackson
(f Sitzungsberichte d.Math.-nat. Cl.d.k. Akad. d. Wissenchaftcn/
Bd. xxxii, 1858). In my " Synopsis of the Distoinidre," which
appeared in the ' Journal of the Linnean Society' for 1861, I
had also placed it amongst the distomes, not considering it

PACHYDERMATA

395

to be a doubtful form ('Proceed. Linn. Soc./ " Zoology,"
vol. v, p. 9). These references exhausted the literature of the
subject up to the time of the issue of my 'Manual' in 1873,
where this fluke is again briefly noticed (p. 13). Several of
Prof. Huxley's specimens have been added to the entozoological
department of the Hunterian Museum. It is clear that all
these notices and descriptions point to the same parasite. The
worm has since been more carefully described by Dr R. H. Fitz,
from a series of dissections and preparations made by Dr H. P.
Quincy, and deposited in the Warren Museum, Boston, U.S.

About the middle of June, 1875, I received a letter from
General Hawkes, of the Madras Staff Corps, dated Secunderabad,
May 12th, 1875, and in reference to the subject before us he
writes as follows : — " My attention has been recently directed
to a very unusual mortality of elephants at this station. Out
of twenty-eight elephants under my charge, no less than twelve
have died within the last sixteen months, whereas the average
annual mortality has been hitherto only two per annum out of
thirty-eight in our establishment. In every case of death there
appeared to exist serious organic disease quite sufficient to
account for such death, but as the mortality increased I had a
post-mortem examination made in each case ; and although
here also organic disease sufficient to account for death was
present in each case, yet in every one of these elephants we
found the liver-fluke in greater or less abundance." General
Hawkes adds : — " Meanwhile I have sent you a small box con-
taining three bottles, one containing the liver-fluke (Fasciola
Jachsoni) referred to in your work on the parasites of domesti-
cated animals. It seems possible that the other two species of
parasites may not have been brought to your notice. Both of
these, namely, the " masuri " and the " soorti," are very common
in elephants. They are both found in the intestines only. The
" masuri," when present in any quantity, cause considerable
disturbance, and the animal instinctively resorts to the eating
of earth, which it consumes in large quantities until the bowels
are acted on and the worm expelled. The soorti is more
common than masuri, and does not seem to inconvenience the
animal very much. When expelled from the animal the soorti
is a round white worm, like most of the threadworms ; the
masuri, on the other hand, is of a delicate flesh color."
Shortly after the receipt of this letter I obtained the entozoa in
a good state of preservation. Accordingly I wrote to General

396

PARASITES OP ANIMALS

Hawkes, stating that the flukes were clearly referable to Fasciola
Jacksoni; that the parasites to which the natives of Hindostan
apply the term u soorti " were evidently examples of Ascaris
lonchoptera (Diesing), previously called strongyles by Rudolphi ;
and that the worms which he called " masuri " were trematodes
new to science. I named the species Amphistoma Hawkesii, in
honor of the donor. The bottle contained as many as forty-nine
specimens. I may here remark that I have made inquiries of
the keepers of the elephants at the Zoological Gardens as to
whether they have ever seen entozoa that were passed by the
animals under their care. They replied in the negative, the
keeper of the African elephants (Scott) having made frequent
inspection of the faeces. I was the more anxious to secure
information on this point since, during my frequent visits to the
menagerie, I had observed that the African elephants were in
the habit of swallowing large quantities of mud and dirt from
small hollows in the ground near the great water-tanks in which
they bathe. Prof. Garrod (who had dissected three elephants)
also assures me that there has been no trace of an entozoon in
any of the Indian elephants examined by him. In one dissected
at Edinburgh the same negative result was obtained. From
the facts at present in my possession, I conclude that the habit
of earth-eating, displayed alike by Indian and African elephants
(and, as stated in my account of the equine parasites, shared by
horses), is not necessarily due to the presence of parasites. I
apprehend rather, that it is resorted to by these animals under
any circumstances of intestinal irritation, whether created by
entozoa or other foreign agents. The notion of the elephant's
intelligent self- cure by eating earth is a very old fable. Captain
Forsyth, as quoted by Mr Fleming, alludes to it in his ' High-
lands of Central India/ and I find the same ideas recorded
by Williamson and Howitt. Forsyth says : — " Elephants are
very liable to intestinal worms. They generally cure themselves
by swallowing from ten to twenty pounds of earth." Captain
Williamson says : — " They are much troubled with worms, for
the cure of which the elephant eats earth. If the dung be
inspected there will be seen an amazing number of moviug
objects, which much resemble pieces of chewed sugar-cane."
Some excellent practical remarks are added, testifying to the
value of the native remedy called Kallah-nimok, or bit-noben,
which is a saline purgative. In Lieut. Ouchterlony's essay
(quoted below) no allusion is made to the subject of worms.

PACHY DEEM ATA

397

General Hawkes afterwards supplied me with further infor-
mation. In a letter from Secunderabad, dated July 30th, 1875,
he says: — "As regards the liver-fluke (F. Jacksoni), it appears
from your treatise to have been first observed in 1847. The
only other published notice that I have been able to find of it
is contained in a letter to a newspaper, dated c Kan goon, 16th
July, 1867/ and is signed 'K. B/ In this letter the unusual
mortality of seven elephants in about fifteen days is attributed
to the presence of this liver-fluke, the two other parasites
(Amphistoma and Ascaris lonchoptera) being also present in the
intestines." " Now (continues General Hawkes), in every case
at which I was present flukes were found in greater or less
numbers in the gall-ducts of the liver, and the Amphistoma was
also as constantly present in the intestines, the soorti (Ascaris
lonchoptera), contrary to the general experience of the elephant
attendants, being less frequently met with, though from its
color and slender shape it is not so easily detected among the
huge mass of faeces as the larger Amphistoma." Speaking of
the amphistoma General Hawkes says: — "This internal parasite
is well known to all who possess elephants. It is alluded to by
Dr Gilchrist in his treatise on the ' Diseases of Elephants/ first
published in 1841, but he merely mentioned it under its local
name, masuri, and made no attempt either to describe it scien-
tifically or to ascertain its place in the natural system. As far
as my experience goes it is only found in the intestines. These
parasites appear to be very generally present in the elephant.
When their numbers are few the ' host ' is probably not much
inconvenienced, but when present in any great quantity they
undoubtedly cause much irritation. When this is felt, the
animal, as before remarked, instinctively resorts to a simple and
effectual remedy. He eats a quantity of earth, which purges
him thoroughly and expels the amphistoma. The mahawats
are of opinion that whilst the elephant is eating earth to relieve
himself of the pests the daily allowance of rice should be
scrupulously withheld ; and they say that if the rice, which is
given uncooked, is eaten by the animal under these circum-
stances, excessive purgation is induced, which frequently results
in death. How far this opinion is founded on fact I am unable
to say, but the mahawat's name for this disease means ' fast-
ing/ and bears testimony to the generally received notion of
the necessity of withholding the rice when the animal is eating
earth."

398

PAIUSITES OF ANIMALS

When describing the parasites of the horse (p. 358), I spoke
of Collins' amphistome from that animal, but in the letter
addressed to me from Simla, 22nd March, 1875, Mr. Collins made
no allusion to the earth-eating habit. He wrote : — " I forward
you by this mail parasites found in the colon of a horse that
died, a subject of fever peculiar to this country. There were about
a thousand of the parasites, and nearly the whole of them were
situated close to the csecum, and were loose in the gut. Not
having seen parasites at all similar to these, I have forwarded
them for identification. They were of a brick-red color when
first obtained." These explicit statements by Mr Collins are
interesting from many points of view. One has only to place
his specimens side by side with those from the elephant in
order to satisfy one's self that the two forms are distinct. For
the reasons already stated I provisionally called the worm
Amphistoma (Jollinsii. It is probable that other veterinary
surgeons have encountered this entozoon in India ; but, unless
they can point to some published account of the fact, Mr Col-
lins is entitled to be considered as its discoverer. Doubtless
many other European residents in India, Ceylon, and Burmah,
must, like Dr Gilchrist, be well acquainted with the rnasuri as
such, though unaware of their zoological position.

In a record of the post-mortem examination of one of the
victims of the Secunderabad epizooty, the veterinary surgeon
said : — " No doubt disease of the lungs and subacute inflamma-
tion of the bowels were the immediate cause of death, but the
large number of flukes in the liver and the intestinal parasites
(i. e. the amphistomes) account in a great measure for some of
the symptoms shown, and these symptoms accord in many
respects with those shown in elephants that died in Burmah
during the epizooty (rot) in 1867, as recorded by R. B., no-
tably, refusal of food, standing with mouth open, restlessness,
and puffiness about the head and shoulders. The liver pai*a-
site is no doubt the same referred to by E. B., and is that
termed by Dr Cobbold Fasciola Jacksoni" In reference to a
later case the same officer remarks : — " I carried out the post-
mortem examination with special reference to inquiry as to the
probability of the mortality amongst elephants at this station
being of parasitic origin. This was suggested to me by the
former case. The post-mortem appearances differed in every
respect. There were flukes in the liver, but in no great
quantity, and the structure of the liver was sound. Although

PACHYDERM AT A.

399

not assisted by this case in attributing the mortality to para-
sitic origin, I am strengthened in my opinion that the death of
the previous elephant was due to disease caused by the presence
of the liver fluke." This report, by Mr W. S. Adams, is
to some extent in harmony with later information. An epi-
zootic outbreak amongst elephants has occurred in England, at
Sanger's Circus, and I had opportunity to examine one of the
dead animals. In my own opinion, and in that of Mr F. Smith,
the veterinary surgeon who attended the animals professionally,
the disease was due to parasites. I obtained large quantities
of Amphistoma Hawhesii from the intestinal canal, and also
other worms. The death of one of the elephants was made the
subject of litigation, when, as might be expected, great diver-
sity of opinion as to the cause of the fatal issue prevailed.

Mr Smith, an old pupil of mine, regarding the amphistomes
and strongyles as the cause of death, wrote to the effect that
" some of the worms were found between the coats of the
intestine, and others on the free surface of the gut, whilst the
excretory ducts of some of the glands were found blocked with
them." The animal examined by myself on the 24th of August,
1876, yielded numerous examples of Amphistoma Hawkesii,
Ascaris lonehoptera, and Dochmius Sangeri, the last species
being so named by me after the owner of the circus who lost
the herd of elephants by the epizooty. The male Dochmii
measured f and the females | of an inch in length. Here I
must reluctantly quit the helminths of elephants, adding only an
expression of surprise that Dr Max Schmidt should have had so
little to say concerning them in his otherwise instructive memoir
on 'The Diseases of Pachyderms ' (quoted below).

I have but a few words to offer respecting the ectozoa. A
species of mite has been described whose generic position
appears doubtful. I allude to Homopus elephantis of Fiirsten-
berg, or Symbiotes elephantis of Gerlach. According to Megnin
it is a nywphe adventive or hypope of a variety of Tyroglyphus
siro. This acarus is abundant in old forage. Another ectozoon
is Hcem atomy zus elephantis. It differs from the lice proper in
many respects, but, according to Piaget, the reproductive organs
resemble those of Hamatopinus. In ' Science Gossip 1 for June,
1871, Mr H. C. Richter d escribes t{ a new form of parasite,"
which is called Idolocoris elephantis. The insect, which was
one line in length, was found upon an elephant in Ceylon.
According to Walker it not only constituted the type of a new

400

PARASITES OF ANIMALS

genus, but of an altogether new family of the Hemiptera Hete-
roptera, coming very near to the bed-bugs (Acanthidce) . It is a
huge sucking louse. From the discussion which followed, it
seems that the parasite had several times been seen before, and
was none other than E. Piaget's Eamatomyzus elephantis.
Excellent figures accompany Richter's and Piaget's descrip-
tions. Notwithstanding Piaget's explanation, I think the
specific name, longirostris, would have been a more appropriate
appellation.

Bibliography (No. 51). — (Anonymous), "Diseased Elephants/''
see ' Lancet/ Sept. 2, 1876 ; also " Report of the Case at Law
(Jamrach v. Sanger)," given in the ' Veterinarian/ Dec, 1877,
p. 886. — Oobbold, T.S., " Description of a species of Trematode
from the Indian Elephant, with remarks on its Affinities,"
'Quart. Micros. Journ./ Jan., 1869 ; see also ' Entozoa/ supp.,
1869, p. 80. — Idem, " On the Destruction of Elephants by
Parasites, with remarks on two new species of Entozoa and on
the so-called Earth-eating habits of Elephants and Horses in
India," ' Veterinarian/ Oct., 1875. — Idem, "Further Remarks
on Parasites from the Horse and Elephant, with a notice of new
Amphistomes from the Ox/' ibid., Nov., 1875. — Diesing (1. c,
in text). — Fitz, B. H., "Anatomy of Fasciola Jacksoni," ' Rep.
of Boston Soc. Med. Sci./ in the 'New York Med. Journ./
Nov., 1876. — Fleming, G., "The Diseases of Elephants" (chiefly
from Captain Forsyth's work on the ' Highlands of Central
India'), 'Veterinarian/ March, 1873, p. 181— Hegnin, "Mem.
sur les Hypopes," in Robin's 'Journ. de l'Anat. et de la
Physiol./ 1874 (H. elephantis), p. 248. — Ouchterlony , J. W.,
"An Essay on the Management of the Elephant, and its Treat-
ment in ordinary Diseases," ' Rep. of Vet. Med. Assoc./ Nov.,
1872, and pub. in 'Veterinarian/ Jan., 1873, p. 65. — Piaget, E.,
" Description d'un parasite de l'elephant," ' Tijschrift voor
Entomologie/ 1869, p. 249. — Bichter, E. C, " A new form of
Parasite (Idolocoris elephantis)," ' Science Gossip/ 1871, pp. 131,
185, 211, 278. — Schmidt, Max, "Die Krankheiten der Dick-
hauter," ' Deutsche Zeitschrift f. Thiermed. und vergleichende
Pathologie/ f. Nov., 1878, s. 360.— Williamson, T., 'Oriental
Field Sports/ London, 1807, vol. i, p. 138.

The parasites of the Bhinoceridce have been even less studied
than those of elephants. In 1856 Prof. Peters described a
tapeworm from Bruce's rhinoceros (B. Africanus), which he
named Taenia gigantea. In 1870 Dr Murie, under the provi-

PAGHYDERMATA

401

sional name of T. magna, published a description of the strobile
of the same cestode from an Indian rhinoceros (B. unicornis).
From a total misconception of the character of the proglottides,
Murie was led to suppose that the segments of the strobile were
very deep as well as broad; whereas the proglottids are
remarkably narrow, thus partaking of the characters of the
Taeniae of the larger herbivora in general. In a subsequent
paper Peters pointed out these errors. Murie had, in fact,
rolled several segments into one. In 1877 Professor Garrod
encountered the same cestode in Rhinoceros sondaicus, and,
following Peters' example, separated it from the Taeniae proper
{Plagiotcenia gigantea). The idea of generically separating
tapeworms possessing a more or less striking breadth of strobile
is not one which commends itself to my view, seeing that many
of the tapeworms of herbivora closely resemble the rhinoscerine
cestodes in this respect. As Diesing hints, this tapeworm
comes near to T. perfoliata, but Garrod' s and Peters' figures
both show that Plagiotcenia wants the neck-lobes. The pre-
sence of cephalic appendages may be regarded as generically
distinctive, but it does not appear that Blanchard separated the
perfoliate tapeworm of the horse from the Taeniae proper on this
ground. Therefore, in my account of the equine tapeworms, I
have not adopted his genus Anoplocephala. I may remark, in
passing, that if the distinctions, as between armed and un-
armed, or between proboscis-bearing (Rhynchotceniada) and
non-proboscis-bearing tapeworms (Arynchotceniada) , are to be
maintained, they should be expressive of divisional or sub-
ordinate value. Dr Weinland's arrangement, having reference
to the thick- and thin-shelled ova (Sclero- and Malaco-leptidota),
is, perhaps, preferable. The whole subject of classification
requires revision, but it should be undertaken by some helmin-
thologists practically acquainted with a large number of cestode
types. As Garrod has well observed, PJagiotaenia enjoys a
wide geographical distribution, infesting alike Indian and
African hosts. Prof. Garrod, I observe, speaks of the head of
the mature tapeworm as the scolex — an extension of the mean-
ing of a term not usually recognised. In this, however, he
only follows Peters' unfortunate example.

The wide distribution enjoyed by Peters' Plagiota3nia is
probably equalled by that of the rhinocerine stomach-bot
{Gastrophikis rhmocerontis, Owen). This parasite was origin-
ally described in 1840, and since that time it has been fro-

2(3

402

PARASITES OF ANIMALS

quently encountered both in India and Africa. To Mr Spooner
Hart, of Calcutta, I am indebted for a large number of speci-
mens; their size exceeding that of any other bots that have
come under my notice. Probably this parasite infests the
stomach of rhinoceroses generally; at all events, it occurs in
B. unicornis, B. bicornis, and B. simus. At present the imago
is unknown. The longest larvEe in my possession measure 1£",
but Brauer records specimens up to 35 mm. in length by 10
mm. in thickness. In African hosts M. Delegorgue found
these parasites in prodigious numbers.

Bibliography (No. 52). — Brauer, " Bot of the Rhinoceros,"
'Monogr. der CEstr./ 1863, s. 92.— Oobbold, "Note on Para-
sites presented by Messrs Danford, Hart, and others/' 'Vete-
rinarian/ 1875, p. 513. — Ooquerel and Salle, in ' Ann. Sqc
Eutom. de France/ 1862 (quoted by Brauer). — Delegorgue,
' Voyage dans FAfrique ' (quoted by Brauer). — Garrod, "On
the Taenia of the Rhinoceros of the Sunderbunds (Plag. gig.,
Peters)/' < Proc. Zool. Soc./ Nov. 20, 1877, p. 788.— Hope, 'in
' Trans. Entom. Soc./ 1840, p. 259.—Joly, M. N., " Recherches
Zool. (&c.) sur les (Estrides (&c.)/J in 'Ann. des Sciences (&c.)
de Lyon/ 1846 (quoted by Brauer). — Marie, J., " On a probably
new species of Taenia (T. magna?) from the Rhinoceros/'' 'Proc.
Zool. Soc./ 1870, p. 608.— Peters, W., "Note on the Taenia
from the Rhinoceros, lately described by Dr J. Murie," ' Proc.
Zool. Soc./ 1871, p. 146. '

Very little has been written respecting the parasites of the
Hippopotamida and Tapiridce. I think it was Livingstone who
first di'ew attention to the fact that the river-horse or sea-cow
is much infested by tapeworms, but I have not seen any
published description of the worm. Dr. Murie, during his
sojourn in Egypt, found a solitary bot embedded in the soft parts
surrounding the eye, and judging from his figure the species is
new to science. Provisionally I speak of it as the Hypoderma
Muriei. In the paper (quoted below) Murie appends a list of
all the animals in which bots have been found. Though chiefly
taken from Brauer, it is useful and tolerably complete. So far
as I am aware no cestodes have been desci'ibed as infesting
tapirs ; nevertheless, at least five other kinds of helminth have
been found in Tapirus Americanus. Of these, two are flukes
(Amphistoma asperum and A. pyriforme), and three are nema-
todes (Sclerostoma monostechum, Spiroptera mediospiralis, and
S'p. chrisoptera). The three species first named occupy the caecum,

PACHYDERM ATA

403

whilst the others are found in the stomach. According to
Molin' s description, both species occupy tuberous excrescences
of the mucous membrane, thus reminding us of the similar
habit enjoyed by Sp. megastoma in the horse. The Sp. chry-
soptera is a comparatively large species, the males measuring
an inch, and the females as much as an inch and a half in
length. Both of the spiropteras were obtained from tapirs by
the indefatigable Natterer, Sp. mediospiralis being also pro-
cured by him from the aguti. If I have read Molin correctly,
as many as thirty-four examples of S. mediospiralis were taken
from a single excrescence in the stomach of the tapir. Up-
wards of a hundred specimens were procured, collectively, from
three similar stomach-excrescences in Dasyprocta aguti. These,
and the other tapirine parasites above mentioned, were originally
discovered in Brazil.

Bibliography (No. 53). — Diesing, " Neue Gattungen Binnen-
wiirmern nebst enem Nachtrage zur Monographie der Amphis-
tomen," in ' Annalen d. Wien. Museums/ Feb., 1839, s. 236. —
Idem, ' Systema/ Bd. ii, s. 306. — Molin, " Una monografia del
genere Spiroptera," in c Sitzungsb. der math.-naturw. CI. d. k.
Akad. d. Wissensch./ Bd. xxxviii, s. 1001, 1859. — Murie, "On
a larval CEstrus found in the Hippopotamus," ' Proc. Zool. Soc./
1870, p. 78.

The osculant position of the anisodactyle pachyderms (Hy-
racidae), formerly classed as rodents, renders it desirable that
their parasites should be briefly noticed in this place. Pro-
bably these animals, zoologically speaking, come nearest to
the rhinoceroses, but Prof. Owen showed that, anatomically,
they possessed marked affinities with the sloths. The klipdas
or dasse (Hyrax capensis) is infested by a tapeworm, of which
hitherto the proglottides only appear to have been seen (Taenia
hyracis, Pallas). Under the name of Ccenurus serialis a larval
cestode has been described by Gervais, the same parasite being
called Arynchotania critica by Pagenstecher (" Zur Naturges-
chichte der Cestoden," in 'Sieb. u. K611. Zeitschrift A variety
of nematodes have also been observed in the Cape hyrax. Of
these, the so-called Physaloptera spirula is classed as doubtful
by Molin and Diesing. Hemprich and Bhrenberg furnished
brief descriptions of four other nematodes. Two of these
worms were placed in the genus Oxyuris (0. pugio and 0.
flavellum) , and the other two in the new genus Crossophorus,
which they formed for their reception (0. collaris and G.

404

PARASITES OF ANIMALS

tentaculatus) . The whole of these nematoids were obtained
either from the caocum or large intestine.

An able article in the ' Natural History Eeview ' for July,
1865, attributed to Professor Huxley, expressed very clearly the
popular notion as to the great danger of the flesh of swine
considered as a source of human parasites. No doubt the
filthy pachyderms in question (Suidce) are much infested by
helminths, some of which gain access to man, but swine are
neither attacked by a greater variety of entozoa than other
domesticated animals, nor are they so frequently a source of
human tapeworms as cattle. In the article above quoted the
following passage occurs : — " Of all animals, feral or domestic,
the common pig is beyond all doubt the most fertile source of
human entozoa ; at least, of important parasites, Trichina sjn-
ralis and the tapeworm would, there is good reason to believe,
cease to infest us, did not this favorite quadruped act the part
of a communicating medium." This paragraph was evidently
written under the impression that "the tapeworm" most
commonly found in man was derived from the hog. So far
back as 1864 I showed that this was an entire mistake.

Flukes are rare in swine; nevertheless, Fasciola hepatica
and Distoma lanceolatum are occasionally present in the
domestic hog, and the peccaries (Dicotyles) are infested by an
Amphistome (A. giganteum). This large species, §" in length,
formed the basis of an admirable account of the anatomy of
this genus of worms which the learned Vienna helminthologist,
Diesing, wrote before he was deprived of his eyesight. The
merits of that respected systematise s investigations have, I
think, been much underrated, in consequence, no doubt, of the
artificial character of his system of classification. For all that,
his writings remain invaluable. Turning to the cestodes of
swine, there is not, so far as I am aware, any evidence of the
occurrence of sexually-mature tapeworms either in the hog or
its allies; but the frequency of larval cestodes, known as
measles (Oysticercus tela cellulose?), was well known to the early
Jewish writers. In the first part of this work I devoted as
much space as I could spare to the consideration of Cysticerci
in general, and the pork-measle in particular ; but an exhaus-
tive knowledge of the subject in relation to hygiene can only
be acquired by consulting the principal original memoirs (quoted
in the Bibliographies Nos. 13 and 14). In a Westphalian ham,
part of which was sent to me for examination, I calculated that

r A C H Y DE RM AT A

405

each pound of the flesh must have contained upwards of 600
Cysticerci. I was informed by the donor, Dr Prior, that in
spite of the disgusting state of the meat much of it had been
eaten by the well-to-do family who purchased the ham. Cysti-
cerci occasionally occupy the brain of the pig in considerable
numbers. Florman recorded a case of this kind where their
presence gave rise to vertigo in all respects resembling the gid
ordinarily produced by Coenurus in the sheep. As regards the
larger cestode larvae, Cysticercus tenuicollis and Echinococcus
veterinorum are of frequent occurrence. One not unfrequently
encounters the former in the mesentery, whilst the liver of the
hog is sometimes so crowded with hydatids that scarcely any of
the glandular substance of the organ remains visible. It is
surprising how little the infested bearers appear to be incon-
venienced in such cases. In the winter of 1859, and in the
autumn of 1860, I found large cystic entozoa in an African Wart-
hog and in a Red River hog.
These animals had died at the
London Zoological Society's
Menagerie ; and as the worms
appeared to me at the time to
be quite distinct from the ordi-
nary slender-necked hydatid,
they were named, respectively,
Cysticercus phacochari cethiopici
and C. potamochceri penicillati.
The solitary example from the
wart-hog was found in a cyst
near the colon ; whilst of the five
large bladder- worms obtained
from the Red River hog, one in-
fested the liver and the other ,
four were lodged in the folds JiRlver hog- Masniiiei1 60 diameters, ori-
of the mesentery. The caudal

vesicle of the worm from the wart-hog measured 3£" in
diameter, the vesicle of the other bladder-worm being much
longer. A reference to the original figures will show that
these forms are distinct. Swine are largely infested by nema-
todes. The best-known form is Ascaris lumbricoides, which
Dujardin regarded as distinct (A. suilla). The hitherto dis-
puted identity of this worm with the human lumbricoid being
no longer questionable, the importance of the entozoon in rela-

illlllHli

iiliWwff/

iiillli

406

PARASITES OP ANIMALS

tion to luinbricoid endemics must at once be obvious ; I have
already, however, dwelt upon this subject when treating of the
human parasites. In like manner, the subject of the flesh-
worm disease, which is due to Trichina spiralis, cannot be
discussed in this place, as I have fully entered upon it in
connection with trichinosis in the human subject. What may
be the nature of the small threadworms found by Leidy in the
extensor muscles of the hog I cannot say, but Diesing inferred
that they might represent a distinct species {Trichina affinis).
As regards the allied genus Trichocephalus, the common species
infesting swine (T. crenatus), appears to be rarely absent. It
not only infests the common domestic and wild hog, but the
peccaries and wart-hogs. These entozoa are probably harmless
to their bearers. In reference to them Krabbe says : — " When
the eggs are expelled with the excrement and pass into water,
then the embryos, after several months' furlough, and there
undergoing further development, are transferred to the swine's
intestinal canal/' If I rightly understand the paragraph
(' Husdyrenes Indvoldsorme,' p. 28), Krabbe states that the
embryos are still within their egg-coverings when infection
takes place. The maw-worm of the hog is known as Spiroptera
strongylina. It was described and figured by Grurlt. The
males measure \" and the females §" in length. Specimens
of this worm were supposed to have been found by Natterer in
Dicotyles albirostris ; but it seems that the worms in question
represent a distinct species, if not an altogether new genus.
In the year 1864 Professor Simonds placed in my hands a
very singular nematode, to which I gave the binomial term
Simondsia paradoxa. Numerous examples of this worm were
found by Prof. Simonds occupying cysts within the walls of
the stomach of a hog which had died at the London Zoological
Society's Menagerie. In my introductory treatise I wrote of
it as follows : — " The worm in question has been regarded by
Mr Simonds as a species of Strongylus, but I am inclined to
think that its affinities will place it nearer to the genus Spire-
ptera. At present I have only examined the female, which is
characterised by the possession of a multitude of large tentacle-
like appendages surrounding the neck. These processes, by
their aspect, remind one of the so-called branchial projections
on the back of Eolis, but in this worm I believe them to be
special folds formed for the lodgment of unusually developed
uterine organs. The female worm is about §" in length."

l'ACIIYDERMATA

407

In the interval that has elapsed I have been unable to supply-
further particulars, and unfortunately the original drawings of
the worm have been lost. The habits of the parasite remind
us of Spiroptera megastoma infesting the walls of the stomach
of the horse. Not improbably this singular entozoon may turn
out to be identical with Molin' s Spiroptera sexalata, and if so,
it may correspond with Spiroptera strongylina. However,
Diesing afterwards recognising, as I had done, the desirability
of separating this last-named worm from the Spiropterse proper,
formed for it his new genus Physocephalus. He then called the
worm Physocephalus sexalatus. If, as is probable, my Simondsia
and Diesing's Physocephalus are identical, the species found by
Simonds ought to be recognised by the generic title which
Diesing proposed. His genus was established about four years
before I described my Simondsia. Diesing was evidently led
up to the recognition of the generic distinction of the worm by
Molin' s examination and description of the worm. As, in my
original account of the worm found by Simonds, I spoke of
numerous appendages to the neck, it is evident that further
investigation is necessary to clear up the question of identity.
According to Molin and Diesing the male Spiroptera sexalata
measures rather beyond \" and the female beyond \" in length.
Neither Diesing nor Molin speak of Natterer's worms as being
found encysted. In fact they were free. Molin simply re-
marks : — " Io ne esaminai in oltre 6 esemplari maschi e 77
femine raccolti in parte dal muco che revestiva le pareti dello
stomaco, ed in parte dal pasto contenuto nello stesso organo di
un Dicotyles albirostris femina ai 24 Aprile, 1826." After all
that has been said it may be that my Simondsia paradoxa
and Diesing' s Physocephalus sexalata are quite distinct, and that
like the large- and small-mouthed maw-worms of the horse
(Spiroptera megastoma and S. microstoma) they play a corre-
sponding role. Before very long I hope to set this question
definitively at rest.

Passing to the strongyloid nematodes one of the most remark-
able and important species is Stephanurus dentatus. In the
' Annalen des Wiener Museums ' for 1839 (s. 232) this worm was
first described by Diesing, who employed the generic title as
expressive of the crown-like figure of the tail of the male worm.
Diesing wrote as follows : — " At Barra do Eio Negro, on the
24th of March, 1834, Natterer discovered this peculiar genus of
worms occurring singly or several together in capsules situated

408

PARASITES OF ANIMALS

amongst the layers of fat in a Chinese race of 8us scrofa
domestica. The males measure from ten to thirteen lines long,
the females from fifteen to eighteen lines, the former being
scarcely a line in breadth at the middle of the body, whilst the
latter are almost a line and a half in thickness. The curved
body thickens towards the tail, is transversely annulated, and
viewed with a penetrating lens is seen to be furnished with
integumentary pores. The oral aperture opens widely. It is
almost circular, and is supplied with six teeth at the margin.
Two of these standing opposed to one another are larger and
stronger than the rest. The tail of the male, when spread out
evenly, is surrounded by a coronet of five lancet-shaped flaps ;
the combined flaps being connected together from base to apex
by means of a delicate transparent membrane. The single
spiculum situated at the extreme end of the tail projects slightly
forward and is surrounded by three skittle-shaped bodies.
The tail of the female is curved upon itself, rounded off, and
drawn out at the extreme end into a straight beak-shaped
point; whilst to both sides of the stumpy caudal extremity of
the body short vesicular prominences are attached. The female
reproductive outlet occurs at the commencement of the second
half of the body. Thus, judging by its external characters this
genus is most closely allied to Strongylus." In reproducing
Diesing's description I have here rendered the translation
somewhat more freely than in my previous record of the dis-
covery given in ' Nature ' (1871). The original description is
supplemented by a brief account of the internal anatomy of
the worm.

So far as I am aware no subsequent notice of this entozoon
appeared until the year 1858, when Dr J. C. White gave some
account of a " find " made in the United States. This re-dis-
covery was reported in the sixth volume of the ' Proceedings of
the Boston Natural History Society/ Dr White says : — " The
worms were found in the leaf -yard of an apparently healthy hog,
in the adipose tissue near the kidney. They occupied a space of
the same about the size of a man's fist and had burrowed through
the mass in every direction, forming canals three or four milli-
metres in diameter, which terminated in cysts. On cutting open
these cavities, which did not communicate with each other, they
were found filled with pus, and in each were two worms, male
and female.-" Dr White expresses his opinion that the worms
gained access to the tissues " by boring through the circulatory

PACHYDNiniATA

409

system while in the embryonic condition." I think that Dr
White deserves great credit for his correct diagnosis of the
species, and all the more so because he was evidently not
acquainted with Diesing's original memoir. He expressly
speaks of the " scanty descriptions " hitherto given of the worm.
As Dr White had accurately determined the species in the
presence of an American Scientific Society, it is remarkable
that neither Verrill nor Fletcher should have identified the
worm.

On the 10th of January, 1871, I received a letter from
Prof. W. B. Fletcher, of Indianapolis, Indiana, U.S.A., and in
it he announced that he had " found a worm " infesting the
hog. The parasite was so abundant in swine that he obtained
it in " nine out of ten hogs " which he had examined. Dr
Fletcher sent me specimens of the worm for description and
identification, when I at once recognised them as examples of
Diesing's Stejphanurus dentatus. As Dr Fletcher's first commu-
nication to myself was undated I do not know precisely when
he first encountered the worm, but it was in 1870. In that
same year Prof. Yerrill received specimens of the worm. He
says that they were received from Dr J. C. White. Failing to
identify the parasites as Stejphanuri, Verrill (making no allusion
to the c Proceedings of the Boston Society ') not unnaturally
supposed he had to deal with an entozoon that was new to
science. Accordingly he immediately described and figured
the worm under the combined title of Sclerostoma pinguicola.
If these data are correctly given, the re- discovery of the worm
in America was due to Dr J. C. White ; its identity with
Stejphanurus being subsequently acknowledged by Diesing, and
afterwards, quite independently, by myself. I gather this
partly from Diesing's ' Kleine helminthologische Mittheilungen '
(s. 281), published as a supplement to his ' Revision der
Nematoden ' (1860—61). Until quite recently Diesing's
recognition of the identity of White's parasites with Stephanuri
was unknown in America. My conclusions arose from an
examination of the actual specimens, whereas Diesing was
entirely guided by White's description. In this connection,
moreover, a still more interesting re-discovery remained to be
recorded. The original announcement which I made in the
' British Medical Journal ' for January 14th, 1871, was followed
by another in the same periodical for September, 1871. As
stated in my second letter and repeated in my notice of Krabbe's

410

PARASITES OF ANIMALS

memoir on « Parasites " (' London Medical Record/ April 2,
1873), the President of the London Microscopical Society
(through Mr Slack, who was at that time the secretary)
forwarded to me a box of microscopic slides received by the
Society from Australia. The slides displayed parasites of
various kinds. Having been requested to identify the parasites
I had the good fortune to recognise amongst them characteristic
examples of Stephanurus dentatus. Thus was first made known
the fact that this singular genus was not confined in its
geographical distribution to the two American continents, but
that it extended to Australia. The order of the principal
" finds " and descriptions may therefore be thus restated.
Natterer discovered the worm in Brazil in 1834. Diesing
described it in 1839. Dr J. C. White re-discovered and
identified the worm in 1858. It was subsequently found by
Dr N. Cressy and by Dr Fletcher. These three observers all
encountered the parasite in the United States (1858—70).
Prof. Verrill re- described the worm as new to science in Sep-
tember, 1870. Diesing confirmed White's diagnosis in 1860.
I identified the worm from Fletcher's "find''' in 1871. Dr
Morris supposed he had discovered a new entozoon in Australia
in July, 1871. The Australian worms were identified by me
as examples of Stephanurus dentatus in October, 1871..

The importance of Stephanurus in relation to porcine epizooty
and the supply of animal food cannot be ignored. As remarked
in my communication to 1 Nature,' it must be quite obvious that
so large a parasite, when present in the hog in any considerable
numbers, would give rise to serious disease, even if it were not
productive of fatal results to the bearer. In one of his nume-
rous communications to myself, Prof. W. B. Fletcher writes
as follows : — " It is my opinion that this parasite is the cause,
in some way, of the hog cholera, which has created such sad
havoc within the past ten years over the pork-producing parts
of America. One farmer told me, a few days ago, that within a
month his loss alone from this cause was over one hundred
head; and sometimes, in one neighbourhood, in a few days'
time, thousands have perished, although this season is not a
cholera year, as our farmers say. I advised one farmer to
burn or bury the dead animals, but he informed me that he
believed that fewer hogs die of the disease after eating the
dead animals than those kept from them. Unfortunately, in
this State there is no law guarding the spread of disease,

PA 0 HYDE KM ATA

411

neither is there any reward of reputation or gain for pursuing
any investigation that would bring pork and beef packers into
disrepute. I myself could not get a pig's kidney or beefs
liver in our city market, because I made investigations in some
Texas cattle (being cut up in our market), which damaged
their sale a few years ago." In a third letter Dr Fletcher
tells me that greater facilities for examining the carcases of
hogs had since been accorded him through the liberality of
a Liverpool firm of pork-packers, who had already killed 75,000
hogs during the summer season, i.e. up to the date of the first
week in July. In hot weather the slaughtering is conducted
in ice-houses. Prof. Fletcher's views receive confirmation from
the statements made by Dr Morris, who speaks of the pigs as
dying from some mysterious disease, and thinks that the worms
may be the cause of the porcine mortality. Writing to the
President of the London Microscopical Society from Sydney
(July 12th, 1871), Dr Morris says: — "It is just possible that
some pigs may survive the irritation such a swarm of young
worms must set up ; others, again, may die from peritonitis,
hence the sudden deaths amongst the pigs." I think Dr
Morris' view is perfectly correct, but whether it be so or not, it
is (as observed by me in ' Nature ') interesting to notice the
remarkable correspondency of the conclusions arrived at by Drs
Fletcher and Morris independently. It will probably not be
difficult to ascertain hereafter whether or not the maladies
respectively termed "hog cholera" and "mysterious disease"
are one and the same disorder, but whatever happens in this
respect, it is now quite clear that this parasite, hitherto little
regarded, and for many years past persistently overlooked, is
extraordinarily prevalent in the United States, and, perhaps,
equally so in Australia ; it being further evident that its presence
in the flesh of swine is capable of producing both disease
and death. The statement of the worthy American farmer that
the swallowing of infested flesh by pigs does not necessarily
involve the pig-eating hog in a bad attack of the so-called
" cholera disease " requires to be further tested, and it also
remains to be proven whether or not the Stejphanurus be capable
of passing through all its developmental changes from the egg
to the adult form within the body of the bearer without having
at some time or other gained access to the outer world. The
comparatively large size of the ova, which I find to be about si5"
or more than four times the size of Trichina-eggs, is not with-

412

PARASITES OF ANIMALS

out significance, but as yet we are entirely unacquainted with the
larvae of Stephanurus. If no intermediary bearers are necessary
to its development, we ought not to have to wait long for a
complete record of the life-history of Stephanurus dentatus. In
conclusion, I will only further remark that since thousands of
hogs are infested by this entozoon the subject is worth further
investigation. I believe that Prof. Fletcher brought the matter
under the notice of the United States National Swine Breeder's
Association, which met at Indianapolis in November, 1872, but
with what success I have been unable to learn. The wealthy
agricultural societies of Great Britain pay little or no regard to
the subject of parasites, although thousands of valuable animals
annually perish from the injurious action of entozoa.

Of the remaining nematodes infesting swine I must particu-
larly mention Sclerostoma {Strongylus) dentatum and Strongylus
paradoxus, the last named being generally regarded as identical
with Dujardin's S. elongatus. The first of these two parasites
infests the small intestines, the male and female worms alike
measuring about ^" in length. The females are sometimes a
trifle longer. The Sclerostoma dentatum is an abundant para-
site, infesting all varieties of swine and also peccaries ; but it
is apparently incapable of serious injury to the bearer.
Schneider selected the male S. dentatum for classificatory
purposes. In this worm the arrangement of the rays of the
hood is simple, forming a good central type. Dr D. V. Dean,
in his excellent report of St Louis Board of Health (1874),
speaks of Strongylus dentatus as if it were the same entozoon
as Stephanurus. The confusion of nomenclature would have
been avoided if Diesing had called the renal worm Stephanurus
Nattereri. I hope this title will yet be adopted to prevent
future mistakes. The lung-worm (S. paradoxus) is by no
means harmless, being a frequent cause of fatal husk in young
pigs. It is a viviparous worm, the females acquiring a length
of whilst the males rarely exceed f" '. Under the title

Gongylonema pulchrum, Molin has noticed yet another filari-
form nematode infesting the wild hog ; and, lastly, the lamented
Russian traveller, Fedschenko, has published a full description
of a new species of Gnathostoma (0. hispidum), which infests
the coats of the stomach alike of the wild and domestic hog.
One of the most interesting parasites of swine is the large
acanthocephalous entozoon (Echinorhynchus gigas). It infests
the small intestines both of the wild and domesticated hog, aud

PACHYDERM ATA

413

it was also obtained by Natterer from the collared peccary of
Tayazou. Common as the great Echinorhynchus is in the
United States (and it is scarcely less so on the Continent) I
believe that few, if any, of the museums in the United
Kingdom of Great Britain contain this large entozoon. It is a
curious fact that it does not exist in the Hunterian Collection,
where, however, there is displayed a very fine set of acantho-
cephalous parasites from whales. When in the year 1865 I
mounted, with my own bands, 200 preparations of entozoa for
the Museum of the Eoyal College of Surgeons, I had not so
much as seen a specimen of this worm. Much scientific interest
attaches to this parasite from the fact that Schneider discovered
that the embryos of E. gigas take up ' their residence in the
larvae of the cockchafer (Melolontha vulgaris). He thinks it
identical with the Echinorhynchus hominis of Lambl. Leuckart
disputes this identity, and compares Lambl's worm with the
Echinorhynchus angustatus of our fresh-water fishes. The
E. spirula of certain Brazilian monkeys and of the Barbary
ape bears a strong resemblance to the species from the hog.
On the strength of Lambl's case — and it is the only genuine
instance of the kind on record — Prof. Leuckart devotes no less
than 125 pages of his great work to the consideration of the struc-
ture and development of the thorn-headed intestinal worms.
This worm demands especial attention. Speaking of the
hog's Echinorhynchus, Prof. Verrill, in his c Connecticut Report/
says that " sometimes the intestine of a hog is found perforated
by so many holes that it cannot be used in the manufacture of
sausages." From Mr George Wilkins I learn that the pig-
slaughterers of our English metropolis are well acquainted with
these perforations, which are sometimes so numerous that the
gut looks as if it had been " riddled " with swan-shot. No
wonder that diseased hogs, afflicted with these formidable
parasites, go about, as Verrill expresses it, " continually
squealing and grunting, especially in the morning." That
they are also " cross and morose, and given to biting and
snarling at their companions," is by no means astonishing.
" In severe cases," remarks Verrill, " hogs afflicted with this
parasite are weak in the loins, and have the membranes in the
corners of the eyes swollen, watery, and lighter colored than
usual." It is some comfort to know that Lambl's human case
is unique, and that so long as people abstain from eating cock-
chafer larvae they are not likely to be infested by Echino-

414

PARASITES OF ANIMALS

rhynchus gigas. In the first book of this work I have given
my reasons for not regarding Welch's " encysted Echino-
rhynchus in man " as a genuine example of this curious genus of
entozoa.

The external parasites of swine are not so numerous as might
be expected from the habits of their hosts. The most common
ectozoon is the hog louse {Hcematopmus suis). This disgusting
little insect is about in length. Almost equally common is
the hog mite. Though hitherto considered as a distinct species
(Sarcoptes suis, Grurlt), it is regarded by Megnin as a mere
variety of Sarcoptes scabiei. As Grerlach and others have
remarked, it is readily transmissible to man. The Sarco]>'><
squammiferus, of Furstenburg, is only another name for this
variety of 8. scabiei. Speaking of this scab-insect Megnin
says : — " This parasite was first encountered by Spinola and
Grurlt, and afterwards by Muller." He then adds : — " A
Ceylon wild boar died at the menagerie of the Museum of
Paris of a chronic affection of the skin which had transformed
its integument into a vast fo'c/ieu." Lastly, as regards the
protozoal parasites I can only remark that the psorosperms
(spoken of as Raiuey's corpuscles or as Miescher's utricles) are
often very abundant in the flesh of otherwise perfectly healthy
swine. Having dwelt upon the character of such organisms
in the first moiety of this work, I will only remark that the
full significance of these singular bodies yet remains to be
determined. Rainey's notion that they represented early stages
of cysticercal growth is altogether untenable. According to
Behrens, as quoted by Davaine, psorosperms are especially
abundant in the flesh of swine which have recovered from the
disease called mal rouge. On the subject generally, the writings
of Rivolta, Waldenburg, Eimer, and Siedamagrotsky are espe-
cially trustworthy. Full references to these and other authorities
are given in the synopsis of the 2nd edition of Davaine's well-
known treatise.

Bibliography (No. 54) . — (Anonymous), " On Parasitic Maladies,
especially Measles, of the Pig," from ' Scottish Farmer and
Horticulturist/ in ' Edin. Vet. Rev./ p. 688, 1861.— Ballard, E.,
" On Diseased Meat, and what to observe in cases of suspected
Poisoning by Meat or Sausages (infected with Entozoa, &c.),"
'Med. Times and Gaz./ Jan., 1864<.—Bowditch, H. J., " ~Rnvr
Pork as an Aliment (without reference to the question of
Entozoa.— T. S. C.)," ' Boston Med. and Surg. Journ./ vol. lv,

PACHYDERMATA

415

1857; see nlso 'Comments/ vol. lvi, pp. 23 and 69, 1857. —
Cobbold, " On the Discovery of Stephanurus in the United
States and in Australia/1 in 'Nature/ Oct. 21, 1871, p. 508,
and in ' Brit. Med. Journ./ Jan. and Sept., 1871 ; also in the
'Monthly Micros. Journ./ Nov., 1871— Idem, "Internal Para-
sites of the Hog/'' in ' Manual/ chap. xii. — Idem, " On Cystic
Entozoa from the Wart-Hog and Eed River Hog./' ' Proc. Zool.
Soc./ 1861.— Idem, "On Simondsia," 'Entoz./ p. 79. — Idem,
" Note on Worms in the Langs of a Pig/' in the ' Field ' for
Jan. 9, 1864. — Idem (in relation to Cysticerci or Measles,
see Bibl. Nos. 13 and 14, and, for remarks on psorosperms,
Bibl. No. 41). — Gressy, N., ' On the Diseases of Domestic
Animals in Connecticut (2nd and 3rd Ann. Reports)/ Hartford,
U.S., 1873-74. — Idem, "The demands of Agriculture on Vete-
rinary Science," in 'Rep. of the Mass. Board of Agric./ 1874.
— Idem, 'Find of Sclerostoma ' (quoted by Verrill). — Grisp,
" Note on Hydatid Cysts in the Abdominal Cavity of various
Hogs," 'Path. Soc. Trans./ 1863.— Dardel (see Bibl. No. 14).
— Davaine " Ladrarie chez le pore," in his ' Traite/ 2eme edit.,
p. 668 (see also the writings of Delpech, Cuardia, and especially
Reynal, quoted at p. 674). — Dean, D. V., "On Meats and
Parasites," in ' Seventh Ann. Rep. of Board of Health of the
City of St Louis/ 1874, p. 58 et sea. — Diesing, ' On Stepha-
nurus ' (quoted in text above). — Dupuy, "Hydatid in a Pig,"
from ' Journ. Theorique et Prat./ in the ' Veterinarian/ vol. iv,
1831, p. 285. — Fedschenho, 'Description of new Species of
Tetrastemma, Prorhynchus, and Grnathostoma ' (in the Russian
language), Moscow, 1872. — Fleming, A., " Measly Pork as Food,
for Man," ' Edin. Vet. Rev./ vol. i, p. 485, 1858-59. — Idem,
" On the Measle of the Pig, and on the Wholesomeness, as
Food for Man, of Measly Pork," 'Dubl. Quart. Journ./ 1857.
— Fletcher (quoted in text above). — Florman (quoted by Ru-
dolphi, ' Synops./ p. 620, 1819; and by Davaine, 1. c, p. 723,
1878), in ' Vetensk. Acad. Nya Handlingar/ 1810, pp. 179-182.
— Oairdner, W. T., " Case of Tapeworm occurring in connection
with the Eating of Raw Pork," ' Edin. Month. Journ./ 1856,
and in the 'Veterinarian/ vol. xxix, p. 228, 1856. — Gamgee, J.,
" On Diseased Meat," ' Pop. Science Rev./ Jan., 1861. — Gordon,
"On Tapeworm from Unwholesome Food," 'Med. Gaz./ 1857.
— Gross, 8. D., " Note on the frequency of Acephalocysts in
Swine at Cincinnati," in his ' Elements of Path. Anat./ p. 118,
181-5. — Gurlt, F. I<\, ' Lehrbuch der path. Anat. der Haus-

410

PARASITES OF ANIMALS

Saiigethiere/ 1831, s. 46, 51, 112, 385. — Heller (see Bibl.
No. 13). — Krabbe, ' Husdyrenes Indvoldsorme ' (1. c, in text;
see also review in ' Lond. Med. Rec./ April 2, 1872, p. 206). —
Leidy, " Note on Trichina spiralis from the Pig," from ' Rep.
Acad. Philad./ in ' Ann. Nat. Hist./ vol. xix, 1847. — Leuckart
(see Bibl. No. IB).— Lewis (Bibl. No. 13).— Martin, J., " Case of
Hydatids in the Liver of a Sow," ' Trans. Yet. Assoc./ pp. 330
and 364, 1842-43.— Megnin (Bibl. No. U).—Molin, ' Una
Monog. del Gen. Spiroptera/ Wien, 1860. — Morris, "Report
on Australian Parasites," ' Month. Microsc. Journ./ Nov., 1871.
— Percy, 8. B., " On Diseased Meat in relation to Public
Health (Prize Essay)," ' New York Med. Journ./ 1866.— Idem,
" On the Food of Cities (an Address)," f New York/ 1864.—
Perroncito (Bibl. No. 13).— Putz (Bibl. No. U).—Bainey (Bibl.
No. U)—Bigetti (Bibl. No. 14).— Sawer, A., "Trichina," in
'Bost. Med. and Surg. Journ./ 1865, p. 16. — Schmidt, Max
(see Bibl. No. 51). — Tartivel (Bibl. No. 14). — Thudichum
(Bibl. No. 13).—Tommasi (Bibl. No. 13).— Verrill, "On
Sclerostoma," 'Amer. Journ. Sci. and Arts/ Sept., 1870. —
Idem, " The External and Internal Parasites of Man and
Domestic Animals," from 1 Rep. of the Conn. Board of Agri-
culture/ 1870, p. 109.— Walker (see Bibl. No. 20, 6).— Wheeler,
E. G., " Worms in the Lungs of Swine," ' Bost. Med. and Surg.
Journ./ 1841.— White, J. O., "On Stephanurus," ' Proc. Bost.
Nat. Hist. Soc./ vol. vi, p. 428, 1858.

Part XI (Cetacea).

The parasites of whales are excessively numerous. Unfortu-
nately only a few of the species have been carefully studied,
and much confusion necessarily exists as to the number of
distinct forms. This statement is especially applicable to the
entozoal group, which comprises upwards of a score of species.
Probably Yan Beneden has examined more of these parasites
than any one else, and what little is known respecting them is
for the most part due to his investigations. I have myself
encountered and described several new species — a circumstance
which Prof. Yan Beneden appears to have altogether over-
looked.

Commencing with the flukes, the first species I notice is
Distoma lancea. The late C. M. Diesing's description of this

OETAOEA

417

Fig. 67. — Distoma
lancea. Original.

worm was based upon specimens obtained by Natterer in
Brazil. The worms were discovered in the biliary ducts of a
male dolphin dissected at Barra do Rio Negro
on 29th. December, 1833. Natterer calls this
cetacean the tacuschi, and in a letter to Diesing
names the species Delphinus tacuschi, in order
to distinguish it from the D. amazonicus of Spix
and Martius. Prof. Flower has shown that
Spix and Martius's D. amazonicus is referable to
the inia or Bolivian dolphin (Inia Geoffroyi).
The views of Flower, Natterer, and Diesing are
thus far in agreement ; and the geographical
position of Barra shows that Natterer' s dolphin
could not be the inia, since, as Blyth long ago
remarked, this last named cetacean <( inhabits
only the remote tributaries of the Amazon and
the elevated . lakes of Peru." Several other
dolphins from Brazil have been described, one
of which Mr Gray named Steno tacuxi. I think
that Gray's cetacean answers to the Delphinus
tacuschi of Natterer; but Prof. Flower is of
opinion that Gray's species is an ordinary
Delphinus. In this case it may, he thinks, probably be
referred either to the D. fluviatilis or to D. pallidus. Which-
ever view is correct, it is clear that Natterer's parasite was
obtained from a fluviatile cetacean, and not from an oceanic or
even an estuary form. In Diesing' s original description it is
stated that Natterer found the Distoma lancea " once only,"
when numerous examples were secured. To Dr Anderson I
stand indebted for a solitary specimen, which he procured from
the short-snouted dolphin (Orcella brevirostris, Owen). The
obliging superintendent of the Calcutta Museum obtained this
Distoma on the 3rd of January, 1873. He removed it from the
duodenum, but it had probably escaped from the liver. Be
that as it may, I easily recognised the species by the sinuosities
of the margin of the body. Dr Anderson's parasite does not
exhibit these marginal irregularities so distinctly and sharply
as they are shown in Diesing's figures. Diesing remarks that
the internal organs may be seen through the transparent body.
The uterine organs, crowded with ova and of a purple color,
are represented by him as branched after the fashion of a
raceme. The artist has been misled. The uterine channel is

418

PARASITES OF ANIMALS

not branched. Dr Anderson's specimen showed two large oval
testes placed one above the other in the middle line, and rather
higher up than is usual with those distomes that have the
organs presenting this simple form. The ducts were not visible.
The yelk-forming glands were particularly well marked, con-
sisting of two laterally-disposed masses, the left gland extend-
ing higher up than its fellow. The so-called yelk-cells or
capsules were well seen. The oval-shaped eggs were tolerably
distinct, yielding a length of from pole to pole, by about
§±o" in transverse diameter. The worm, when unrolled, did not
exceed at most, whereas some of Natterer's specimens
measured i" in length. The neck had lost that rounded
character which Diesing called skittle- shaped (Tcegelformige).
The ventral acetabulum is very nearly twice as large as the
oral sucker. Diesing represents the ventral sucker as circular ;
but in Anderson's specimen this organ was broadly oval.

The next fluke I have to notice {Distoma Gampula) is better
known to me. In the twenty-second volume of the ' Linnean
Society's Transactions ' I first described this new fluke, having
secured numerous examples from the peripheral branches of the
biliary ducts of a porpoise (Phocana communis) . The apparently
healthy cetacean was shot by Mr Jardine Murray in the Firth
of Forth, in April, 1855. I mention its condition because
the bile- ducts were found to be diseased in a way similar to
that ordinarily observed in cases of fluke-rot affecting sheep,
cattle, and other animals. In my MS. note-book I remarked :
" The liver-ducts were in several places thickened and knotted
near the surface of the organ. On opening these they were
found to be loaded with small distoniata." It was added that,
so long as the flukes were alive, they displayed under the
microscope a " double and peculiar intestinal tube/' the skin
being clothed with spines arranged throughout with perfect
regularity. When the superficial ducts were dissected out they
presented a distinctly beaded appearance, the enlargements of
the lumen being occupied by flukes closely packed together.
At least twenty were found in one spot. One of these enlarged
ducts is figured in my recent paper to the Linnean Society
(quoted below). The most striking feature connected with the
structure of Distoma Oampula is the twisted condition of the
digestive canals. They present a zigzag appearance, the lateral
folds being so sharp that they seem to constitute, as it were, a
transition between the ordinary simple intestinal tubes of a true

CETACEA

419

Distoma and the branched tubes seen in Fasciola. This led
me originally to place the worms in a distinct genus {Campula).
Perhaps there were no sufficient grounds
for this generic separation ; but in all
Dr Anderson's specimens obtained from
the liver-ducts of the Gangetic dolphin
more or less decomposition of the con-
tents of the intestinal tubes had occur-
red, consequently the angular appearance
of the folds is entirely lost. From the
other characters presented by the worms
I believe that these flukes from the
Ganges are specifically identical with
those originally obtained from the por-
poise of the Firth of Forth. For
reasons elsewhere stated at full length
I have merged my genus Campula into
that of Distoma. Thus, Campula ob-
longa is a synonym only. I cannot
here treat of the morphology of trema-
tode organisation as it deserves ; but
in relation to the question of transition-
forms I may remark in passing that an
extreme degree of intestinal folding
seems as if it must result in branching.
This, I think, would happen should any
departure from the central distome type
be rendered necessary by the exigences yiq. 68. — Distoma Campula. original,
of the creature. At all events, the

spirally-twisted and branched digestive organs constitute diffe-
rent ways in which nature attains one and the same end.
I may add that this coiled condition of the tubes in D.
Campula is by no means unique, since I have seen it in
other trematode forms, as, for example, in my D. compactum
from the Indian ichneumon. Dr Anderson's specimens of
D. Campula furnish a good general view of the reproductive
organs. They show that the single, relatively narrow, and
unbranched uterine canal is of great length, and coiled upon
itself in a very tortuous manner. In this way the duct passes
from side to side, crossing the central line of the body at least
a dozen times, whilst every fold is likewise bent upon itself to
such an extent as to increase its length to at least four times

420

PAEASITBS OF ANIMALS

that of the animal. In short, the uterine folds may be described
as passing from side to side, each separate coil being twisted
upon itself so as to form secondary coils. In the fluke here
drawn I have accurately represented every winding of the duct,
from its vaginal outlet above to its termination, where it is
joined by the ovarian and vitelligene ducts in the ordinary way.
Only the merest traces of these smaller channels were visible ;
but the two oval testes were well defined, occupying a position
somewhat lower down than usual. There was a third organ,
apparently the ovary. This was less well defined, and situated
higher up in the middle line. The vitelligene glands occupied
the usual position. The terminal cells or capsules with their
efferent ducts were well seen in several specimens. The water-
vascular system was constantly visible, or at least that part of
the main channel which expands into a large vesicle imme-
diately above the central point of the tail. At this part several
of the specimens ruptured. In all of the worms the lower end
exhibited a sort of tail, resulting from post-mortem changes.
None of the Edinburgh specimens of Campula displayed either
the slightest trace of this projection or of the water-vessel con-
nected with it. The uterine duct was filled with eggs. Ap-
proximately, the ova gave a measurement of T~ " from pole to
pole by 2^0" in breadth. Although in Anderson's specimens the
integumentary spines had fallen off, they are still attached in
my original specimens from Edinburgh. The spines average
ji57/ in length. With their shafts directed downwards they
separately presented the form of a long cone, the base of which
was onlv T7^" broad. After describing the above-mentioned
trematodes I received a letter from Dr Anderson, in which he
enclosed a sketch of a parasite taken from the small intestine of
another Platanista. The illustration evidently represented a
new species of cetacean fluke which I called Distoma Anderson',
with the following diagnosis : — " Body oblong, smooth exter-
nally, uniform in thickness, six times as long as broad; head
with lateral projections ; ventral sucker large and prominent ;
neck much constricted; tail evenly rounded off, blunt. Length
I", breadth about ±" '." This worm, which was discovered by
Anderson, in March, 1873, is figured in my memoir communi-
cated to the Linnean Society. Only one parasite was found.
The figure in question shows that in this species the testes are
globular and placed high up in the middle line of the body. A
small lobed gland immediately above the testes is probably the

CETACEA

421

ovary. The vitelligene glands are largely developed. In the
year 1858 Van Beneden described a large fluke from the pike-
whale (Balanoptera rostrata). The specimens were from Bs-
chricht's collection and had been removed from the liver. As
some of the examples measured no less than 80 millimetres,
Yan Beneden described them as " the largest known distomes."
This is probably correct, but the great human fluke (D. crassum)
reaches 2\" , and the giraffe's fluke (Fasciola gigantea) 3 inches
in length. The curator of the Australian Museum, at Sydney,
Mr Gerard Krefft, mentions a Distoma which himself and Mr
George Masters obtained from Delphinus Forsteri. Not impro-
bably it represents a new species. Of the single-suckered
flukes, Creplin in 1825 obtained Mono stoma plicatum from the
intestines and oesophagus of a northern whale. This cetacean
was obtained on the coast of the island of Kugen, in the Baltic.
It has been variously spoken of as Balcena horealis or B. ros-
trata, but by Yan Beneden this cetacean is called Balcenop-
tera musculus. The flukes exceeded \" in length. Another
species of monostome (M. delphini) was vaguely indicated by
Blainville as occupying the cutaneous follicles of Delphinus
Dalei, which cetacean is a synonym of Micropteron sowerbiensis.
The same worm is supposed by Yan Beneden to infest the
bottle-head (Hyperoodon butzkopf), and perhaps it was the same
or a similar worm which Poelman found in the flesh of Lage-
norhynchus Eschrichti. By naturalists imperfectly acquainted
with helminths, the monostomes are apt to be confounded with
Cysticerci ; nevertheless, these widely different types may
coexist in the same host. The presence of larval cestodes has
been indicated in various whales. Thus, F. Cuvier and Yan
Beneden state that Surgeon-Major Carnot, in 1822, found an
enormous quantity of small hydatids in the nasal sinuses of a
porpoise (Phoccena compressicaudata). These are supposed to
be Cysticerci. In like manner Mr F. D. Bennett, in 1887,
obtained numerous capsuled Cysticerci from the skin and
blubber of Catodon (Physeter) macrocephalus. It is unfortunate
that so few of the cetacean helminths find their way into the
hands of persons competent to decide upon their true character.

Mr Bennett's " find " was originally stated to have been
made in Balcena mysticetus, but Yan Beneden refers it to the
northern sperm-whale or blunt-headed cachalot. The naturalist
Bosc noticed a larval cestode found in the fatty tissues surround-
ing the reproductive organs of Delphinus dclphis. He called it

422

PARASITES OF ANIMALS

an hydatid (Hydatis), and Rudolphi placed it with the Cysticerci
(0. delphini). According to Van Beneden the parasite in
question is probably a sexually-immature example of the Phyl-
lobothrium delphini described by his son. Edouard Van Beneden
found this scolex in great abundance in a dolphin (Delphinus
delphis), which he dissected at Concarneau in 1868. The
sexually-mature state of this worm is, as the Belgian savans
remark, to be looked for in some one or other of the larger
sharks. The Phyllobothrium has also been found in the black
fish, tursio, or high-finned cachalot (Physeter tursio). M.
Gerrard Krefft has described a cestode from the stomach of a
dolphin [Delphinus Forsteri), which he terms Tania Forsteri.
The strobile only measured 2\" in length. It is just possible
that the worm may be identical with the species found by
Schott. Unfortunately M. Krefft did not find any ova, and his
figures do not indicate the position of the reproductive pores,
if, indeed, they were present. In this place, therefore, it is
fitting to remark that, under the name of Tetrabothrium tri-
angulate, Diesing has furnished the diagnosis of a small cestode

found by Schott in Delphinus
restrains off the coast of Portu-
gal. The strobile measured only
two or three inches in length,
and showed a uniserial dispo-
sition of the reproductive pores.
Remarking on this species Van
Beneden has stated that this is
the only sexually-mature tape-
worm hitherto encountered in the
intestines of the cetacea. This
observation, made in 1870, is
somewhat unfortunate, because
I had already, in the year 1855,
described a very large and ma-
ture form of cestode (Diphyllobo-
thrium stemmacephalum)irom the

F,a M-Diphjllobothrium stemmaoephalum. a, Common porpoise (DelphinUSpho-

f&Sa, SJr^AXS^ coma). As stated by me to the
krged. original. Linnean Society in December,

1857, the small intestine of this porpoise was completely choked
for the space of eight or nine feet by fine tapeworms so closely
packed together that the gut presented the appearance of a

•1

CETACEA

423

solid cylinder. The same porpoise yielded the flukes already
described (D. Campula). As afterwards remarked in my treatise
on the 'Entozoa' (1864), four of the tapeworms measured, respec-
tively, from 7' to 10' in length, the fifth example being relatively
small (18" only). For a full description of the worm I must
refer either to the Linnean ' Transactions ' or to my introductory
volume whence the figures here given are taken. Five of the
finest examples of this remarkable cestode have been added to
the small collection of entozoa which I prepared for the Museum
of the Middlesex Hospital Medical College. The head of this
large cestode is excessively minute. The same cetacean host
not only yielded these new cestodes and flukes, but also great
numbers of the well-known strongyloid lung-worms, called
Prosthecosacter inflexus and P. convolutus. Another species
(P. minor or Pharurus minor) also infests the porpoise, and a
fourth (P. alatus) the narwhal. As I have elsewhere observed
(' Entozoa/ p. 91), the three first mentioned forms are readily
distinguishable from each other by their relative size and length,
and also more especially by the form of the tail. The females
of P. inflexus attain a length of nine inches, and those of
P. convolutus may be upwards of 1\" in length, whilst those of
P. minor do not exceed an inch. The species described by
Leuckart, from Monodon monoceros, is only half an inch long.
All the forms infesting the porpoise were met with by Professor
Quekett, and one of them has been carefully anatomised by
Professor Busk. Probably several other species will be dis-
covered when the lungs and cranial sinuses of the larger cetacea
are carefully examined for this purpose. The form (P. convo-
lutus) here represented is the least known of the three infesting
the porpoise. This species has been dissected by Kuhn and
Eschricht, whilst the other species have not only been examined
by these authors, but also by Raspail, Dujardin, Von Siebold,
Van Beneden, Leidy, and several other helminthologists.
Some of Professor Busk's examples of the male worm (P.
convolutus) were fully fifteen lines long, yet, from the condi-
tion of the internal reproductive organs, he was led to believe
that they were not quite fully grown. I cannot here repeat
the anatomical details given in my former work, but I may add
that all the species of this genus reproduce viviparously. If
the worms are examined in the fresh state the young may
occasionally be seen escaping by the vagina. Professor Van
Beneden noticed this phenomenon in Prosthecosacter inflexus,

424

PARASITES OF ANIMALS

and the same tiling was observed by Busk in P. convolutus.
In the instance here drawn (Fig. 71) one of the embryos is in
the act of emerging, its caudal extremity being still lodged
within the vulva of the parent. In the fresh worm one may also
see, under the magnifying glass, numerous young worms coiled
together within the oviduct ; the last-named organ widening out
into a capacious sac at a little distance above the end of the
tail. The embryos measure about — ' by 3^5" in breadth.

Fio. 70.— Tail of the male Prosthecosacler convolutus. Highly magnified. After Busk.

Higher up, within the uterine and ovarian ducts, the ova may be
seen in all stages of development, according to the particular
region of the tube under examination. In their full-grown
condition the eggs have a longitudinal diameter of Tiy' by a
transverse measurement of about p^o"-

As regards the development and migrations of the young
worms, it is highly probable that the embryos enter the bodies
of various fishes before they have acquired sexual maturity.

CETAOEA

425

Thence they will be passively transferred to the stomachs of
cetacea, whence they bore their way through the tissues to the
bronchi and pulmonary vessels. Though usually found in these
situations they also infest the cranial sinuses. Frostlnecosacter
■minor is frequently lodged within the cavity of the tympanum.
Professor Quekett and myself, working independently, found
examples of P. injiexus occupying the chambers of the heart.
Under the name of Filaria ivflexicaudata, Prof. Von Siebold
has described yet another pulmonary nematode from the porpoise.

It occupied cysts in the lung. Like the strongyloids above men-
tioned, the females are viviparous, but the males are destitute
of any caudal hood. In the whale (Balcenoptera) killed off the
isle of Eugen, and already alluded to, M. Rosenthal obtained a
large number of Filariae {F. crassicauda, Creplin). To employ
Dujardin's words the worms were situated "dans les corps
caverneux du penis d'une Balana rostrata." The males and
females measured respectively 6£ and 13 inches in length.

42G

PARASITES OF ANIMALS!

Several forms of ascarides are known to infest cetaceanB. The
species called Ascaris simplex by Rudolphi was originally pro-
cured from the oesophagus and stomach of the dolphin of the
Ganges and afterwards by Albers from the common porpoise.
According to Diesing the worms obtained by Dussumier from
a dolphin, taken off the Maldive Islands, must be referred to
the same species, but Van Beneden maintains that Dussuniier's
" find " refers to a distinct species, which he calls Ascaris
Bussumierii. To this view I cannot see any objection, but I
think that Van Beneden's retention of Lebeck's Ascaris
delphini as distinct from A. simplex is untenable. Speaking of
examples of this entozoon received from Calcutta, I have re-
marked in the ' Zoological Society's Proceedings ' that Dr John
Anderson's collection of parasites showed four specimens
of this species. The worms had been obtained from the
intestines of Platanista gangetica. Singular to say, all the
examples were of the female sex, the two largest measuring
about 1§" from head to tail. The smaller worms did not
exceed one inch in length. In connection with these specimens
(all of which were carefully examined by me on the 28th of
September, 1875) I have only to add that they presented the
peculiarly flexed state of the chylous intestine described by
Dujardin. As that helminthologist had accurately surmised,
the Ascaris delphini of Rudolphi must be regarded as identical
with this species. It is impossible to say how many distinct
species of cetacean lumbricoid worms exist. Messrs Krefft and
Masters found a species of Ascaris infesting a Delphinus Forsteri
taken off Sidney, New South Wales. Creplin also, in 1851,
described a species (A. angulivalvis) from the intestines of
Balana rostrata. The males are less than three inches long,
the females measuring 3|". The late C. M. Diesing received
from Prof. Steenstrup a notice of some nematodes taken from
a narwhal (Monodon monoceros), which appeared to the
Vienna authority to be scarcely different from Creplin's worm.
Under the title Conocephalus typicus Diesing has both figured
and described a remarkable nematode, two inches in length,
which possesses the power of retracting its conical, or, rather,
mushroom-shaped head within the body, somewhat after the
manner of certain Echinorhynchi. His description is based
upon museum specimens that were obtained from the stomach
of a dolphin (probably Delphinus del phis) captured in the
Atlantic Ocean. In addition to the above nematodes some few

OBTAOEA

427

others remain to be mentioned. Thus, the Peritrachelius insignis
of Diesing was obtained by Natterer from the stomach of
Delphinus amazonicus (Spix and Martius). The largest males
measured 3V and the females h\" .

Another singular parasite, named by its discoverer, Koussel
de Vauzeme, Odontobius ceti, was found by him in the mucus
covering the bristles (fanons) of Baltena australis. The
separate worms measured about ±" in length only, but they
occurred in very great numbers. Lastly, Van Beneden points
to certain filiform worms found by Pallas in the cavity of the
ear of Beluga leucas as probably representing another distinct
species of nematode, which he designates Strongylus Pallasii.
I suspect, however, they will only have been fine specimens of
Prosthecosacter minor. Be that as it may, it is as well to be
reminded that Albers and Mehlis, and also Klein, Camper, and
Bosenthal, found P. minor within the tympanic cavity of the
common porpoise. When looking into the Museum stores of
the Eoyal College of Surgeons, I found many choice examples
of the genus Prosthecosacter. Some few were evidently Hun-
terian, but others had been separately contributed by Professors
Owen and Quekett.

The acanthocephalous entozoa are well represented in whales.
One of the commonest species is Echinorhynchus porrigens, of
which I possess specimens given me by Prof. Goodsir, who
procured them from a Balanoptera rostrata, captured in the
Firth of Forth. The Hunterian Collection contains examples
of this worm, recorded as having been obtained from Baleena
mysticetus ; and also several Echinorhynchi from the pike-whale.
Professor Owen regards these last-named entozoa as distinct
(E. balanocephalus, Owen). Probably Hunter's whale-worms,
which resembled E. fillicollis of the eider duck, and which Van
Beneden has characterised as representing a distinct species
(E. mysticeti), were examples of E. porrigens. The specimens
set up by me for the Museum of the Eoyal College of Surgeons
were all of Hunterian origin. The whole subject of cetacean
Echinorhynchi requires revision, and would well repay special
investigation. The small, oval-shaped entozoa found by Murie
in a whale which Van Beneden refers to Balamoptera musculus
were probably examples of a distinct species of this genus (E.
Muriei). The Vienna helminthologist has described a small
species (E. turbinella) obtained by Hyrtl from Hyperoodon
butzlcopf. The male worms measured \" in leugth. Another

428

PARASITES OP ANIMALS

small species (F. pellucidns) was discovered by Leuckart in the
intestines of Delphinus delphis, the males measuring less than
\" , and the females about \ an inch. Lastly, under the name of
Echinorhynchus brevicollis, Van Beneden has indicated another
species found by Malm in the intestines of a curious whale
(Balcenoptera Sibbaldii) captured alive off the coast of Sweden.
The Louvain savant refers to the " take " of another example
of this rare whale in the Firth of Forth about the same period.
A good many whales have been captured of late years off our
English and Scottish coasts, but, unfortunately, very little effort
has been made to collect the numerous entozoa which they
undoubtedly will have contained.

The external parasites and fellow-boarders or messmates of
Cetacea are almost as numerous as helminths. In this work,
however, little account can be taken of them. Every naturalist
is familiar with the common Gyamus balcenarum, and voyagers
tell us that the whales are sometimes so densely covered by
these lice that they impart to the skin a white color, and so
enable the fishermen to see their victims at a great distance.
The Gyami and Gaprella are closely allied forms of leernodipo-
dous crustaceans. Professor Liitken, of Copenhagen, has
enumerated about a dozen distinct species of Gyami which are
parasitic upon whales. Some of the Pycnogonida are said to
attach themselves to whales. In their young state they are
known to be parasitic upon polyps. I obtained specimens of
these in 1856. The barnacles found on whales are genuine
messmates ; when once they have attached themselves to the
skin, they remain permanently fixed. Like the Gyami or true
whale-lice, these parasitic cirrhipeds are so numerous that almost
every cetacean host maybe said to carry its own species of louse
and its own species of barnacle. The classification of these
creatures is an admitted difficulty, even amongst skilled crus-
taceologists. The genera of cirrhipeds that are parasitic upon
whales chiefly belong to the genera Goronula, Diadema, Tubici-
nella, and Gonchoderma, but in addition to these, many lernaeans
of the genera Penella and Lerneonema are found on whales, and
also, according to Van Beneden, at least one species of Acarus.
This mite (Acaridina balanarum, Van Beneden) is found on
Balesna australis. Here I must stop. The limits of this work
preclude my attempting a more extended notice or enumeration
of the crustacean and arachnidan parasites.

Notwithstanding the known differences existing between the

CETAOEA 429

phytophagous manatee-dugong group and the true whales, the
parasites of this remarkable order of mammals (Sirenia) will be
most conveniently noticed in this place. Not much is known
respecting them. A single species of Amphistome (A.fabaceum)
of the usual size has been described and figured by Diesing
from the caecum and large intestine of Natterer's manatee
(Manatus exunguis), the same mammal yielding a rather pecu-
liar nematode, Heterocheilus tunica t us. This worm possesses a
complicated set of cephalic lobes and oral papillae, which at first
induced Diesing to call the genus Lobocephalus. These struc-
tures have been fully described and figured in Diesing's account
of the anatomy of the worm in the ' Annals of the Vienna
Museum/ The males measure and the females up to 1\"
in length. From the stomach of an Indian dugong (Halicore)
Riippell and several other naturalists obtained lumbricoid worms
(Ascaris halicoris, Owen), the males of which measured 2£" and
the females from four to five inches in length. BuppelPs
specimens were from the Red Sea and Owen's from Penang.
From the now extinct Bhytina Stelleri similar worms were
obtained by Steller, who called them Lumbrici caudidi. Fol-
lowing Brandt's nomenclature the species has since been
recognised as Ascaris rhytina. The worms measured half a
foot in length, and occupied the stomach and duodenum. They
were obtained by Steller in July, 1742, the last of the Rhytinas
having been seen in 1768.

Bibliography (No. 55). — JBaird, " Entozoa of Cete," 'Brit.
Mus. Catalogue/ Index, p. 120, 1853. — Idem, "Ascaris hali-
coris," in 'Ann. Nat. Hist./ 1860, p. 329; see also "Description
of a rare Entozoon from the Stomach of the Dugong," ' Proc.
Zool. Soc./ 1859. — Beneden, P. J. van, "Les Cetaces, leurs
commensaux et leurs parasites/' ' Bullet, de l'Acad. Koyale de
Belgique/ p. 348, 1870. — Idem, 'Animal Parasites/ 1876. —
Beneden, E. van, in ' Comptes Rendus/ 1868. — Bennett, B., in
'Proc. Zool. Soc./ 1837, p. 30. — Brandt, " Asc. dugonis and
A. rhytinee," in ' Bull. P.-Math. de l'Acad. Imp. des Sci. de St
Peters/ torn, v, p. 192. — Oobbold, " Trematode Parasites from
the Dolphins of the Ganges, Platanista gangetica and Orcella
brevirostris," ' Linn. Soc. Journ./ Zool. Div., vol. xiii, p. 35,
1876. — Idem, "Entozoa of Delphinus phoccena," 'Linn. Soc.
Trans./ vol. xvii, p. 167, 1858.— Idem, "Descript. of Asc.
Andersoni," in ' Notes on Entozoa/ part iv, in ' Proc. Zool.
Soc./ March, 1876, p. 296. — Idem, ' Catalogue of the Speci-

430

PARASITES OP ANIMALS

mens of Bntozoa in the Museum of the Royal College of
Surgeons (Preps. Nos. 39-43 and 98-101)/ 1866 ; see also Owen,
for an account of the Hunterian specimens of Echinorhynchi. —
Greplin, F. 0. H., " Note on a Filaria and a Monostoma from a
Whale (Balcena rostrata)," from f Nova Acta Acad. N. C./ xiv,
in ' Zool. Journ./ 1832-34, vol. v, p. 381 ; and in ' Ersch. and
Grube's Encyclop./ 1846, s. 172. — Idem, " On Asc. angulivalvis,"
in ' Wiegmann's Archiv/ 1851, s. 158. — Diesing, "On Hetero-
cheilus and Awphist. fabaceum," in 'Neue Gattungen von
Binnenwiirmen (u. s. w.)/ ' Annalen des Wiener Museums/
1839. — Idem, " Descr. of Conocephalus," in ' Revision der
Nematode^ (1860), from ' Sitzungsb. d. k. Akad. d. w. math.-
naturw./ 1861, s. 669. — Dujardin, ' On Odontobius/ 1. c, p. 292.
— Idem, ' Fch. porrigens,' p. 504. — Idem, ' Fil. crassicauda,'
p. 50. — Idem, 1 Stenurus/ p. 266. — Idem, 'Asc. simplex,' p. 220.
— Krefft, " Parasites of Forster's Dolphin," in his ' Australian
Entozoa/ from f Trans. Etom. Soc. of New South Wales/
Sydney, 1871. — Lebech, "Asc. delphini" (quoted by Rudolphi),
Synops., p. 296, from 'Neue Schriften der Berl. Gesellsch.
.Naturf. Freunde/ Bd. iii, s. 282. — leuchart, B., "On Pharurus,"
in ' Wiegmann^s Archiv/ 1848, s. 26. — Murie, in 'Proc. Zool.
Soc./ 1865, p. 213. — Owen, "Asc. halicoris," e Zool. Soc. Proc./
1838, and in art. "Entoz.," ' Todd's Cyclop/— Idem, "On
Echinorhynchi," in his ' Catal. of the Contents of the Roy. Coll.
Surg. Museum,-' part iv, fasc. i, p. 44 ; and for the " External
Parasites of Wales," ibid., p. 74, 1830. — Quekett, J., " On the
Anatomy of four species of Entozoa," abstract from 1 Proc. of
Micros. Soc./ in 'Ann. of Nat. Hist./ vol. viii, 1842; also in
'Micros. Journ. and Struct. Rec./ p. 125, and in the original
series of ' Trans, of the Micros. Soc. of Lond./ vol. i, p. 44,
1 844. — Idem, in ' Baird's Brit. Mus. Catal. of Entoz./ p. 3,
1853. — Boussel de Vauzeme, "On Odontobius," f Ann. des Sci.
Nat./ 1834, p. 326, and in ' Isis/ 1836.— Ruppell, "Entozoa
Dugonis," in 'Abhandl. der Senkenbergschen Museums/ i,
s. 106. — Siebold, Von, " On Filaria infiexicaudata/ in 'Wieg-
mann's Archiv/ 1842, s. 347.

Part XII (Marsupialia) .

The well-known fact that, in respect of their habits, the
pouched mammals epitomise, as it were, the non-marsupinl
quadrupeds, would naturally lead us to look for a repetition of

MARSUPIAL J A

431

corresponding type-forms of entozoa as guests within their
bodies. Speaking generally, the inference is correct ; but very
few of the entozoa hitherto found in marsupials correspond, as
species, with those infesting man and non-pouched mammalia.
A noteworthy exception occurs in the case of the common liver-
fluke, which is abundant in the great kangaroo (Macropus
major) . This fact was well known to Bremser and all the older
helminthologists, and it has since been confirmed by numerous
observers resident in Australia. The late Dr Eowe, an acute
observer and successful stock-breeder, who wrote chiefly in
connection with the sanitary bearings of parasitism, remarked
that " the native animals of Australia are much infested with
internal parasites. Some of those now found in the kangaroo
and the smaller marsupials may have been derived from
our domestic animals ; but tapeworms and other internal
parasites have been met with in animals occupying regions
wholly unsettled." Precisely so. That is just what we should
expect. The Australian indigenous mammals have their own
entozoa as a matter of course, and, in addition, they have con-
tracted a few species from the domestic animals introduced into
the country. On the whole, however, it cannot be said that
the parasites of marsupials are of much practical consequence
to agriculturists, since, with the exception of flukes, and pro-
bably hydatids, the Australian marsupials do not appear to
harbor any entozoa that are likely to prove injurious to man
and his domestic companions. The amount of fluke-germ dis-
tribution by kangaroos must be infinitesimal as compared with
that proceeding from sheep and other kinds of " stock there-
fore on the score of parasitism alone it is not desirable to
hasten the slaughter of kangaroos. From the scientific stand-
point, it is to be regretted that the naturalists of New South
Wales and other colonies have done so little towards defining
the various species of marsupial entozoa. Mr Krefft, in his
interesting brochure on Australian entozoa, describes a few
tapeworms, and also points to several round worms which may
be new to science, but with the exception of the common fluke
no trematode appears to have been encountered by himself or
Mr Masters in the various marsupials which they examined in
the neighbourhood of Sydney and Queensland. Dr Bancroft,
of Brisbane, has placed in my hands a small collection of
entozoa, several of which have been obtained from marsupials,
but their identification remains partly in abeyance.

432

PARASITES OF ANIMALS

Besides the liver-fluke, the only marsupial trematodes at
present fairly described appear to be Hemistoma alatum, and
two species of Rhopalophorus (R. coronatus and R. horridus).
All of these were obtained by Natterer from the opossums of
tropical America. One of these flukes was described at some
length by Rudolphi, who called the species Distoma coronatum,
and gave its length as varying from two to four lines. Diesing,
in one of his best illustrated monographs, has shown that
the opossums in question are infested by two distinct species of
fluke, which must be generically separated from the distomes.
These singular Rhopalophori are furnished with a pair of armed
retractile proboscides (Bohriisseln), which must form powerful
organs of anchorage. The worms are found attached to the
walls of the stomach and small intestines.

The tapeworms of marsupials are more numerous than flukes.
Thus, we have Rudolphi' s Tcenia f estiva, eight to ten inches in
length, occupying the gall-bladder and hepatic duct of Macropus
giganteus. Dr Bancroft's collection contains two almost perfect
examples of a tapeworm which he procured from a small
streaked kangaroo (Hahnaturus Derby anus). These I have
identified as T. /estiva. In this worm the reproductive papillae,
not hitherto observed, are biserially arranged. Fragments of a
tapeworm (T. didelphidis) are preserved in the Vienna Museum,
taken from the intestines of the American Didelphis murina.
From different species of wallaby (Halmaturus) Mr Gerard
Krefft has given more or less complete descriptions of two
tapeworms (Tcenia fimbriata, and T. Mastersii), and a probable
Bothriocephalus (B. marginatus). I am not in a position to
pronounce upon the distinctness of these Australian Teenies ;
but I may observe that Krefft' s T. Jimbriata comes very near
to another species which Dr Bancroft has given me. The
Brisbane savant obtained the worm from a koala (Phascolarctos
cinereus). Provisionally I call this species Tcenia geophiloidrs,
in allusion to its general resemblance to a long millipede. The
single, perfect strobile, with the head attached, measures
thirteen inches in length. Prof. Leidy has furnished a de-
scription of another tapeworm (T. bipapillosa) from a wombat
(Phascolomys), and Mr Krefft has described yet another species
obtained from the common vulpine opossum (Phalangista vul-
pina). The single example in Krefft' s possession measures
four inches in length. He has named it Tcenia phalangistce.
Some of the American opossums (Didelphys brachyura, and D.

MAESUPIALTA

433

quica) have been found to harbor a species of ligula (L. reptans,
Diesing) in the sexually-immature state. Lastly, I find in
Bancroft's collection several tapeworms obtained from that
small and interesting monotreme marsupial commonly known
as the Australian hedgehog, echidna, or porcupine ant-eater of
the colonists (Tachyglossus setosus). The strobiles, which are
nearly perfect, average three inches in length, and are made of
very narrow and closely-set proglottides. This species is evi-
dently new to science, and as such I propose to call it Ttenia
phoptica, in allusion to its thick-set appearance and its conse-
quent burdensome character to the bearer. The largest pro-
glottides measure fully §" in width. There can be little doubt
that the presence of any considerable number of such compara-
tively large tapeworms must seriously incommode, if they do
not actually prove fatal to their unfortunate hosts.

So far as regards mere variety of species, the nematode
fauna of marsupials is probably far in excess of that of the
trematodes and cestodes. I cannot therefore do much more
than enumerate the species. The Hunterian Museum of the
Royal College of Surgeons contains the original " worms found
alive within the capsular ligaments of the knee-joint of a
kangaroo," which have been indicated as Filarice macropodis
gigantei. It would, in my opinion, be far better to call the
worm, after its discoverer, Webster's filaria (F. Websteri). Dr
Bancroft has likewise encountered this same parasite in the great
kangaroo. From the abdominal cavity of a wallaby Leidy has
also obtained a filaria (F. spelcea). The American opossums
(Didelphys) are much infested by Ascaris tentaculata, which is
found in the csecum, and many of them also harbor a small
whipworm (Trichocephalus minutus). Another tolerably abun-
dant nematode (Physaloptera turgida) was obtained by Natterer
from the stomach of Didelphys azaree, D. myosurus, and D.
cancrivora. The only other nematode mentioned in Diesing's
'Systema' is Aspidocephalus scoleciformis. This is found in
D. murina, and D. domestica. From the opossum of the
United States (D. virginiana) Prof. Leidy has also obtained
Asc. tentaculala, Trichocephalus minutus, and Physaloptera (Spi-
roptera) turgida. According to Molin there are grounds for
separating some of these stomach-worms into distinct species.
Thus, he has recognised the examples found by Natterer
in Didelphys myosurus as belonging to the genus Histiocephalus,
as emended by himself. If the separation be allowed, then we

434

PARASITES OP ANIMALS

must add to the list Molin's Histiocephalus subulatus. From
Bancroft's collection I am certainly in possession of two dis-
tinct kinds of nematode taken from the stomach of Halmaturus
Derbyensis. I have also two species of nematode from Ma-
cropus giganteus. Their identification, however, is a matter for
future consideration. Lastly, as regards the acanthocephalous
parasites, only one species appears to have been described.
This worm (Ecliinorhynchus micro cephalus) was obtained by
Olfers in Brazil, from the intestines of Didelphus philander.
It also occurs in D. virginiana. Being a tolerably large species,
that is to say 3" in length, it seems surprising that it has not
been found in the American opossums generally.

Bibliography (No. 56). — Gobbold (see Eowe). — Diesing,
' Syst./ 1. c, p. 519; also monogr. ' Binnenwiirmen ' (1. c,
Bibl. No. hb)—Krefft, G., 'Australian Entozoa' (I.e., Bibl.
No. bh).—Leidy, ' Proc. Philad. Acad./ 1856.— Idem, " Tape-
worm from the Wombat," ibid., 1875, p. 6. — Molin, " Una
monogr. del genere Physaloptera," p. 10, " Una monogr. del gen.
Dispharagus/ and ' Una monogr. del gen. Histiocephalus/
p. 37, aus dem xxxix Bd., d. ' Sitz. d. m.-nat. CI. d. k. Akad.
der Wissensch./ s. 479-507 und s. 637-672, I860.— Bowe, J.,
" Parasitism in Australia " (in which the author sought to
benefit stock-owners by utilising, prophylactically, my pub-
lished opinions), repr. from the 'Melbourne Leader ' in the
'Veterinarian/ May, 1874. — Budolphi (in ref. to Dist. coro-
natum), ' Synops./ p. 116 and 686. — Webster (respecting his
" Filaria/' see), ' Catalogue of the Hunterian Collection of
Bntozoa/ p. 7, prep. 49, 1866 ; also the old ' Catal. of the
Museum Koy. Coll. Surg.' (by Owen), part iv, fasc. i, p. 37,
No. 170; also Diesing's 'System/ p. 280, and Froriep's
' Notizen/ Bd. xlii, s. 328.

SECTION II.

To give an air of completeness to this treatise, I shall devote
the few pages remaining at my disposal to a brief summary of
the general facts of parasitism as witnessed in birds, reptiles,
fishes, and evertebrated animals. For details I must refer to
the separate original works and memoirs quoted in the appended
bibliographies.

AVES

435

Part I (Aves).

A prodigious number of entozoa are known to infest birds.
So far from birds being less victimised than mammals, the
contrary is the case. Every now and then avian epizootics,
due to parasites, sweep off hundreds of these attractive hosts,
and in some cases even nestlings are not secure from entozoal
invasion. It might be supposed that predacious birds would
be more liable to invasion than the graminivorous species.
Such is not the case. The eagles, hawks, vultures, and owls
certainly harbor a great variety of helminths, but as much may
be said of the grain-feeding game birds, and still more of the
water birds. Pheasants and land-fowl, grouse and partridges,
are largely infested ; whilst, of water-fowl, herons and plovers,
rails and snipe, ducks and geese, cormorants and divers, gulls
and awks, play the role of host to a practically infinite variety
of parasitic guests. The presence of the worm-guests does not
imply any previously diseased condition of the host. Shoot any
water bird, say an oyster catcher {Hcematojpus) , or, still better,
a grebe (Podiceps) , and then carefully examine its intestinal
contents. You will probably find in its interior flukes and
round worms, tapeworms and Echinorhynchi. Capture and
examine a frog or a salamander. The result is the same, except
that the cestodes would probably be absent. As for fishes, if
entozoa be a proof of cachexia, then it follows that the normal
condition of all piscine hosts is a diseased state. Examine any
tolerably well-grown salmon, trout, pike, perch, roach, chub,
carp, or barbel, and probably any one of them will contain at
least three different kinds of parasites, each of which will be
present in more or less considerable numbers. From what is
stated above it would be obviously futile to attempt even an
enumeration of the species of avian entozoa — a remark which
applies almost equally to the other groups of hosts that remain
for consideration. Confining our attention to a few of the more
noteworthy facts, I may observe that we have no very trustworthy
data respecting the power for mischief possessed by flukes.
From what we know of their destructiveness in man and certain
other mammals, it would be hazardous to pronounce them
harmless. Scientifically, they furnish particulars of great
interest. One of the most striking facts of recent study relates
to Zeller's discovery that the little cercariae (0. exfoliata)
which are contained in a peculiar sporocyst (Leucocldoridiwm

436

PARASITES OF ANIMALS

paradoxum), infesting the tentacles of a snail (Succinea
amphibia), are in reality larval forms of a fluke (Distoma
macrostomum) which, resides in the intestinal canal of warblers
(Syloiadce). By experiment Dr Zeller reared the Distoma in
question in the intestines of whitethroats (Ourruca garrula),
in blackcaps (0. atricapilla) , and in wagtails (Motacilla).
Six days after transference the Cercarise acquired sexual
maturity. The odd thing is, that as the sporocystic Leucoch-
loridia resemble insect larvee, they are attacked and swallowed
by the birds under delusion. It is a curious example of
mimetism in favour of the nuke's welfare. To this and
other equally brilliant scientific results helminthologists were
gradually led up by the earlier researches of Steenstrup and
Van Beneden. As remarked in my ' Entozoa/ ever since
Steenstrup's discovery of the fact that Cercarige found in the
bodies of water-snails were larval flukes, a peculiar interest has
attached itself to this subject. Not only were the conclusions
which he elicited novel in themselves, but they formed a basis
for the enunciation of that interesting " law of alternate genera-
tion " with which the famous Danish naturalist's name will ever
remain associated. In all essential particulars Steenstrup's
statements have been verified.

By way of illustrating the phases of development through
which the distomes pass I cannot do better than recapitulate in
an abridged form the account I have previously given of
Distoma (Echinostoma) militate of the snipe and curlew. This
account is based on the investigations of Van Beneden, Von
Siebold, and Pagenstecher. I regret that it is out of my power
to reproduce the illustrations that have already appeared on
this head in my introductory treatise (see ' Entozoa/ figs. 5 to
9 inclusive). Our Echinostoma militare produces oval-shaped
eggs, which give birth to a free ciliated embryo, and this embryo
produces a sporocyst or scolex by internal budding. When
the sporocyst separates itself from the embryo it presents a
very simple appearance, but showing already a csecal digestive
tube. The tail end is fissured, indicating an early stage of
formation of caudal appendages. In the next stage we have a
well-developed head and body, the tail becoming strongly pro-
nounced. Limb-like caudal lobes project on either side, and
an oral sucker makes its appearance in front. This sucker
communicates with the oesophageal bulb and passes directly
into the digestive caecum, which contains a variable number of

AVES

437

rounded particles. At this stage, also, incompletely developed
CercariEe may be seen in the perivisceral cavity. These Cer-
cariEe are at first shapeless organisms, but after passing through
a series of gradations they ultimately assume a definite form,
which, in many cases, is sufficiently distinctive to enable us to
refer the CercariEe to particular species of Bistoma. The older
writers regarded many of the cercarians as adult flukes. In
the early state these larvae are furnished with tails. They may
be seen lodged within the cavity of the body of the sporocysts,
being twisted and folded in various attitudes. The CercariEe
not only exhibit a cephalic and ventral sucker, but also a dark
forked line representing the digestive system. At a still fur-
ther stage other structures come into view, until the perfect
Cercaria displays an oral sucker, a pharyngeal bulb, an
oesophagus, two alimentary ventral sucker, a water-

vascular system consisting of two main excretory ducts, and a
contractile vesicle, by means of which the ducts communicate with
the external sui'face. The tail is conspicuous and furnished with
a fringe. The alimentary organs conform to the general trema-
tode type, but before passing into the sexually-mature condi-
tion other changes are undergone. The CercariEe part with
their tails, and subsequently they encyst themselves on or
within the surface of the body of some mollusk. Their pupa
condition is thus arrived at. The pupa itself differs from the
cercaria in presenting a double crown of hooks surrounding the
head, but the other organs correspond with those already
described. According to Van Beneden the hooks make their
appearance immediately after encystation. In this condition it
is next transferred to the intestine of some higher animal, and
in this final situation it gradually acquires all those organs the
possession of which will entitle it to be called a sexually-mature
or adult distome. In the immature fluke we may now discern
the mouth, the buccal or cephalic sucker, the pharyngeal bulb,
the oesophagus, the digestive cseca, the coronal spines, the
contractile vesicle, the aquiferous system of vessels, the matrices
of the yelk-formiug glands, and also a central mass of cellules,
from which all the other reproductive organs will in due time
be developed. In the adult Echinostoma militare the upper
third of the body is clothed with little spines. Taking this
example as illustrative of the ordinary mode of fluke develop-
ment we find that a change of hosts is necessary, and that in
the intermediate state they occupy the bodies of mollusks.

438

PARASITES OF ANIMALS

Thus, for the continuation of the species, there must needs be
a contemporaneity of vertebrate and evertebrate hosts. Surely
no reasonable person can ascribe this concurrence to merely
fortuitous circumstances. In this connection I may remark that
Villot, in his account of the migrations of the trematodes, states
that the cercarian forms of Distoma l&ptosomum and D. brachyso-
mium of Tringa alpina occur in Scrobicularia and Anthura. These
parasites are also found encysted in the gizzard of Tringa.
The tapeworms of birds are undoubtedly injurious to their

bearers. All the worms appear to be
armed with cephalic hooks ; at least, such
is the case with the species described
by Krabbe, who has supplied figures of
the hooks drawn to a scale. Dr Krabbe's
beautiful monograph is a perfect model
of its kind. In the accompanying figure
the hooks have fallen (Fig. 72). On ac-
count of the frequency of their occur-
rence, some persons have supposed that
tapeworms are not injurious to their hosts,
forgetting that it is not the mere fact of
the existence of tapeworms, but their
excessive numbers during particular sea-
sons that give rise to avian epizootics.
The same rule holds good with other
parasites. Of course, in fledgelings, as
also obtains in yearlings amongst our
domesticated animals, a very few para-
sites are sufficient to prove destructive
to the bearer. Thus, as regards the
so-called " grouse-disease," during one
season it may be due to tapeworms, dur-
ing another to strongyles, during a third
to excessive abundance of both these
parasites. Unfortunately, other avian
epizootics, not necessarily due to para-
sites of any kind, may be mistaken for
helminthic epizooty. The same thing
happens amongst quadrupeds. We have,
for example, parasitic equine epizootic outbreaks, and likewise
non-helminthic equine epidemics (as in the case of the Egyptian
horse plague of 1876). The true nature of any epizooty can

Fig. 72. — Head of Tmnia paradoxa.
a, Proboscis retracted ; b, end of
the rostellum expanded. Highly
magnified. From an oyster-
catcher (Hiemalopus oslralcgus).
Original.

AVES

439

only be determined by competent investigation. That was
well shown in the grouse epidemic of 1872. In that epizooty
the greater number of the birds succumbed to the injuries
produced by a nematode worm (Strongylus pergracilis, Fig. 73),

Fig. 7S.—Stront/ylus pergracilis. a, Head and neck; b, c. d, e, tail of the male in various
positions; f, tail of the female; g, section showing termination of the oviducts; h, three
eggs. Highly magnified. Original.

but, without doubt, the occasional presence of numerous tape-
worms {Tcenia calva) hastened the consequent fatality. The
following table, giving the results of examination of twelve
diseased grouse from the Earl of Cawdor's estate, will show
how inconspicuous a part tapeworms played in the epidemic
of 1872. For further details I must refer to my brochure on
the ' Grouse Disease/ and to some other memoirs quoted in the
bibliography below.

No. of
specimen.

Condition of
bird.

State of viscera.

Tapeworms
present.

Strongyles present.

Good

Full

None

Abundant.

Emaciated

Putrid

Two

Very numerous.

Good

Full

None

Abundant.

One

Shrunken

Two

Very numerous.

Emaciated

Much shrunken

None

Thin

Shrunken

»>

Emaciated

Distended

Many

Thin

Rather full

Several

Ahunchint.

»»

None

Very numerous.

440

PARASITES OP ANIMALS

I have said that tapeworms prove fatal to young birds,
even to nestlings. A notable instance of this is recorded by
Mr Eames. The parasites were examples of Taenia angulata.
Apart from the epidemiological aspects of the subject, it is not
uninstructive to notice the variety of helminths that infest the
common fowl and game birds. Accordingly, I append a similar
but more extended list than that previously given in the
supplement to my introductory treatise :

xjutozoa of game birds arid the
common fowl.

ommon
fowl.

heasivnt.

J- <u
<u —

« r?

Red
rouse.

Grey
artriuee.

Red
artridge.

— "

P-<

Monostoma verrucosum, Zeder...

Distoma oxycephalum, Rudolphi.

j, ovcitum, Rudolphi

„ lineare, Zeder

„ dilatatum, Miram

...

Filaria Mansoni, Cobbold

Ascaris vesioularis, Froelich ...

Spiroptera hamulosa, Diesing ...

„ helicina, Molin

Dispharagus nasutus, Dujardin .

Slrongylus pergracilis, Cobbold

Sclerostoma syngamus, Diesing .

Trichosoma longicolle, Rudolphi

„ infundibuliforinis, Goeze .

In regard to this list I can only afford space to remark that
several of the species are possibly mere varieties. Some of the
worms are of great interest. It occasionally happens that
Distoma ovatum is found in the albumen of the fowl's egg, and
it is even more common to obtain Ascaris injlexa from the same
situation. For a recent example I am indebted to Dr Walker,
of Peterborough. Spiroptera helicina resides in the feet, occa-
sioning enlargement of the joints and consequent distress to
the bearer. Probably the most important in the list is my
Strongylus pergracilis. Hero I may mention that on the 10th
of April, 1878, I received a letter from Dr Manson, of Amoy,

AVES

441

announcing his acquaintance with a filaria infesting the eye of
the fowl. On the 9th of May I also received from Dr Manson
the head of a bird showing examples of the worm. As the
species is new to science I have proposed to call it Filaria
Mansoni, after the discoverer. The male is §" and the female %'
in length. Of the injurious nematodes, giving rise to avian
epizooty, probably one of the most destructive is Ascaris
maculosa infesting pigeons. On the 9th of October, 1873, I
received a letter from Dr J. Alexander Macdonald, of Woburn,
Bedfordshire, stating that he had forwarded to me a pigeon
which had been found dead on the previous morning. It
seems that the owner of the bird had erected a large pigeon-
house, and had imported a number of Antwerp smerles, these
birds all continuing in a perfect state of health until about a
week before the above-mentioned date, when, to use Dr Mac-
donald's words " first one and then another was attacked, and
so on, until four or five of the pigeons had died after a few
hours' illness." The suddenness of these attacks not unnatu-
rally suggested poisoning ; and, accordingly, says my informant,
the owner " had the curiosity to open one of the birds, when,
to his astonishment, he found the intestines stuffed with
worms." Two days later I received a letter from Dr Mac-
donald, stating that several others of the flock had died, and it
further appeared to him probable that the daily list of sick and
dying would continue to increase. On the 14th of the month
three more of the birds were dead. On the 4th of the follow-
ing November, the same correspondent obligingly informed me
that the epidemic had been "at last mastered." It seems that
altogether twelve birds had perished. On the 9th of October
one of the birds was carefully examined by me, and the results
were so interesting that I am constrained to give a few of the
particulars originally communicated to theZoological Society. As
stated in my paper, the whole intestinal tract of the dead bird was
crowded with these ascarides. The small intestine was inflamed
throughout, and showed several large ulcerated patches ; never-
theless, there were no traces of emaciation. From this it was
evident that the parasites had grown quickly, the malady
having a correspondingly rapid formation. The distribution of
the parasites was curious. One specimen, two inches long,
reached from the crop to the proventriculus. The cavities of
this organ and of the gizzard were crammed with worms com-
pletely blocking the passage. Three of the worms had also

412

PARASITES OF ANIMALS

placed themselves within the pyloric opening, their bodies
partly lodging within the duodenum. The duodenum itself
was crowded with worms, their numbers somewhat decreasing
downwards. I removed thirty-six worms from the oesophagus,
proventriculus, and stomach, besides 166 others from the intes-
tinal canal, thus obtaining a total of no less than 202 nematodes
from this small host. Considering the large size of these
entozoa, the extent of infection must be pronounced remarkable.
The largest females measured 2\" in length. One of the most
interesting facts — serving to exemplify a well-known habit of
lumbricoid worms generally — consisted in the circumstance
that two of the parasites had succeeded in perforating the
horny lining membrane of the gizzard. The injuries had been
accomplished during the life of the host, for the walls of the
gizzard were inflamed opposite the perforations. There was a
little half digested food within the stomach, the debris of which,
when placed under the microscope, showed several ova. There
were no free embryos, neither had the development of the
freed eggs proceeded beyond yelk-segmentation. Free eggs
were also found both in the small and large intestine. The
eggs measured about ^k" by — g" in diameter. Eeferring to
my paper for further anatomical details, I can only add that,
despite these facts, the ascarides in question do not appear to be
a very frequent source of epizooty. It was remarked by Dujar-
din that Heister, at Rostok, and Grebauer, at Breslau, found this
parasite abundant at the beginning of the eighteenth century ;
but, according to examinations conducted at Vienna, the worm
was found in the common pigeon in only eleven instances out
of 245, and thrice only in thirty-eight examples of the ring-
dove ; moreover, the examination of eighty-seven other pigeons
and doves of different species yielded entirely negative results.

The Dublin helminthologist,
Bellingham, noticed the occur-
rence of this parasite in Ireland.

Any attempt so much as to
enumerate the species of nema-
toids infesting birds would carry
me far beyond the aim and scope
of this treatise. One of the com-

pI0. 74— Tail of the male Mcarh vesiartaris. , . ,

From a ring-necked pheasant. Original. monest species IS Ascaris (ilere-

ralds) vesicularis. Many hundreds of forms have been described

by Dujardiu, Diesing, Molin, Krabbc, and other systematists,

AYES

413

and it remains for some future laborer to condense the facts
which are dispersed throughout a very wide-spread literature.
As regards the particular species of nematoids that are either
actually known or conjectured to be injurious to birds I can only
find space to repeat some of the particulars which I have else-
where recorded in respect of Sclerostoma syngamus. In 1799
a letter from Dr Wiesenthal, of Baltimore, U.S., was published
in the ' Medical and Physical Journal/ containing an account of
a parasite infesting the trachea of fowls and turkeys in America.
The communication is dated May 21st, 1797, and is the first
public record concerning the entozoon. Dr Wiesenthal says :
" There is a disease prevalent among the gallinaceous poultry in
this country, called the gapes, which destroys eight-tenths of
our fowls in many parts, and takes place in the greatest degree
among the young turkeys and chickens bred upon old-established
farms. Chicks and poults, in a few days after they are hatched,
are found frequently to open their mouths wide and gasp for
breath, at the same time frequently sneezing and attempting to
swallow. At first the affection is slight, but gradually becomes
more and more oppressive, and it ultimately destroys. Very few
recover ; they languish, grow dispirited, droop, and die. It is
generally known that these symptoms are occasioned by worms
in the trachea. I have seen the whole [windpipe] completely
filled with these worms, and have been astonished at the animals
being capable of respiration under such circumstances."

Any one who has witnessed the gapes will at once recognise
the accuracy of Wiesenthal' s description ; and so far as the
phenomena of the disease are concerned, very little more
has been added in the numerous accounts which have since
appeared. On the 1st of August, 1808, the English naturalist,
George Montagu, communicated to the Wernerian Society
a paper entitled " Account of a species of Fasciola which
infests the trachea of poultry, with a mode of cure." Mon-
tagu does not appear to have been aware of the existence
of any previous record. He gave a scientific description of
the parasite, which led to its being noticed in the systematic
works of Eudolphi, Dujardin, and Diesing, but the best accounts
of the worm are due to Von Siebold. Sclerostoma syngamus
has been found in the trachea of the turkey, domestic cock,
pheasant, partridge, black stork, magpie, hooded crow, green
woodpecker, starling, and swift. In July, 1860, I obtained a
fowl suffering from the gapes, and operated upon it in the

444

PARASITES OF ANIMALS

following manner: — A small portion of wool having been
dipped in chloroform and placed in front of the nostrils the bird
soon became insensible. The skin of the neck was then divided
and the trachea slit up to the extent of a quarter of an inch.
With a pair of common dissecting forceps, I removed seven
Sclerostomata. Six of these parasites were sexually united,
the odd worm being a female. After I had closed the external
wound with a single thread the bird woke out of its artificial
sleep, when it soon recovered its legs, and ran about the table
vigorously. Moreover, in a very few minutes it devoured the
contents of a saucer partly filled with bread and milk. An
occasional gape was caused by an accumulation of frothy mucus
within the injured trachea, but this obstruction the bird soon
got rid of by shakes of the head and sneezing. The only sub-
sequent inconvenience to the bird arose from emphysematous dis-
tension of the cellular tissue of the head and neck. This was
relieved by puncture, the emphysema ceasing to form after the
external wound had healed. Some months afterwards I destroyed
the bird, and on dissecting the neck, a distinct cicatrix was
found indicating the site of the operation on the trachea. The
divided cartilaginous rings, six in number, were united only by
a thin layer of connective tissue. The female worms gave an
average length of f " , the males scarcely exceeding \" . The
mouth is furnished with six pi'ominent chitinous lips. In both
sexes the surface of the body is quite smooth, but the tail of
the female exhibits a tendency to fold upon itself. The
lower part of the body suddenly contracts to form a short,
narrow, mucronate, pointed tail. The male is usually found
rigidly affixed by means of a strong, membranous, sucker-like
bursa, which proceeds from the lower end of its body. In
regard to the peculiar mode of union of the sexes, it becomes
an interesting point to ascertain whether there be an actual
incorporation of the substance of the copulatory organs during
or after the act of impregnation. In my specimens none of
the three pairs were organically united, and I succeeded in
separating one pair very readily. Dujardin speaks of them as
being soldered together, whilst the statements of Von Siebold
are still more explicit. In connection with this subject the
latter observer makes the following comment (' Wiegmann's
Archiv/ 1836, s. 106) : — " The two sexes of almost all round
worms are united only at the time of copulation. The male of
Hctcroura androjplwra has also the habit of remaining connected

AYES

445

with its mate beyond the period of copulation ; here, thus,
there is a continuous union of the sexes without a growing
together ; and in Syngamua trachealis there is ultimately a lasting
continuity of the sexes by means of an actual growing together.-"
Having confidence in Yon Siebold's statement, I concluded that
the sexual union in my specimens had only recently been
effected. Admitting this to have been the case, one naturally
asks how the mature eggs can make their escape. Clearly, the
eggs can only escape by an eventual breaking up of the body
of the parent. The eggs of Sclerostoma syngamus are com-
paratively large, measuring 5^" in length. Many of the ova
contained fully formed embryos, and in the centre of the lower
third of the body of one of them I perceived an undulating,
imperfectly formed intestinal tube. By whatever mode the
young escape the shell, it is clear that they are already suffi-
ciently developed to undertake an active migration. A change
of hosts is probably necessary, but in the first instance they
either enter the substance of fungi or other vegetable matters,
or they bury themselves in the soil at a short distance from the
surface. In view of checking the destructive influences of this
parasite, the following methods have been recommended.

First. The simplest plan consists, as Dr Wiesenthal long ago
pointed out, in stripping a feather from the tube to near the
narrow end of the shaft, leaving only a few uninjured webs at
the tip. The bird being secured, the webbed extremity of the
feather is introduced into the windpipe. It is then twisted
round a few times and withdrawn, when the worms are found
attached. In some instances this plan succeeds entirely.

Secondly. The above method is rendered more effectual when
the feather is previously steeped in some medicated solution
which will destroy the worms. Mr Bartlett employs salt for
this purpose, or a weak infusion of tobacco ; and he informs
me that the simple application of turpentine to the throat
externally is sufficient to kill the worms. It should be borne
in mind that the bird itself may be injuriously affected by these
drugs if they are carelessly employed.

Thirdly. The treatment recommended by Mr Montagu
proved successful in his hands, although the infested birds were
old partridges. One of his birds had died of suffocation ; but
he tells us that " change of food and change of place, together
with the infusion of rue and garlic, instead of plain water, to
drink, and chiefly hemp- seed, independent of the green vege-

446

PARASITES OF ANIMALS

tables which the grass plot of the menagerie afforded, recovered
the others in a very short time."

Fourthly. The plan I employed in my experiment. This is
only desirable in advanced cases, where suffocation is impend-
ing. It will afford instant relief, as the trachea may be cleared
of all parasitic obstructions.

Lastly. The essential point to be observed is the total de-
struction of the worms. This will help to put a stop to future
epizooties. If the parasites are merely killed and thrown
away carelessly, the eggs will sustain no injury. Decomposi-
tion having set in, the young embryos will sooner or later
escape their shells, migrate in the soil or elsewhere, and ulti-
mately find their way into the air-passages of birds in the same
manner as their parents did before them.

In this place I must not omit to mention the remarkable cir-
cumstance, quoted in my pamphlet on the grouse disease, that
Prof. Wyman, of Boston, found Eustrongyli surrounding the
cerebellum in seventeen out of nineteen snake-birds or water-
turkeys that had been shot in Florida. These viviparous
nematodes apparently occasion their avian bearers no inconveni-
ence. No doubt, as Wyman observes, their presence must be
regarded as a normal state of things : but should they occur in
excessive numbers, then we can hardly doubt the result.

As regards acanthocephalous parasites, although not so nume-
rous as the nematodes, it is extremely probable that they
play a similar role. Parasites which prove fatal to swine are
scarcely likely to be harmless in birds. On the 18th of
February, 1875, I received from Sir Joseph Hooker eleven
worms for identification. Mr Charles P. Hooker, his son,
subsequently informed me by letter that he had found them in
a Kedwing (Tnrdus iliacus) which he dissected in January, 1875.
The worms (EcJiinorhynchus transversus) occupied the large
intestine, probably to the number of one hundred in all.
Hitherto this parasite has been found abundantly in the black-
bird, thrush, and in most of the Turdidce ; but not in the red-
wing. It has also been obtained from the starling and red-
breast. The presence of so many of these armed parasites in
one small host could hardly fail to inflict severe injury on the
bearer.

In concluding this section of my work I can only find space
to make a few acknowledgments. Most of the rare, new, or
interesting avian entozoa which I have examined and described

AVES

447

have been received either from the Zoological Gardens, or from
personal friends. In pai-ticular I may mention the collections
sent to me by Mr Charles Darwin, Mr Eobert Swinhoe, Mr
Charles W. Devis, Dr Murie, Dr John Anderson, and Mr
Spooner Hart, of Calcutta. A great many correspondents have
contributed single specimens, many of which I have already
incidentally acknowledged in .these pages. In this place I
must particularise the new species {Ascaris Cornelyi) which I
described from specimens sent to Mr Sclater. This worm in-
fests the vulturine pintado {Numida vulturina). Mr Darwin's
collection contained fine examples of Filaria horrida from the
American ostrich {Rhea). When dissecting birds at the Zoolo-
gical Society's Menagerie, I obtained (in addition to the parasites
already mentioned) Distoma (equate from the American owl
(Strix perlata); Taenia multiformis from the night heron (Ardea
nyctocorax) ; T. infundibuliformis from a horned pheasant
(Phasianus) ; T. Icevis and T. lanceolata and also Ascaris
tribothrioides from a dusky duck {Anas obscura) ;
Fustrongylus papillosus (fig. 75) from the larus
crane {Qrus antigone); Trichosoma longicolle from
the horned pheasant, and T. brevicolle from the
Sandwich Island goose (Bemicla Sandruichensis) .
This bird was also infested by Spiroptera crassi-
cauda and Ascaris dispar. From the ring-necked
pheasant {Ph. torguatus) and from the black-backed
Kaleege {Euplocomus melanotics), and also from a
cheer pheasant {Ph. Wallichii), I obtained abund- P°a-
ance of Ascaris vesicularis. From the ashy -headed fie^'o^gS"1"
goose {Chloephaga poliocephala) examples of Str.
tubifex and Str. nodularis. This bird also yielded a new species
{Str. acuticudatus) . From a tinamou {Tinamus) I obtained the
Ascaris strongylina of Rudolphi {Str. spiculatus, mihi). From
amongst our British birds I have obtained Asc. deprossa, Tri-
chosoma falconum and Hemistoma spathulum, from the kite
{Falco milvus). Of these three worms, the last named was
also found in the long- eared owl {Strix otus), whilst the first
likewise occurred in a kestrel {Falco tinnunculus) and in a honey
buzzard {Pernis apivorus). I may add Filaria attenuata from
a peregrine {F. peregrinus) ; and F. leptoptera from a sparrow
hawk {Accipiter nisus). From the redshank {Totanus calidris)
I obtained Tcenia variabilis, and from the curlew {Numenius
arcuata) T. sphoerophora j and from various gulls {Larus glaucus

448

PARASITES OP ANIMALS

and L. tridactylus) the Tetrabothrium cylindraceum. Also
from the grey gull Echinostoma spinulosum. From the red-
throated diver I procured Tetr. macrocephalum. I found this
tapeworm also in the guillemot {Uria troile), together with a
nematode {Ascaris spiculigera) two examples of which were
lodged in the right auricle of the heart. From a capercaillie
(Tetrao urogallus) I have obtained a species of Ligula, and
likewise numerous examples of Trichosoma longicolle. Of
necessity, this brief notice only comprises a small part of the
avian entozoa contained in my collection, many of which I have
not had time to describe, whilst, as regards others, I can only
say that they remain in abeyance for examination and identifi-
cation.

Mr Brotherston has recently recorded an interesting find
(made Nov. 25th, 1874) of nematodes in the legs of the lesser
grebe (Podiceps minor), and also (Feb. 27th, 1878) in the water-
hen (Gallinula chloropus). Both limbs of both birds were in-
fested. The worms of the grebe were spirally coiled amongst
the muscles and tendons near the lower end of the tibia, and
when unrolled measured about an inch in length. The para-
sites of the waterhen were similar in appearance. Not impro-
bably these were all sexually-imperfect female examples of
Filaria acuta hitherto found in the abdomen of grebes.

The British Museum contains many interesting specimens
purchased from the collection of Yon Siebold, but they are
practically inaccessible to investigators. The entozoa in the
Hunterian Collection, though few in number, are in an excellent
state of preservation, and at all times accessible to visitors.

The ectozoa of birds are too numerous to be dealt with in
these pages. Eeferences to recent papers by Haller, Megnin,
and Westwood will be found below. The insects infesting the
domestic fowl have been alluded to in connection with the
occurrence of poultry-lousiness in the horse.

Bibliography (No. 57).— (Anonymous), "On the (gape)
Diseases of Fowls," the f Veterinarian/ p. 267, 1841. — (Anon.,
initialed " Q ."), Letter on the "Grouse Disease," in the ' Times/
Sept. 5, 1874. — (Anon.), "On Grouse and Partridge Disease,"
being annotations in the c Lancet/ Sept. 4, 1875, pp. 360 and
361. — (Anon.), "Grouse Disease," letter signed " R." (probably
from Lord Ravensworth), in ' Land and Water/ Aug. 16, 1873.

(Anon.), "Grouse Disease," article (from "W. C.") in the

'Field/ Aug. 2, 1873 (criticising my brochure— T. S. C). —

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449

Arhng, " Note on Tsenia from the Fowl/' ' Rec. Med. Vet./
1875. — Baird, W., "Descr. of Tania calva from the Grouse/'
in 'Brit. Mus. Catalogue/ p. 83. — Blavette, "Descr. of a
Verminous Disease among Fowls/' 'Veterinarian/ p. 649, 1840.
— Brotherston, A., " Parasitic Worms in Legs of Grebe (Podiceps
minor) and Water-hen (Gallinula chloropus)," ' Science Gossip'
for April, 1878, p. 88, and in ' Proceedings of the Berwickshire
Naturalists' Club/ vol. viii, p. 288. — Carter, B. (see Cobbold).
— Chapman, H. C, " Description of a new Tapeworm from
Khea/' ' Proceed. Phil. Acad./ 1876, p. U.—Chatin, J., "Etude
sur des helminthes nouveaux ou peu connus (treating of a
Cyathostoma, from Anas tadorna and Sclerostoma pelecani),"
' Annales des Sciences Naturelles/ 1875. — Cobbold, ' The Grouse
Disease, a statement of facts tending to prove the Parasitic
Origin of the Epidemic/ London, 1874. — Idem, Letter in the
' Field/ Sept. 9, 1872. — Idem, " Contributions totour Knowledge
of the Grouse Disease, with description of a new Species of
Entozoon," ' Veterinarian,' March, 1873 ; see also an article
(by Brudenell Carter) in the ' Times ' for Sept. 5, 1874; repr.
in the 'Veter./ Oct., 1874. — Idem, " Remarks on the Entozoa
of the Common Fowl and of Game-birds, especially in relation
to the Grouse Disease," the ' Field/ Sept. 14, 1867, and 'Brit.
Assoc. Rep./ 1867. — Idem, " On Sclerostoma and the Disease
it occasions in Birds/' 'Linn. Soc. Proc./ 1861 ; repr. in the
'Field/ June 22, 1861, and in ' Edin. Vet. Rev./ vol. iii,
p. 439, 1861, also in Tegetmeier's work on ' Poultry / see also
a comment on " Gapes," by " Umbra," in the ' Field/ June 29,
1861. — Idem, "Parasites from the Zoological Gardens," 'Intel-
lectual Observer/ 1862. — Idem, "Notes on Entozoa (species
Nos. 1, 7, 9, 12)," in ' Zool. Soc. Proc./ 1873-76.— Idem,
'Linn. Soc. Trans./ 1858. — Idem, "On Entozoa of Birds and
Fishes, collected by Mr Charles W. Devis," ' Zool. Soc. Proc./
1865. — Idem, "Note on Entozoa in the Crested Grebe," the
'Field/ March 29, 1873.—Colquhoun, W., 'Remarks on the
decrease of Grouse, and on the Grouse Disease (Gapes)/
Edinburgh, 1858 ; see also a notice in ' Edin. Vet. Rev./ vol. i,
April, 1859. — Crisp, E., " Note on Hydatids in an old Honduras
Turkey," ' Path. Soc. Trans./ 1863.— Idem, " On Sclerostoma,"
'Rep. of Path. Soc./ in 'Med. Times and Gaz./ Oct. 26, 1876,
p. 474. — Idem, " Note on Filaria in the Heart of a Peregrine
Falcon," ' Path. Soc. Trans./ 1854. — Davaine, ' Syngame de la
trachee/ p. 37, and 'Synops.' cxiv, in his ' Traite.'— Levis, C. W.

450

PARASITES OF ANIMALS

(see Cobbold). — Dicsing, ' Re vis. der Myzelminthen/ Abtheil.
" Tremat.," 1858. — Idem, " Tapeworm from Podiceps," in his
'Zwanzig Arten von Cephalocotyleen/ Wien, 1856. — Idem, see
various species in his ' Revisionen/ given in Bibl. No. 58. —
Fames, 0. J. L., " On Tapeworm in Blackbirds," letter to the
'Lancet/ June 9, 1877, p. 863.—Farquharson, B., "The
Grouse Disease (due to a contagious fever)," letter to the
' Lancet/ Sept., 1874. — Fergusson, J., " On Grouse Disease,"
letter in the ' Times/ July 16, 1878.— Gentles, T. W., "Tape-
worm in Birds," letter to the ' Lancet/ Jan. 18, 1868, p. 106.
— Haller, G., " Freyana und Picobia, zwei neue Milbengat-
tungen," in ' Sieb. u. Koll. Zeitsch./ 1877, s. 181. — Johnston,
D., " On the Grouse Disease," letter in the ' Lancet/ Sept. 20,
1873, p. 441. — Krabbe, 'Bidrag til Kundskab om Fuglenes
Beendelorme/ Copenhagen, 1869 (contains a summary in French,
' Recherches sur les Tenias des Oiseaux'). — Linstow, 0. von,
" Enthelminthologica " (containing descriptions and notes on
worms from birds, fishes, reptiles, and mollusks), ' Archiv fur
Naturgeschichte/ 1877. — Idem, "New Flukes (Dist. vitellatum
and D. macrojphallus) from Totanus hypoleucus, and. D. ccslebs
from Fringilla," Beobacht. in < Arch. f. Naturg., '1875, s. 189-
193. — Maclagan, "Note on the Grouse Disease," ' Proc. Roy.
Soc. of Edin./ April 20, 1874, p. 378,— Marion, " Revis. des
Nemat. du Golfe de Marseilles," ' Compt. Rendus/ 1875.—
Megnin, P., " On Harpirhynchus and other Mites," in ' Rev. f .
Thierheilk./ Oct., 1878, s. 146. — Idem, "Memoire sur les Chey-
letides Parasites (Picobia, &c.)," ' Journ. d'Anat. et de Physiol./
1878 ; see also 'Rev. f. Thierhielk./ Sept., 1878 et seq. — Moliu,
in his various monographs (quoted in Bibl. Nos. 54, 56, and
elsewhere). — Montagu, G., "Account of a species of Fasciola
which infests the Trachea of Poultry, with a mode of Cure,"
'Memoirs of the Wernerian Nat. Hist. Soc./ vol. i, p. 194,
1811. — Perrier, 'On Syngamus ' (brochure, Paris, 1875). —
Pulteney, B., " On Ascarides discovered in Pelicanus carbo and
P. cristatus," 'Linn. Trans./ vol. v, 1800, p. 24.— Boll, "Beitrag.
zur Entwickelungsgeschichte der Tsenien," ' Verh. d. Wiirzb.
p. m. Ges./ Bd. iii, 1852, s. 51. — Sanderson, J. B., " On Grouse
Disease," 'Brit. Med. Journ./ May 15, 1875.— Small, M.,
" Worms in the Eyes of Geese," from the ' Irish Farmer's Gaz./
in the 'Veterinarian/ 1862, p. 19.— Tait, L., " Tapeworm in
Birds," letters to the 'Lancet' for Jan. 25, 1868, p. 145, and
Feb. 8, 1868, p. 214; see also " Pediculus," ibid., p. 180.—

REFTILJA

451

Tegetmeier, " On the Grouse Disease," in the ' Field/ Sept. 12,

1874. — Thick, "Letter on the Cure of Gapes/' in 'Land and
Water/ Aug., 1867, p. 77. — Vaughan, "The Grouse Disease/'
in the ' Field/ Aug. 23, 1873. — Villow, A., "Sur les migrations,
et les Metamorphoses des Treuiatodes," ' Oomptes Eendus/

1875, and 'Ann. Nat. Hist./ 1875 (chiefly on flukes of the
sea-lark, Tringa alpina). — Idem, " Sur le syst. nerveau," &c.,
ibid., 1875. — Idem, " On the Helm. Fauna of the Coast of
Brittany (chiefly from birds)," 'Ann. Nat. Hist./ from ' Comp.
Eend./ 1875, p. 1098, and from 'Arch, de Zool. Experim. et
gen./ 1875. — Idem, "Sur Pappareil des Trematodes (Dist.
insigne)," ' Compt. Rend./ 1875. — Wedl, K., 'Anatomische Beo-
bachtungen ueber Trematoden/ Wien, 1858 (contains excellent
descriptions of numerous flukes, chiefly from birds) . — Westwood,
J. 0., "New Flea on a Fowl (Sarcopsillus)," 'Entom. Month.
Mag./ xi, p. 246, 1875. — Wiesenthal, A., "Account of a Parasite
infesting the Trachea of Fowls and Turkeys in America," ' Med.
and Phys. Journ./ vol. ii, p. 204, 1799. — Wyman, " On Eustron-
gyli within the Cranium of Water-Turkeys," ' Proc. Boston
Nat. Hist. Soc./ 1868. — Youatt, " On a Verminous Disease in
Poultry " (being a letter to him), ' Veterinarian/ p. 648, 1840.
— Zeller, " On Leucochloridium," from ' Zeitsch. f . wissensch.
Zool./ 1874, s. 564, and from ' Bibl. Univ. Bullet. Sci./ 1874,
p. 366, in 'Ann. Nat. Hist./ Feb., 1875.

Paet II (Reptilia).

Much that I have advanced in respect of the parasitism of
birds holds good in the case of reptiles. I cannot recapitulate.
The saurians, ophidians, and chelonians are extensively infested,
but in this respect the amphibian frogs, toads, and salamanders
are probably the most victimised. If, on the one hand, com-
paratively few tapeworms have been found in reptiles, it may,
on the other hand, be said that the Echinorhynchi come into
prominence, causing serious injury to reptilian hosts. Serpents
and chameleons are particularly liable to have their lungs
infested by acanthocephalous entozoa, these organs being also
attacked by pentastomes. I have received evidence of fatal
epizooty amongst chameleons from this source ; and I have been
requested to suggest a remedy. To prevent outbreaks of
entozoal disease is one thing j to offer a radical cure when the

452

PARASITES OF ANIMALS

parasites are firmly anchored within the pulmonary organs is
quite another nfatter.

As remarked in my ' Entozoa/ the trematodes display a great
partiality for batrachians, more than half a dozen different
species of fluke being known to infest the common frog. Flukes
are likewise tolerably abundant in the saurian and chelonian
reptiles. I regret that I cannot find space so much as to enu-
merate the species. As one would naturally expect, the frog
has been exhaustively anatomised and examined for entozoa,
and it was this creature that supplied Leuckart and Mecznikow
with the materials which led to their well-known discovery
and controversy respecting the development, dimorphism, and
parthenogenetic phenomena exhibited by Ascaris nigrovenosa.
I cannot give the facts in detail. Female examples of the worm
live in the lungs of the frog. Their young, as embryos, pass
into the damp earth and mud, where they grow up into
sexually- mature forms different from the parent worms found in
the frog. These free adult worms, male and female, produce
rhabditiform embryos which present characters of their own
and attain a certain stage of growth. At this stage they
are conveyed into the lungs of the frog where they arrive at
sexual maturity. As there are no male worms in the frog, it is
probable that the embryos of these parasitic females are agamo-
genetically produced by internal budding, the sexual influence
of the free males being, as it were, continued onward without
actual contact with the parasitic females. Amongst the
interesting parasites of the frog one must also mention Amphi-
stoma subclavaium and Pohjstoma inter ■gerrimum. The former
worm resides in the large intestine and the latter in the urinary
bladder. The larvse (Oercaria diplocotylea) of this amphistome
reside in or upon the body of water-snails, and, like the
cercarian larvae of polystoma, they are furnished with eyes. I
state this fact on the authority of Pagenstecher j and, since I
cannot devote a special section to the entozoa of mollnsks,
I repeat, in part, the valuable results which Pagenstecher
published many years back and which have a permanent value
in relation to the origin of parasitic diseases resulting from
flukes. In the memoir quoted below, Pagenstecher gives the
following conclusions (Schlussbcmerfruv gen) :

« (a). — The eggs of the trematoda vary in respect of size,
form, and color, being either furnished or not with a lid, and
accordingly distinguishable. In the mature condition they

EEPTILIA

453

contain a ciliated or a non-ciliated embryo of unequal growth,
this embryo partly increasing in size even after its birth. In
various conceivable ways the eggs themselves, or the embryos
which have quitted their shells, arrive in and upon the bodies of
mollusks, where they are consequently found. In this situation
the egg opens, or the ciliated covering decays, and the contained
motionless germ — which in itself offers no distinctive charac-
ters— having become free, grows into a nurse, or forms several
nurses within itself.

"(b). — Whilst some of the trematodes display a highly
organised nurse condition, others exhibit only a simple kind of
germ-sac. Both, forms, nevertheless, appear to occur in one
and the same species, probably depending upon external causes.

" (c). — The organised nurses (or rediae, as they are termed)
have a mouth, and a strongly marked muscular oesophagus,
which is continued into a short or prolonged, single, blind
intestine, or the latter may be double. The expulsion of
animals developed within them I have only seen to take place
through an opening at the hinder extremity. Old rediae lose
their structure. I did not observe any vascular system. Tailed
trematode larvas (Cercariae), as well as rediae themselves, are
developed within the rediae, this variation of nurse-contents
probably depending on the season.

" (d). — No independent new germ-sacs are developed within
the simple unorganised germ-sacs (sporocysts), and only such
trematode larvae as are capable of arriving at sexual maturity
are furnished with special appendages.

" (e). — When the immature contents of both, nurse forms
(i.e. of sporocysts and rediae) are accidentally set free, and are
situated within the organs of nutrition of the living host, then
they appear prepared to develop themselves anew into nurse
forms ; and, moreover, cercariae whose development has not yet
attained a definite stage — and even their tails also — appear to
enjoy a similar capacity. Some nurses are likewise capable of
multiplication by division and budding.

" (/). — Some germ-sacs have the property of developing
within themselves cercaria-like larvas — which are different from
the true cercariae — from whose body the development of a
(Mstoma may take place, while their single or double tail-like
appendages in all cases develop anew into germ-sacs. To this
class belong Bucephalus and Distoma duplicatum.

" (</). — All the cercaria at present known are destitute of

454

PARASITES OP ANIMALS

eyes, but other forms of trematode larvae are furnished with
visual organs. Accordingly, I never found eyes in young
distomata whilst they were in their last dwelling-place, but eyes
are certainly present in the young forms of Polystoma and
Amphistoma. The supposition that a spontaneous wandering
is associated with eyes is not yet confirmed in my experience.

" (h). — As a means of distinguishing the different forms of
Cercariae, amongst other indications, their places of dwelling
may be useful, because each mollusk only harbors a limited
number of species. Notwithstanding, Professor Filippi is in
error if he believes that every species of mollusk carries only a
single armed form of cercaria. A migration of the cercaria is
indispensable to its perfection.

" (i). — Many larval trematodes form cysts round themselves,
probably by means of a special organ of secretion, and also by
the epidermis. Their future destiny necessitates this. The
sporocysts apparently fulfil towards the larvae, which are deve-
loped within them, a similar purpose, namely, a protection
against the stomachal digestion of the new host. In the pupa
condition the development of the larva, which has now thrown
off the tail, makes greater or less progress, according as to
whether it is surrounded by nourishment or not. In particular,
while in this stage, the different kinds of hooks for migratory
purposes make their appearance, always, without doubt, after
the shedding of the skin. Other trematodes pass through this
tail-less sexually-immature stage without any cyst. I have not
yet seen any larval trematode forms which had been produced
in sporocysts Or rediae without appendages ; they appear to
occur, nevertheless.

« — As the larv8e exist only in a few hosts — and most of
them dwell only in one species of animal — so, also, the con-
tinued progress towards sexual maturity only succeeds in the
case of certain well-defined larval organisms, but the digestion
of the cysts and liberation of the larvae may be accomplished in
various animals.

<< fym — The armed Cercariae appear to be larvae of the spine-
covered distomes of amphibia; for, as examples, the Oercaria
ornata becomes transformed into JDistoma clavigerum, and G.
armatd into Distoma endolobum ; the Dist. duplicatum and
Oerc. diplocotylea are, apparently, the juvenile forms of Dist.
cygnoides and Amphistoma subclavatum. The Dist. echiniferum
of Paludina could neither be advanced in development in the

EEPTILIA

455

frog or duck, nor could all the other larvae which I subjected to
experiment be developed either in the green or brown frogs.

" (m) . — When young trematodes arrive at the right place for
their maturation, then the male generative structures develop
before the female organs, and in the subsequent excess of egg
production the form and structure of the animal becomes
obliterated.

" (n). — The yelk-molecules surrounding the germinal vesicle
are not directly transformed into an embryo/''

As regards the acanthocephalous parasites of reptiles, I may
observe that Echinorhynchus anthuris is very common in the
lesser water newt (Lissotriton punctatus) . In the accompany-
ing illustration (Fig. 76) I have represented the free ovarian

Fig. 76. —Echinorhynckus anthuris. 1, Attached to the intestine ; 2, specimen enlarged •
3, ovarian vesicle, including germs ; 4, germs in various stages ; 5, vesicle with germs
more advanced; 6, 7, eggs in their capsules; 8, free egg. Original.

egg-bearing bodies, the development of the ovum, and the
adult worms. For anatomical details, however, I must refer to
my earlier treatise (' Entozoa/ p. 100 et seq.).

Amongst the species of entozoa that were found by me at the
Zoological Society's Menagerie I may mention Distoma coro-
narium and Ascaris lineata, from the intestines of Alligator
mississippiensis ; Dist. Boscii, from an American snake {Coluber);
an immature nematode, from the heart of Coluber Blumen-
bachii ; and Echinorhynckus inflexus, attached to the intestines of
a snapping turtle {Chelydra serpentina). I may add that the

450

PARASITES OF ANIMALS

lungs of the alligator also contained examples of Diesing's
Pentastoma oxycephalum. An Egyptian hooded snake (Naia
hage), which died at the Zoological Gardens in 1859, furnished a
new species of pentastome (P. multicinctum) . Dr George Harley
described and anatomised this worm with remarkable care.
To Harley's memoir Prof. Leuckart did ample justice in his
work on the ' Pentastoma/ Several new species of reptilian
entozoa have recently been described by Dr von Linstow ; and
Dr Solger has found a new trichosome (T. recurvum) beneath the
skin of a young crocodile (probably Croc, acutus). For further
particulars I refer to the revised descriptions and additions
by Diesing, Molin, Schneider, and other systematists.

Bibliography (No. 58). — Baird, W., "Description of a new
Entozoon from the Diamond Snake/' in ' Proceed. Zool. Soc'
for 1865, p. 58, and in ' Ann. Nat. Hist/ for July, 1865, p. 52.
— Blanchard, " On Polystoma," ' Ann. des Sci. Nat./ 3e ser.,
viii, p. 331. — Canton, LJ., " An account of some Parasites
attached to the Conjunctivae of the Turtle's Eyes/' ' Quart.
Journ. Micr. Sci./ and ' Dublin Med. Press/ 1861 (with remarks
by myself). — Colloid, 'Notes/ &c. (1. c, Bibl. No. 57), and
in ' Linn. Trans./ 1857. — Crisp, U., " Note on Cysticerci and
Trichocephali from an Alligator/' ' Path. Soc. Trans./ 1854. —
Diesing, ' Revision der Cercarieen/ 1858; ' Revis. der Myzel-
minthen/ 1858; 'Nachtrage (u. s. w.)/ 1859; 'Revis. der
Nematoden/ 1860; 'Revis. d. Turbellarien/ 1861; 'Revis. d.
Cephalocotyleen/ 1863. — Idem, ' Monographie d. Gatt. Amphis-
toma und Diplodiscus/ and ' Nachtrage zur Monog. der Amph./
1839— Dujardin, 'Hist. d. Heltn.' (1. c, pp. 320, 526, &c.).—
LJlerth, " On Myoryldes Weismanni from the Muscles of the
Frog," trans, by Busk, from ' Siebold und Kolliker's Zeit-
schrift/ in ' Lond. Micr. Journ./ Jan., 1864. — Gastaldi, ' Cenni
sopra alcuni nuovi Elmint./ Torino, 1854 (new flukes from
frogs and salamanders). — Gliige, "On Entozoa in the Vessels of
Frogs/' from ' Oomptes Rendus/ in 'Micr. Journ. and Struct.
Rec./ p. 207, 1842 ; see Griibe and Valentin. — Qr&be, "On the
Entozoa of the Frog, and on the Pathology of that Batrachian/'
from ' Comptes Rendus/ in ' Micr. Journ. and Struct. Rec./
p. 246, for 1842 ; see also Mandl. — Harley, G., " On the Anatomy
of a new Species of Pentastoma found in the Lung and Air-sac
of an Egyptian Cobra," ' Proc. Zool. Soc./ June, part xxv,
p. 115, 1857. — Leuckart, ' Bau und Entwickelungsgeschichte
der Pcntastomen/ Leipsig, 1860. — Linstow, ' Enthelminth.' (1. c,

PISCES

457

Bibl. No. 57). — Macalister, A., "On the presence of certain
Secreting Organs in Nematoidea," ' Ann. and Mag. of Nat.
Hist.' for 1865. — Idem, "On the Anatomy of Ascaris dactyluris,"
' Proc. Nat. Hist. Soc. of Dublin/ vol. iv, 1865. — Mandl, "De-
velopment of Entozoa (Ascaris nigrovenosa of the frog)," from
' Eep. of French Acad, of Sci./ in ' Month. Journ. of Med. Sci./
vol. ii, p. 1081, 1842. — Molin, ' Monog. del gen. Myzelminth ;
Mon. del gen. Physaloptera ; Mon. del gen. Histiocephalus ;
Mon. del gen. Spiroptera ; ' Wien, 1859-60. — Pagenstech&r, 'Tre-
matodenlarven und Trematoden/ Heidelberg, 1857. — Sibbald,
J., " On the Nematoideum natricis," 'Path. Soc. Trans./ vol. viii,
1857. — Solger, " Ueber eine neue species von Trichosoma/-'
'Arch. f. Naturg./ 1877. — Valentin, "On Parasites in the
Bladder of the Frog" (from ' Eepertorium '), in ' Micr. Journ.
and Struct. Eecord/ 1842, p. 183.— Vogt, C, " On Filaria in
the Vessels of the Frog," from ' Miiller's Archiv/ in ( Micr.
Journ. and Struct. Eec./ p. 241, 1842. — Wedl, F., " Beitrage
zur Lehre von den Haematozoen," ' Sitzungsb. Akad./ Wien,
1850 (from the blood -of frogs, &c.).—Zeller, E., " Weiterer
Beitrag zur Kentniss der Polystomen/'' ' Sieb. und Koll. Zeit-
schrift/ 1875.

Paet III (Pisces).

Swarms of entozoa infest fishes, and it is hard to say
whether they are less numerous in the inhabitants of fresh
water than in those of salt water. More attention has been
paid to the helminths of the fishes than to the internal parasites
of birds and reptiles, consequently, the number of known
species must be estimated by many hundreds. In like manner
a great deal has been written respecting the ectozoa of fishes.
These parasites, often called fish -lice, belong chiefly to the
haustellated crustaceans, and are better known by the title of
Epizoa. No account of them can be afforded in this treatise,
but some trifling notice of the literature of the subject will be
given below.

About a hundred distinct species of fluke have been described
as infesting fishes. Not many of these worms possess more
than a zoological interest ; nevertheless, from that point of view
certain types are very curious. Most of the species dwell in
the stomach and intestines, but, as more or less remarkable
exceptions, I may mention Distoma seriale infesting the kidney

458

PARASITES OP ANIMALS

of Salmo umbla, D. longum (Leidy) from the pharynx of Esox
cstor, D. polymorphum from the urinary bladder of the common
pike (Esox lucius), D. obesum from the gall bladder of iSalminius
and other Brazilian fishes (Xiphostoma, Leporinus), D. tomatum
attached to the gills of Ooryphcena hippuris, D. rosaceum attached
to the palate of Lota communis, and D. contortum attached to
the gills of Orthagoriscns mola. Most of the forms found
encysted are sexually-immature worms. To these belong
D. annuligerum, found by Nordmann in cysts in the vitreous
humour of the eye of the perch (Perca fluviatilis), and D. embryo
from the liver and peritoneum of Acerina vulgaris. One of the
largest and most remai'kable of the flukes inhabiting marine
fishes is the Distoma clavatum, found by Tilesius in the stomach
of Pelamys, by Pohl in Thynnus, and by Bosc in Coryphcena.
In the last-named fish it has been found adhering to the gills,
in the liver, and in the intestines. In August, 1865, 1 obtained
this parasite from a sword-fish (Xiphias gladius), and in the
same piscine host I also found examples of four other species
of helminths (Tetrarhynchus attenuatus, scolex of another tetra-
rhynch, Bothriocephalus ptMcatus, Ascaris incurva). Believing
Distoma clavatum to represent several forms hitherto regarded as
distinct, I append a few particulars respecting it. Five examples
of this worm were obtained by me from the stomach of a sword-
fish. Generally they varied in length from four lines to two
inches. They differed somewhat in shape, but all had the so-
called head and neck directed backwards. Below the ventral
sucker the two largest specimens were distended with eggs
and black pigment. All of them likewise exhibited more or
less well-marked transverse rugae, the last ring surrounding
an orifice which represented the outlet of a large contractile
vesicle. The eggs averaged J^" in length.

When revising the entozoa of the Museum of the Royal
College of Surgeons I encountered many parasites without
labels attached. Amongst these were several flukes, which,
though differing from each other in size and shape, appeared to
be identical. One of these specimens turned out to be the
particular Distoma clavatum described and figured by Professor
Owen in the ' Zoological Society's Transactions/ Several of
the others I made out to be part of a series contributed by Mr
George Bennett, who also gave specimens to the British
Museum, but the College Museum stores contained yet a third
group of specimens of uncertain history. The large fluke

PISCES

459

described by Prof. Owen was formerly in the collection of the
Kev. Lansdown Guilding. In Dr Baird's catalogue the specimens
presented by Mr Bennett are stated to have come from the
stomach of a bonito, and probably Mr Guilding's specimens
may be referred to the same " host." Be that as it may, the
specimens differ from each other in a very striking manner.
In the year 1730 M. Garsin first described this worm under the
generic title of Eirudinella. He says : — " Cet insecte tire de
Testomac de la Bonite ne «vecut qu'environ deux heures.
Expose a Tair il etoit languissant, et reprenoit de la vivacite
dans de Feau de mer. II diminua sensiblement de volume
pendant qu'il vivoit encore." M. Garsin's description is accom-
panied by three figures. His specimens do not appear to have
exceeded \\" in length. In 1774 Pallas described a trematode
{Fasciola ventricosa). It measured two inches in length. All
that he says regarding its source is as follows : — " Ex Amboyna
missum fuit singulare hoc molluscum, quod ad aliud quam Fas-
ciolarum genus referre non potui, in quo quasi gigas erit." He
remarks upon its pale white color, and notices particularly the
soft elastic body proper, which when wounded gave out a dark
matter resembling soot. This material, when examined with
the microscope, appeared fresh ; it was not the result of decom-
position. Pallas also gives many other details, accompanied by
a figure. In 1790 Menzies likewise described and figured a
fluke about two inches long. He calls it Fasciola clavata : —
" It is of whitish color, somewhat pellucid, discharging at its
mouth a black-colored fluid, which can easily be perceived
through its body. I have often found it," he adds, " in the
maws of the bonito, between the tropics in the Pacific Ocean."
Notwithstanding the similarity of description, Menzies does not
appear to have recognised the identity of his worm with that
described by Pallas. Prof. Owen, however, subsequently estab-
lished this identity, and referred to this species as the Fasciola
clavata seu ventricosa. On the other hand, the British Museum
Catalogue represents Pallas' s worm as specifically distinct from
that of Menzies, but as identical with the specimen described
by Prof. Owen from Mr Guilding' s collection.

In 1802 Bosc described and figured a trematode under the
title of Fasciola fusca. This he obtained from the intestines
of a dorado. In form it differs considerably from the fore-
going species. Bosc's description runs as follows : — " Brune,
la partie posterieure tres-renflee, presque ovale, la partie

460

PARASITES OF ANIMALS

anterieure mince, cylindrique, inegale, aveo deux petits tenta-
cules en dessoas. Le sucoir do 1'anus tres grand." Bosc
recognised the identity of this worm with the Distoma coryphena
of Rudolphi, and systematists generally have adopted his
synonymy. In the British Museum Catalogue the Fasciola
fusca and F. ventricosa of Pallas are regarded as one and the
same species. The existence of two small tentacles is certainly
peculiar.

In 1827 Nardo obtained two very large flukes from the
stomach of a fish captured in the Gulf of Venice during the
month of September. He calls the fish Frostostegus prototypus,
which appears to be the same as the Luvarus imperialis of
Pafinesque. One of the parasites, being five inches in length,
he named Distoma gigas. His description is as follows : —
"Distoma teres, rubrum, retractile; poro ventrali minimo cujus
apertura magna, rotunda, ciliata ; poro antico terminali, parvo ;
collo brevi, retrorsum divergente, extensili, apice angusto, basi
lato ; cauda longa, postice incrassata et in apice obtusa oscula
donata." The alleged ciliated character of the ventral sucker,
was perhaps due to a wrinkled state of the lip. Apart from
this character, I see no reason for supposing this parasite to be
distinct from the Distoma clavatum procured by Mr Guilding,
or the Fasciola ventricosa described by Pallas. The intestines
of the fish harbored another parasite (D. Raynerianum) .
Unfortunately, Nardo gives no figure of Distoma gigas. It is
the longest fluke known to science.

In the year 1835 Professor Owen communicated the memoirs
already alluded to. In his paper he discussed questions relating
to the structure of Distoma clavatum, and threw much light
upon its anatomy, but I believe that the large " lateral cavities "
described by Owen are neither more or less than the somewhat
unusually distended alimentary cseca.

In 1845 Dujardin placed the worm with the true distomes,
yet, at the same time, expressed grave doubts as to whether it
were, in any sense, a fluke. " Ce ver," he remarks, " n'est
certainement pas un distome ni meme un trematode. Si sa
forme exterieure et ses deux oscules lui donnent quelque resseni-
blance avec les distomes, sa structure musculeuse la rapproche
davantage des Gordius, et son tegument ressemble a celui des
siponcles." M. Dujardin examined the specimens preserved
in the Paris Museum, and with regard to one particular
example, described as " Fasciola, trouve dans la mer de Nice/'

PISCES

461

he says, it presents " une certaine analogie avec le pretendu
Distoma clavatum." Dujardin himself was somewhat puzzled
by the resemblance in question. He does not appear to have
examined fresh specimens, yet he mentions the species as
tolerably common in the bonito, and occasionally present in the
tunny. At all events, it appears that the rightly so-called
Distoma clavatum is not unfrequently
taken from the ocean in the free
state. In concluding my notice of
this remarkable worm I can only add
that after examining numerous specie
mens both in the fresh and preserved
states, I have formed the opinion that
the following specific names all refer
to one and the same parasite : —
Distoma clavatum, Rudolphi ; D. cory-
phcence, Eud. ; D. gigas, Nardo ; Fas-
ciola clavata, Menzies ; F. coryphcence,
Bosc; F. coryph. hippuridis and F.
Scombri pelamidis, Tilesius ; F.fusca,
Bosc; Hirudinella marina, Garsin ;
H. clavata, Baird. In this list of syno-
nyms we may probably also include
Rudolphi's Distoma tornatum.

In addition to these distomes there
are numerous piscine flukes which
may fairly be relegated to other
genera. Thus, provisionally, I ele-
vated Dujardin's sub-genus Echino-
stoma into a separate genus ; and on
what I considered sufficient grounds
I established several other new genera
from amongst the more curious flukes
that had been described as infesting
fishes (Wedlia, Kolliheria). In the
genus Echinostoma the oral sucker is
either surrounded by a circle of little
spines, or it occupies the centre of a
disk, which is cleft at the ventral or
anterior aspect. In the latter case
the disk is either bordered both laterally and above by spines,
or there are two large lobed appendages, whose margins are

F;g. 77. — Echinostoma Mspidum.
Magnified. Original.

462

PARASITES OF ANIMALS

furnished with spines. In other respects this genus nearly
corresponds with the distomes, the simple digestive tubes
bifurcating immediately below the oesophageal bulb. The speci-
men of JEch. hispiclum here drawn (Fig. 77) was taken by me from
the spiral intestine of a sturgeon (1855), in which fish it occurs
very abundantly. The figure represents a back view of the

head and a lateral view of the body,
the neck having been slightly twisted.
The ventral sucker is concealed, but
the transparency of the skin permits
a view of the internal organs.

Another remarkable genus, estab-
lished by Von Siebold, is Gastero-
stoma. In this genus the ventral
sucker has taken the position usually
assigned to the oral opening ; the
latter being near the centre of the
body. The digestive caeca also dis-
appear, leaving only a short sto-
machal cavity, which reminds one of
the same viscus in imperfectly or-
ganised sporocysts or redise. When
G. gracilescens first came under my
observation I followed Rudolphi in
describing it as a distome (_D. graci-
lescens). The anatomy of the genus
has been illustrated by Yon Siebold ;
from whose observations also it may
be inferred that the larvae are various
forms of Bucephali. Prof. Molin de-
scribes the water- vascular or respi-
ratory apparatus as consisting (in G.
fimbriatum) of a broad central tube,
occupying the entire length of the
body and opening externally at the tail .

Amongst the more remarkable
fluke-types may be mentioned Van
Beneden's Nematobothrium (N. fila-
rina), occupying the branchial cavity
of Sciana aquila, also Holostoma
clavus, found by Molin in the intes-
tines of Gadus merlucius, also KdlWc&ria filicollis, occupying

Fig. 78 — Oasterosfoma grdciUscetu.
Maguilied. Original.

PISCES

463

open follicles in the branchial cavity of Brama Baii. The sexes
in the last-named genus are distinct ; male and female worms
together occupying each cyst. The genus Monostoma is also
largely represented amongst fishes. Prof. Wedl found a species
(Jkf. Wedlii) occupying follicles in the intestinal mucous mem-
brane, and also adhering to the fin rays of Rhombus lavis. I
also found a species (M. dubium) in a cyst attached to the
ovary of Gasterosteus spinachii. Several species of amphisto-
matoid worms were found by Natterer in Brazilian fishes
{Cataphr actus, &c), some of these representing distinct genera
(Aspidocotylus, Notocotylus) , to which I found Sonsino's remark-
able fluke (Gastrodiscus Sonsinonis, mihi) from the horse to be
very closely allied. In this connection must also be men-
tioned Griibe and Wagener's curious Amphiptyches urna, found
attached to the branchiae, and also in the intestines of Chimtera
monstrosa.

In addition to the above families and genera of digenetic flukes
infesting fishes we have the monogenetic tristomes and poly-
stomes. As remarked in my introductory treatise, the Tristo-
midas display a leech-like aspect, in consequence of which they
have been placed either along with the Malacobdellidce, or in
some other allied family of the suctorial annelids. The tris-
tomes are not strictly entozoa, yet their internal organisation
conforms more to the Trematoda than to the Hirudinidce. Thus,
they support two small suckers anteriorly and one large sucker
posteriorly, the body being externally smooth and devoid of
annulations. The tristomes have therefore no anus. In some
species the large caudal sucker is sessile, in others it is stalked
or pedunculated, being in either case bordered by a membranous
fold (Dujardin). All the species are hermaphroditic. They
attach themselves to the gills of fishes or to the general surface,
selecting especially the neighbourhood of the fins. Some species
are parasitic on crustacean parasites that are themselves attached
to marine fishes. In the genus Udonella the mode of develop-
ment is known to be simple and direct. According to Yan
Beneden, the embryos are large and acquire the form and
characters of their parents whilst they are still within the egg-
shell. They are ready to assume an independent existence the
moment they quit the shell. The eggs are oval, the chorion
being prolonged into a single filamentary process or " holdfast/'
Van Beneden compares a group of them to a " bouquet of vorti-
cells." On quitting the shell the embryonic Udonella at once

464

PARASITES OF ANIMALS

attaches itself to the Caligus, and there acquires the adult con-
dition. The Polystomidce comprise a variety of remarkable
genera. T accept this family as the equivalent of Dujardin's
first group of trematodes which he termed " Onchobnthriens,"
rejecting only his genus Diporpa, which is a juvenile condition
of Diplozoon. In this family Van Beneden includes the genera
Calceostoma and Gyrodactylus. In all the polystomes we have
a more or less ramified intestine, but the reproductive organs
conform to the general trematode type. All are hermaphroditic,
the eggs being supplied with filamentary appendages, in some
only at one pole of the shell, in others at both ends. The
water-vascular system is conspicuously developed. All the
species are supplied with prehensile hooks.

In the Diporpa condition of Diplozoon there are two super-
numerary hooks, associated with a dorsal sucker at the centre of
the body, and it is by means of these organs that a conjugation
between two such juvenile forms is effected. These two indi-
viduals become organically united for life, after the fashion of
the Siamese twins. After conjugation the sexual organs appear.
In Onchotyle appendiculata the lower end of the body merges
into a curious appendage, which is placed almost at a right
angle with the body itself, and in this way, as Van Beneden
justly remarks, the entire animal resembles a little hammer,
the resemblance being very much heightened by the circumstance
that one end of the appendage is cleft so as to correspond, as it
were, with the notch which we employ in the action of nail- draw-
ing. The Onchotyle appendiculata was first discovered by Kuhn
attached to the gills of a dog-fish (Scillium catulus), but it has
since been found ectoparasitically lodged upon other marine
fishes. With the Gyrodactylidce I include Van Beneden's genus
Calceostoma. The gyrodactyles have been classed with the
Polystomidee. Amongst the characters standing out most
prominently are those having reference to peculiar hooks which
project from the great sucking disk. In Calceostoma this
mechanism is reduced to a single horny structure placed at the
margin of the caudal sucker in the central line. In some
Gyrodactyli the hooks are very numerous. In Gyrodactylus
elegans the caudal sucker supports a pair of large laterally-
curved hooks, which are placed back to back in the centre of
the disk, being connected at their upper ends by a supple-
mentary semi-lunar bar. A series of tentacles serve to increase
the prehensile action of the sucker. In many species the males

PISCES

465

are supplied with accessory horny developments. The genus
Gyrodactylus has been studied by Nordmann, Yon Siebold, Gr.
Wagener, Van Beneden, and especially by Wedl, who records
the following results : — {a.) " Gyrodactylus is found on the gills
of fresh- water fishes under numerous specific forms, G. elegans
being also found by Creplin and
Siebold on the fins. Moreover, as
I have found nearly every species
of fish supporting a particular gyro-
dactyle representative, it would seem
that each finny creature supplies its
own Gyrodactylus. Sometimes two
of them are parasitic upon the same
gill, being frequently associated with
Trichodince, as well as with the still
unintelligible Psorospermice. (&.)
The clasping apparatus at the pos-
terior end of the body must — in an
animal so soft and constantly ex-
posed to the passage of regular
currents — be comparatively strongly
developed and accommodated to the
peculiar dwelling-places, and pro-
bably the varying character of the
latter supplies a reason why there
should be so great a difference in
the mechanism of the hooks belong-
ing to the disk, (c.) The hooked
apparatus affords a very valuable
and mathematically precise means
of diagnosis in the determination

of Species. This differentiation Fig. 79.— Gyrodactylus eUgans, containing
Z t n t -i • an eml)ry°- a> a> (Esophagus ; a, testis ;

may be accomplished fc>v observing" h, h, sucker ; i, i, huge, hooks ; spines.

, r P , Magnified. After Van Beneden.

whether there are two or tour large

hooks ; whether there be one or two connecting portions, and
by noticing their several forms and relations to one another ;
and whether, again, there are hooklets or not, remarking in the
first instance their position, form, distribution, and so forth.
(d.) The integument is sometimes wrinkled transversely, at
other times appearing to be smooth. (e.) The muscular
apparatus is, in certain cases, very strongly developed. In the
majority of instances special muscles are inserted into the

466

PARASITES OF ANIMALS

handles of the hooks, and they are also very frequently directed
into the transverse muscles of the skin. In Gyrodactylus cras-
musculus we find a protrusor penis and retractor palp arum
medius. (/.) Except in the case of G. ehgans, four so-called
eye-spots are observed at the anterior extremity of all Gyro-
dactyli. As Siebold says, they answer the purpose of light-
refracting organs. The palpi, which in G. crassiusculus are
seen to contain muscular bundles, appear to be retractile touch-
organs, extending more or less prominently forward. (g.)
Observations in regard to the alimentary canal are at present
incomplete, for only in the case of G. cochlea did I find a
single gullet demonstrable, (h.) Gyrodactylus becomes sexu-
ally developed, and cannot be regarded merely as a kind of
' nurse/ "

So much for Wedl, whose views I have elsewhere recorded
at great length. The genetic relations subsisting amongst the
Gyrodactyles have given rise to much controversy. Observing
the singular mode of reproduction in G. elegans, Yon Siebold
arrived at the conclusion that Gyrodactyles in general were
only nurse-forms of some higher organism, and he pointed
out, with undeniable accuracy, all the birth-stages of the young
one as it apparently pullulated within the parent and sub-
sequently emerged an almost perfect Gyrodactyle. Von Siebold
also remarked that the so-called " daughter," at the time of
birth, nearly equalled the " parent " in respect of size, whilst,
moreover, it contained within its interior another very young
Gyrodactyle, or, in other words, a " grand- daughter." Van
Beneden interpreted these facts very differently. I have myself
noticed the second generation, or daughter, to contain in its
interior evidences of a third generation. This I observed in
specimens obtained from the tails of Gasterostei caught in the
Serpentine, Begent's Park. Indications of the third progeny
were seen whilst the daughter still resided within the body of
the nurse-parent, and the so-called grand-daughter became
much larger immediately after birth. In one instance the
" daughter " commenced showing herself by a slight bulging
at the centre of the parent's body, whilst the integument of
the latter yielded on all sides of the bud-like projection, and in
such a manner as to convey the idea of a vaginal opening.
There was an evident struggle on the part of the young one to
free itself from the so-called parent envelope, but the tissues
showed no signs of injury. On partial protrusion it was seen

PISCES

467

that the budding portion corresponded with the centre of the
daughter's body, and this, in a little while, assumed the aspect
of a semicircular band. Subsequently the upper end became
detached, the freed extremity being now recognised as the
head. An interval elapsed before the broad posterior end of
the animal could be disengaged, but immediately after this was
effected the sides of the parent envelope closed in upon the
opening, and all that remained was a small cavity or sac,
indicating the position recently occupied by the daughter.
Altogether the process occupied about five minutes. I care-
fully compared the so-called " parent " with the " daughter,"
but in regard to size I can scarcely say which was the larger of
the two. As before hinted, Yan Beneden demurs altogether to
Von Siebold's views. He does not admit the parent to be a
kind of " nurse," he does not consider the primary young one
to be a " daughter," and, consequently, he does not regard the
embryo seen within the latter as a " grand-daughter." Van
Beneden says : — " According to our researches there is here a
false interpretation; the little daughter is lodged within the side
of its pretended mother, and not in its interior ; instead of
being its mother, it is its sister ; there is a difference of shape
because there is a difference of age ; the Gyrodactyles are vivi-
parous, and as among the Trematodes the eggs are formed one
by one, one embryo is scarcely formed when another commences
its evolution, and the egg-deposition is effected even whilst the
embryo is being produced. The Gyrodactyles are therefore
viviparous worms, which beget a single embryo at a time, as
those of the trematode group, to which they are allied, beget a
single egg at a time, and before the first embryo is expelled
another is already partly developed. There, we believe, lies the
correct interpretation of that phenomenon ; instead of a bud it
is an embryo, which has escaped from an egg. Here, therefore,
we have no phemonenon of alternate generation or of digenesis,
as Von Siebold supposes, but a simple viviparous reproduction."

Passing on to notice the cestodes of fishes, I may remark
that they often display characters very distinctive from those
inhabiting birds and mammals, being commonly furnished with
special tentacular hook-appendages employed as supplementary
organs of boring and anchorage. In the cartilaginous sharks
and rays these cestodes are remarkably abundant, and in certain
osseous species they are scarcely less frequent. The only
noteworthy kinds of fish which are commonly free from the

468

r AHA SITES OF ANIMALS

invasion of tapeworms are the sturgeons, blennies, gobios,
mullets, sparoids, and ScieenoB. Some few of them are infested
by Ligulce, Caryophyllcei, &c. Cuttle fishes harbor a great
variety of tapeworm-larvaa, forming one of the chief sources
whence sharks and rays obtain the same parasites destined to
arrive at sexual maturity within their own bodies.

Among the most interesting cestodes of fishes we may reckon
the pit-headed tapeworms and their allies (Bothriocephalidae).
One of the most common species is Both, proboscicleus which

Fig. 80.— Section of the strobile of Bolliriocephahis prohoscideus. Magnified. After Busk.

is found, often in considerable numbers, lodged within
the pyloric appendages of the' salmon (Salmo salar and 8.
hucho). It acquires a length of two feet. "When in large
numbers it cannot fail to prove injurious to the bearer. In this
connection also must be mentioned B. ilodosus. In the adult
state this worm infests a great variety of water-birds (herons,

PISCES

469

gulls, and divers), but in the young or sexually-immature
txenioid condition it is a frequent inhabitant of sticklebacks
(Gastereosteus aculeatus and O. pungitus), being also found in
the salmon and in the bull-head, or father-lasher (Oottus
scorpio). The immature tapeworm was formerly considered a
separate species (B. solidus). Some years back Creplin dis-
covered the connection subsisting between the two forms, and
re-described the species in its two conditions under the name
of Schistocephahos dimorphus, but it was reserved for Von
Siebold to explain the full nature of this relationship. In his
essay on " Tape and Cystic Worms " he shows that it is not
until the worm reaches the intestine of the ultimate host that
its segments acquire sexual completeness. As Yon Siebold
observes, "the extent of development in each individual will
be found to be in proportion to the time the parasite has passed
in the bird's alimentary canal after its passive immigration/ '
A similar instance, it is added, " occurs in the case of the Ligula
simplicissima, infesting the abdominal cavity of various species
of carp, whose sexual organs are, and remain, undeveloped as
long as the worm resides within the fish ; whilst, when the
latter is eaten by ducks, divers, waders, and other water-fowl,
the entozoon being thus conveyed into their intestine, it
attains perfect sexual development. In the older helmintho-
logical works the sexually- mature Ligula simplicissima is
described under various specific names (L. sparsa, L. uniserialis,
L. alternans, L. interrupta) ." These results have been con-
firmed by later observers, but it is now usual to recognise the
sexually-mature worm as the Ligula monogramma of Creplin.
In 1876 Dr Duchamp published his beautiful memoir on this
subject, treating the entire question exhaustively and adding
important experimental details. M. Duchamp gives a list of
about twenty species of fish that are infested by the immature
worm, and amongst these the Oyprinidce play by far the most
conspicuous part. M. Duchamp has recorded a fatal piscine
epizooty amongst tenches {Tinea vulgaris), occurring in the
ponds of La Bresse. This is produced by Ligula simplicissima,
which escapes by an aperture formed near the vent of the
infested fish. M. Duchamp also gives important anatomical
and embryological details, but the especially interesting part
of his memoir refers to his feeding experiments, seven in
number. He succeeded in rearing L. monogramma in the
domestic duck, by feeding this bird with examples of L. simpli-

470

PARASITES OF ANTMALS

cissima obtained from the abdomen of the tench (Tinea vulgaris).
The interest of these experiments does not cease here, since they
afford a probable clue to the source of human Bothriocr/ihal!,
which in nearly all essential points of structure correspond with
the Ligules. As remarked in the first part of this work,
Leuckart long ago pointed to the Salmonidae as probably
furnishing the intermediate host of this worm ; and he dis-
proved the views of Knoch, of Petersburg, who thought he had
reared Bofhriocephalus latus in the dog in a direct manner. I
have already called attention to the opinion of Dr Fock, of
Utrecht, who thinks the human bearer may become infested by
the consumption of the little fresh-water bleak (Leuciscus
alburnns). Prom the observations of Dr Bertolus, it is
extremely probable that our Bothriocejphalus latus is the
sexually- mature condition of Ligula nodosa infesting the abdo-
minal cavity and pyloric appendages of the common trout
(Salmo trutta).

Another cestode of general interest is the Tricuspidaria
(Tricenophorus) nodulosus, infesting many of our fresh- water
fishes. It varies in length from one to two feet. The seg-
mentation of the strobila is very indistinct, but the reproductive
organs occur at regular intervals. All parts of the body are
extremely contractile, especially the head. The tricuspid hooks
support thin chitinous laminae, which connect the two lateral
horns of each hook to the central apophysis. The object of
this arrangement is to afford additional security to the prong-
like processes. Van Beneden appears to think it an error that
the cusps of the hooks should have been figured in f Regne
Animal ' as directed forwards, and he has drawn the hooks
with the points downwards. In regard to the calcareous cor-
puscles, narrow vessels may be easily recognised passing off
continuously from the capsules in closing the particles. These
vascular prolongations are single, having their course directed
towards the epidermis ; doubtless they open at the surface, but
I did not detect any aperture. I have figured the tubes in
my ' Entozoa ' (p. 182). Dr Guido Wagener figures similar
structures as occurring in Cercaria macrocerca.

Various species of Tetrarhynchus dwell in the bodies of
sharks and rays, whilst their larvas inhabit fishes on which the
plagiostomi feed. Immature tetrarhynchs occur in cuttle-fishes,
but they are most abundant in such fish as the cod, haddock,
turbot, whiting (Fig. 81), flounder, sole, gurnard, mackerel,

PISCES

471

mullet, and conger-eel. A tsenioid scolex constantly infests
the muscles and viscera of the great sunfish. The tetrarhynchs
differ from one another as regards the form of
their proboscides and the relative number and
disposition of the hooks. I must refer to my
' Entozoa ' for a full description, with figures,
of a larval tetrarhynch from the wall of the
intestine of a haddock. Some Tetrarhynchi
exhibit a very complex armature, as may be
seen in Tetrarhynchus longicollis infesting the
tope or penny dog-fish (Oaleus vulgaris). In
this species the hooks are uniform in size,
and arranged in spirally disposed circles car-
rying from twenty to thirty hooks each. In
the tetrarhynch from the whiting the hooks
show much irregularity both as regards size
and arrangement. A remarkable scolex in-
fests the sun-fish (Orthagoriscus mold) ; it is
a true tetrarhynch, but has been variously
classed. According to my view all the follow-
ing titles refer to this parasite : — Gymnorhyn-
chus restarts, Rudolphi ; G. horridus, John
Goodsir ; Acanthorhynchus reptans, Diesing ; Bothriorhynchus
continuus, Van Lidth de Jeude ; Bothriocephalic ptatulus,
Leuckart ; Acanthocephalus elongatus, Rudolphi ; A. macrourus,
Bremser ; Floriceps saccatus, Cuvier ; F. elongatus, Blainville ;
Scolex gigas, Cuvier; Tetrarhynchus reptans, Cobbold.

Five or six examples of the sunfish have been examined by
me in the fresh state, all of them being infested by tetra-
rhynchs. In the fish here drawn (fig. 82) the liver and lateral
muscles were extensively tunnelled by the parasite. In all
instances the anterior part of the worm was found surrounded
by a thick, clear, transparent cyst, which gradually diminished
in thickness towards the tail. When liberated from its investing
capsule the head of the worm presents a quadrilateral figure,
each lateral half being furnished with a bipartite facet. The
retractile boring organs are club-shaped, each supporting about
1600 hooks. Nearly all the hooks display a uniform length and
thickness, but at the lower part of each proboscis there are
two conspicuous circles, the hooks of which are at least twice
as large as the others. The joints of the immature strobile are
well formed, but exhibit no trace of sexual organs. If it be

Yig. 81.— Portion of the
proboscis of a scolex
of Tetrarhynchus infest-
ing Merlangus vulgaris.
Magnified. After Busk.

472

PARASITES OF ANIMALS

asked " what is the object of this perpetual tunnelling/' and
" does the boring cause suffering to the host," I reply : —
" The object of tunnelling is apparently twofold ; first, that
the parasite may constantly obtain fresh nourishment ; and
secondly, that it may acquire another residence." It furnishes
an example of a parasite perpetually striving to perform an act
which it cannot accomplish ; for, in order to arrive at sexual
maturity, it must wait until the sunfish is devoured by a shark.
In regard to the question as to the boring action giving rise
to pain, one cannot, of course, speak with absolute certainty.
When there are many parasites occupying the liver, or other
important viscera, then, doubtless, they create pain, and cause
decay of the organs infested ; thus they enfeeble the vital
powers of the host. At such a time the sunfish would be
easily overcome by its natural enemies, and be the first to
succumb in the struggle for existence. These wandering
tetrarhynchoid scolices never escape the body of the inter-
mediate host until they are passively transferred into the
alimentary canal of the ultimate entertainer. In the sharks
and rays they acquire sexual maturity. From these animals
the proglottides pass into the water in the ordinary way. The
ova are subsequently swallowed by sunfishes and other inter-
mediate hosts, within whose stomachs the six-hooked embryos
cire liberate^ and the scolices become developed in the ordi-
nary manner. As obtains in Cysticercus fasciolaris of the
mouse the seolex of Tetr. reptans becomes teenioid. I have
seen the liver of an adult sunfish so infested by these para-
sites that the whole organ might be fitly described as a
mere bag of worms, the immature strobiles being inextricably
coiled together and defying separation. One of the parasites
which I removed from this particular fish is preserved in the
Hunterian Collection.

In reference to the nematoids of fishes I can say but
little. They are excessively abundant ; sexually-immature filarise
being found in almost every marine fish that one examines.
Even at our dinner and breakfast tables nothing is more
common than to observe the little Filaria piscium spirally
coiled within the tissues of herrings, haddocks, cod-fish, and
whiting. All the sexually-immature nematoids are, as it were,
waiting to be passively transferred to their ultimate hosts.
These final bearers are usually either fishes, birds, cetacea, or
seals. Amongst fresh-water fishes the Cucullanida play an

Flo. 82. — Tetrarhynchus replant. 1, Reduced figure of a sunfisli, allowing the worms in
aim ; 2, head ol a worm in its capsule ; 3, tienioid scolex j 4, section of the immature
gtiobile; 6, prohoscis j 6, row ot hooks; 7, 8, large and small hooks (magnified 200
diameters) , D, head of the scolex viewed from abot c. Original.

474

PARASITES OF ANIMALS

important role. These parasites closely resemble the strongy-
loid Sclerostomata, but the absence of a true bursa seems to
justify their separation into a distinct family. In most of them
the body is truncated in front and much narrowed or drawn
out posteriorly. The head is broad and globular, and furnished
with a powerful muscular pharynx. The mouth is seldom
round; it is often subterminal, opening by a transverse slit.
The tail of the male is recurved, and usually supplied with
membranous winged appendages; sometimes there is a pre-
anal sucking disk. In the female the tail is simple, and more
or less sharply pointed.

The facts relating to the development of these parasites are
especially interesting as having afforded Leuckart and Fed-
schenko a clue to what obtains in the guinea-worm {Dracun-
culus). The Cucullanus of the perch (G. elegans) is a vivipa-
rous species. The embryos are supplied with little boring
teeth, or styles, which enable them to perforate the bodies of
entomostracous crustaceans. Having in a direct manner gained
access to the perivisceral cavity of Cyclops, they remain coiled
within the intermediate bearer until it has been pursued,
captured, and transferred to the stomach of the ultimate or
piscine host. Once liberated within the stomach of the fish the
young Oucullani soon acquire sexual maturity.

The acanthocephalous Echinorhynchi are very abundant in
fishes. They also, like the Oucullani, require a change of

nscES

475

hosts in order to ensure the continuance of the species. No
less than six species of Echinorhynchi are known to infest the
trout (Sahno fario). As many as four species likewise infest
the eel (Anguilla) ; the same number of distinct forms being
also found in the turbot (Rhombus) and ling (Lota), whilst
three species may be met with in the common sole (Solea).
What we at present know respecting the mode of development
of Echinorhynchi infesting fishes is principally due to the
researches of Leuckart. Some years back Dr Guido Wagener

YiQ. %\.—\ Echinorhynchus angnstatus (natural size and enlarged) ; 2, Echinorhxinclivs
twdulosus (natural size and enlarged), with (3) two eggs (magnified 1000 dian/etcrs)
Both species from a trout. After Busk. ;

supplied admirable illustrations of the eggs and embryos of
Echinorhynchi, but he was erroneously led to conclude that the
larvae were developed in a direct manner. The notion of a
simple metamorphosis was entirely disproved by the experi-
ments of Leuckart, who found the growth and development of
the young to be accompanied by a true alternate generation.
He showed this to obtain in Echinorhynchus proteus, a species
abundant in the trout and in many other fresh-water fishes. The

470

PARASITES OP ANIMALS

embryo of this parasite is broad and obliquely truncated at the
ventral surface anteriorly, being gradually narrowed to a blunt
point posteriorly, and at the front part, on each side of the
middle line, there are five or six spines biserially disposed.
Similar characters are seen in E. jilicollis. Prof. Leuckart
introduced a number of eggs into a vessel of water con-
taining several small crustaceans (Gammarus Pulex). These
little animals readily swallowed the ova, and in a few days the
embryos were found emerging from their shells, boring their
way through the intestinal walls, then passing into the general
cavity of the body, and even into the appendages themselves.
During the next fourteen days the embryos within the Gammari
exhibited an increase of size ; and in course of the third week
a further metamorphosis caused the embryos to assume the
readily recognisable characters of a young Echinorhynchus.
Thus, in Leuckart's own words, " the ultimate animal arises in
the interior of the primordial body, by a process which presents
so close an analogy with the production of an embryo, and,
consequently, with the act of generation, that one feels inclined
at once to identify it with such an act, and therefore, also, to
regard the Echinorhynchus as exhibiting an alternation of
generation in its mode of development rather than a metamor-
phosis."

The young Echinorhynchus afterwards grows rapidly, its
several internal organs, proboscideal sac, and muscular apparatus,
gradually coming into view. At last the young entozoon com-
pletely fills the interior of the embryo, the latter having scarcely
undergone any change, and still remaining, of course, within its
crustacean host. What may be regarded as even more extra-
ordinary is the circumstance that the embryonic body next
becomes firmly adherent to the young Echinorhynchus, thus
ultimately forming the true integument of the adult Echinorhyn-
chus. The original skin of the embryo, however, is cast off
" as soon as the Echinorhynchus occupies the whole interior of
the embryo." After this the sexual differences become clearly
established. Leuckart remarks that the passage of the young
Echinorhynchi into their ultimate host is probably unattended
by any striking changes, whilst .the metamorphosis of the embryo,
as thus far detailed, occupies a period of about six weeks. In
general the crustacean hosts appear to suffer little from the
borings of the embryo parasites, but when the latter have
assumed the Echinorhynchus - condition and happen to be

PISCES

477

particularly numerous they not unfrequently prove fatal to the
unsuspecting Gammari. After their transference to the intestine
of the ultimate host a period of about one week more is required
for the completion of their development.

From the large number of species of Echinorhynchi infesting
our fresh-water fishes, they present quite a feature of piscine
parasitism. Almost every perch, chub, carp/ pike, barbel, bream,
or roach that one opens is found to have its intestines occupied
by parasites which exhibit a light yellow color. These are
Echinorhynchi, the common forms being E. proteus, E. angus-
tatus (Fig. 84, No. 1), E. clavceceps, E. globulosus, and E.
tub&rosus. In the Salmonidae, besides several of the above, we
may also find E. clavula, E. fusiformis, and E. pachysomus. As
a group these parasites are more attractive looking than most
other helminths, and they will well repay the zoological collector.
The species infesting marine fishes are almost as numerous as
those found in fresh-water hosts.

Bibliogeaphy (No. 59). — (Anonymous), " Note by ' An In-
quirer ' respecting Worms in Fish," in the ' Lancet/ March 7,
1868, p. 336. — Baer, K. E. von, " On the Tapeworms found in the
"Waters of the Pregel by Linneus/' from the German, in ' Edin.
New. Phil. Journ./ 1829, p. 374, and in the ' Edin. Nat. and Geo.
Sci. Journ./ 1829-30, p. 3J1 ; also from the ' Trans, of Nat. Soc.
of Dantzig/ in the ' Lancet/ 1829. — Badcock; see Slack. — Baird,
'British Entomostraca/ London, 1850. — Beneden, J. P. van, " Les
vers cestoides/' 'Mem. deFAcad. Roy. de Belg./ torn, xxv, 1850.
■ — Idem, ' Mem. sur les vers intest./ Paris, 1858. — Idem, " On
Echinobothrium/' in 'Bull, de l'Acad. de Brux./ 1849.— Idem,
(with Hesse), ' Rech. sur les Bdellodes ou Hirudinees et les
Trematodes marins/ 1863-65. — Idem, 'Rech. sur les Turbel-
laries/ 1861. — Idem, "On a new Lerneonema," ' Bull./ 1. c,
1851, and in ' L'Institut/ 1851. — Bertolus, 'Mem. sur le deve-
loppement du Dibotlirium latum (Bothriocephale de Phomme)/
App. to Dr Duchamp's work quoted below. — Bosc, ' Hist. Nat.
des Yers/ 1802, p. 271. — Bradley, 0. L., "On the occurrence

of Gyrodactylus on Sticklebacks/-' ' Proc. Linn. Soc./ 1861.

Brulle, " Note on the Reproduction of Ligula/' from ' Comptes
Rendus/ in 'Ann. Nat. Hist./ 1855. — Chatin, ' On Amphibdella
torpedinis from the Gills' (1. c, Bibl. No. 57). — Ghavannes,
" On Fluke-larvsB from Coregonus/' in ' Bull, de la Soc. Vaud.
des Sci. nat./ torn. iii. — Claparede, E., " Ueber die Gattung
Tetracotyle/' &c, in ' Zeitsch. f. wiss. Zool./ 1857, s. 99 et

478

PARASITES OF ANIMALS

seq. — Cobbold, " The Sunfish (Orthagoriscus) as a Host/'
' Intell. Observer/ Sept., 18G2. — Idem, "Notes on the Calcareous
Corpuscles of Tricuspidaria," ' Quart. Jouru. Micr. Sci./ 1859.
— Idem, " Notes on Tricuspidaria and Pentastoma," ibid., 1 859.
— Idem, " Note on Gyrodactylus elegans," ibid., 1862 ; see also
Wedl. — Idem, " On Distoma clavatum from the Sword-fish,"
'Proc. Linn. Soc./ 1867 (Zool. Sect., p. 200).— Idem, "Tape-
worms in Trout/' letter to the ' Field/ July 26, 1873 ; see also
P. Francis (below). — -Idem, "Remarks on the Entozoa and
Bctozoa of Fish/' the ' Veterinarian/ Oct., 1867, p. 671.— Idem,
" On Agamonema crenilabri," in ' Science Gossip,'" 1876 ; see
W. W. Wilson.— Idem, in ' Linn. Trans./ 1858.— Idem, ' Note
on Parasites from the Wolf-fish (Annarhicas) and Lump-sucker
(Cyclopterus), collected by Mr Devis ' (1. c, Bibl. No. 57). —
Idem, " Descr. of the Scolex of a Tetrarhynchus," in a paper
on ' Parasite Larvae/ c Intell. Observer/ 1863. — Idem, "Synopsis
of the Distomidae," 'Proc. Linn. Soc/ (Zool. Div.), I860.—
Idem, " Remarks on Bothriocephalus latus, in relation to Dr
Fock's supposition that the Bleak (Leuciscus alburnus) is con-
cerned in its Production/' the 'Veterinarian/ July, 1878. —
Comalia, E., ' Sopra una nuova specie di Crostacei Siphono-
stomi (Gyropeltis doradis),' Milan o, 1859. — Ooughtrey, M., " On
the absence of Tapeworm in the Salmon-trout of New Zealand," -
letter to the ' Otago Daily Times/ dated from the Otago Uni-
versity, Dec. 6, 1875. — Diesing, ' Vierzehn Arten von Bdelli-
deen/ Wien, 1858; see also 'Revisions/ quoted in Bibl. No. 58.
— Donnadieu, A., "Etude sur les Ligules," in ' Archives Zool.
Experiment./ 1876. — Duchamp, G., ' Recherches anat. et
physiol. sur les Ligules/ Paris, 1876 ; see also Bertolus. —
Foch, ' The Bleak as a source of the Broad Tapeworm.' See
Cobbold. — Francis, F., "Tapeworm in Fish" (with report
by myself), in the 'Field/ June 28, 1873. — Idem, in the
'Field' for July 12, 1873 — Garsin, ' Histoire de l'Acad. des
Sci./ Paris, 1730, p. 44. — Goodsir, J., "On Gymnoryitchus
horridus, a new Cestoid Entozoon," ' Edin. New Phil. Journ./
p. 9, 1841. — Houghton, W., "On the occurrence of Gyrodactylus
elegans in Shropshire," ' Ann. Nat. Hist./ 1862. — Huxley, "Note
on Gyrodactylus," ' Proc. Roy. Inst./ April 20, 1852, and
' Edin. New Phil. Journ./ 1852, p. 172. — Enoch, J., ' Entwick-
elungsgeschichte d. Both, probosoideus,' 1862. — Knox, J. F.,
" Note respecting the occurrence of a peculiar Microscopic
Eutozoon in the Textures of the Herring," 'Lancet/ 1838. —

PTSCES

479

KolUher, "Zwei neue Distomen," ' Ber. v. d. K. Zoot. Anstalt
zu Wiirzburg/ 1849. — Idem, ' Ueber Tristoma/ ibid., 1849.—
Leidij, J., " Notice of a Tetrarhynchus (T. tenuicaudatus) in
the Remova;" ' Proc. Acad. N. S. Philad./ Oct. 15, 1878;
and in 'Ann. Nat. Hist./ Feb., 1879. — Ley dig, "Ueber
Argulus," 'Sieb. und K611. Zeitsch./ 1850. — Maddox, B. L.,
" Some Kemarks on the Parasites found in the Nerves (and
other parts) of the Common Haddock (Morrhua ceglefinus),"
' Trans, of the Roy. Micr. Soc./ 1867, p. 87. — Mensries,
'Linn. Trans./ 1790, p. 187. — Miescher, "On Filaria piscium,"
&c, in 'Excerpta Zoologica/ communicated by Dr Prances,
in 'Ann. Nat. Hist./ 1842 — WIntosh, W. C, "Notes on the
Food and Parasites of the Salmo salar of the Tay," ' Proc.
Linn. Soc./ 1863; repr. in the 'Zoologist/ Feb., 1864.—
Muller, J., " Note on a Parasitic Formation (Gregarina) in
the Pike, with a statement from his 'Neurologie der Myxi-
noiden/ that Diplostomum rachineum is to be found alive under
the cerebral membranes of Petromyzon fluviatilis," from 'Miiller's
Archiv/ in ' Micr. Journ. and Struct. Record/ p. 20, 1842. —
Nardo, in ' Heisinger's Zeitsch./ 1827, s. 68, and in ' Isis/
1833, s. 523. — Olsson, P., "Researches on the Flukes and
Tapeworms chiefly of Marine Fishes," ' Entozoa, iakttagna hos
Skandanaviska Hafsfiskar/ Lund (aftr. ur ' Lunds Univ.
Arsskrift/ torn, iii, iv), 1867—68. — Owen, ' Zool. Soc. Trans./
1835, p. 382. — Pallas, ' Spicilegia Zoologica/ fasc. x, p. 18,
1774. — Siebold, G. von, 'Band und Blasenwiirmer/ s. 41, Huxley's
edit., p. 32. — Idem, "On Diplozoon paradoxum," from 'Zeitsch.
f. wiss. Zool/ (by Huxley), in 'Ann. Nat. Hist./ 1851. — Idem,
" Ueber den Generationswechsel der Cestoden nebst einer
Revision der Gattung Tetrarhynchus," ' Zeitschr. f . wiss. Zool./
1850, s. 198. — Idem, " Gyrodactylus, ein ammenartiges Wesen,"
ibid., 1849. — Slack, H. J., " On Bucephalus polymorphic
in 'Monthly Microsc. Journ./ April, J 875, p. 141. — Van
Beneden (see Beneden). — Verrill, A. E., " On the Parasitic
Habits of the Crustacea," from ' American Naturalist/ in
'Scientific Opinion/ Aug. 4, 1869, p. 185. — Idem, "New
Flukes {Tristoma lave and T. comutum) from the Mouth and
Gills of Tetrapturus albidus," 'American Journ. of Science/
p. 40, 1875. — Von Baer (see Baer). — Von Siebold (see Siebold).
— Wagener, R. G., "Helminth. Bemerkungen," in ' Sieb. und
K611. Zeitsch./ 1857— Idem, " Enthelrainthica," ' Miiller's
Arch./ 1851. — Idem, "Ueber Eingeweidowurm (Ainphiptych.es)

480

PARASITES OF ANIMALS

in Chimara monstrosa," ' Mull. Arch./ 1852. — Idem, * Beitrage

zur Entw.-Gesch. der Eingeweidewiirmer (Preisschrift)/ 1857.

Wedl, " On Gyrodactylus " (see reference to my paper on ' G.
elegans'), ' Quart. Journ. Micr. Sci./ 18G2 • trans, from his
' Anhang/ " Ueber die Gattung Gyrod./' to ' Anat. Beobach-
tungen ueber Trematoden/ Wien, 1858. — Idem, ' Haematozoa in
Fishes/ &o. (1. c, Bibl. No. b8).—Wigham, B., "Note on
Holostomum cuticola from Eoach and Bream/' 'Ann. Nat.
Hist./ p. 235, 1851.— Wilson, W. W., "On a Parasitic Worm
infesting a Marine Fish {Crenilabrus rupestris) ," in ' Science
Gossip/ Jan., 1876. — Yarrell, W., "Note on Tristoma coccineum,"
in his work on 'Brit. Fishes/ vol. ii, p. 353, 1836.

Pakt IY (Evertebrata) .

Since a large proportion of all those helminths that require
a change of hosts must needs pass into the bodies of insects,
crustaceans, mollusks, or other evertebrated animals, it is
evident that these lower creatures are almost as liable to be
infested by parasites as the vertebrates themselves. As a rule,
no doubt, the parasitic forms infesting individual evertebrated
hosts are not numerous ; nevertheless the water-snails form a
noteworthy exception. Thus, some ten different species of
parasite are found either in or upon the common Planorbis
corneus ; whilst Lymnceus stagnalis, Paludina vivipara, and
P. impura, each support at least a dozen species. Of course,
the parasites are not sexually mature, since nearly all of them
are Cercarice or larval trematodes. Snails, oysters, mussels,
whelks, and other mollusks afford harbour and anchorage to a
variety of parasites and messmates ; but, fortunately, few or it
may be none of the strictly human parasites require to pass
through these intermediate bearers. Distoma crassum is possibly
an exception. Save the cuttle-fishes, not many evertebrated
animals are infested by sexually-mature worms. One of the
most notable exceptions is that of a nematoid infesting bees.
This worm was known to John Hunter, who spoke of it as
"the animal that breeds in the bumblebee.^ In the year
1836, M. Leon Dufour first applied the term Sphccrxdaria to
this remarkable worm, which he discovered in the abdominal
cavities of two species of bee [Bombus terrestris and B. hor-
torum). The worm was subsequently found by Von Siebold in

EVE11TEBEATA 481

two other species of bee (B. muscorum and B. sylvarum), but it
remained for Sir John Lubbock to demonstrate that this
parasite not only infests these insects, but also Bombus lucorum,
B. layidarius, B. pratorum, B. subterraneus, and Ajpathus
vestalis. I possess specimens from Vespa vulgaris and V. rufa.
Sir J. Lubbock and Mr. Cole have separately given full
anatomical descriptions of the worm. According to Lubbock
the so-called female is about an inch in length, of a whitish
color, and in thickness, being bluntly pointed at either
extremity. Sphcerularia is everywhere covered by small warts
or button-like projections, in all numbering about 800. The
warts are transparent, each, according to Lubbock, project-
ing from ~f to above the general surface of the integu-
ment. There is neither mouth, oesophagus, intestine, nor

Fiq. 85.—Sphcerularia bombi. Showing the supposed male in sitti. After Lubbock.

anus j but in their place a large fatty mass or corpus adiposum.
Sir J. Lubbock remarks that this peculiar organ " is homo-
logous, not with the whole intestinal canal of nematodes, but
only with the intestine ; and we find, in fact, that in Grordius
the oesophagus is very short, and opens at once into the
anterior end of the corpus adiposum ; so that to pass from this
genus to Spharularia it would be necessary to shorten the
oesophagus a little more, and then the wall of the coo-pus
adiposum would be immediately attached to that of the body.
So far, therefore, as concerns the corpus adiposum and the

[•82

PARASITES OF ANIMALS

oesophagus, Sphcerulcvria agrees neither with Gordius nor
Mermis, nor, indeed, with one more than the other ; since, if
it agrees with Mcrmis albicans in the double series of large fat
cells, it has no oesophagus, and in this respect more nearly
resembles Gordius." The reproductive organs consist of a
single ovary, uterus, and terminally situated vulva. These
organs in the full-grown females contain ova in all stages of
development up to the condition of advanced yolk segmenta-
tion ; but it does not appear that embryonic formation takes
place whilst the eggs are still in tdero. " The young animals
are born soon after the eggs are laid. They are about in
length, and ~5" in diameter at the broadest part. Before Sir
J. Lubbock conducted his inquiries the so-called male appears
to have been overlooked. The male, if male it be, is extremely
minute ; that is to say, about 28,000 times smaller than the
female. Notwithstanding this very circumstantial account
based on Lubbock's determinations, Schneider has sought to
show that the facts have been entirely misinterpreted. What
Lubbock regards as the male worm is, in Schneider's opinion,
a female, whilst the so-called female is nothing more than a
gigantic prolapsed uterus which has become many thousand
times larger than the body of the worm whence it proceeded.
It must be allowed that Schneider's description and accompany-
ing figures are very convincing. When revising the entozoa
of the Hunterian Collection in 1866 I explained the specimens
and dissections in accordance with Lubbock's views. In the
following year Prof. Huxley in his College Lectures supported
the view of Schneider, but in his recently published manual
the opinions of the Berlin helminthologist are not so much as
alluded to.

Another point of special interest in connection with the
parasites of insects concerns the development of Mermis albicans.
At or near the time of the maturation of the ova, the parent worm,
hitherto lodged within the body of some insect, buries itself in
the soil. It commences its migration by boring its way out of
the body of the host. Some difference of opinion exists as
to the condition of the parent at the time of its wandering, for
Von Siebold asserted that it quitted its parasitical mode of life
" in order to become sexually mature away from the animal "
infested ; whereas Van Beneden states that the embryos are
always formed at the time of the wandering.

From Von Siebold's experiments it would appear that in-

EVE RTEBRATA

483

completely developed Mermes can become mature whilst still in
the soil ; but the normal condition requires the wandering to
commence, as we have said, at or near the full time of embryonal
development. The embryos are reproduced viviparously, and
being set free, they pass a certain period of their existence in
the soil. Here they grow rapidly, acquire sexual organs, and
subsequently seek to " gratify their immigrative propensities/''
as Von Siebold says, by selecting and penetrating the soft-bodied
larvae of lepidopterous and other insects. This entrance they
accomplish by means of a sharply-pointed dentule or boring
stylet, which at the time of disuse is concealed within the
head. Having once gained access to the host they remain
within its body until the caterpillar has become transformed
into the perfect butterfly, or until their own sexual maturity is
completed. Yan Beneden thinks it probable that the males
quit the host some time before the females, a view which, if
correct, might alone account for the comparative scarcity of the
males. According to Yon Siebold, sexual congress occurs before
the entrance of the worm into the caterpillar. This observation
agrees with the generally admitted fact that hitherto no male
Mermes have actually been detected in the bodies of insects.
The Gordii, like Mermes, become free in damp earth and
penetrate the bodies of certain insects or their larvae. Some
of them gain access to fishes. Like the free nematodes
(Anguillulidat) , many of the Gordii will survive complete desic-
cation. The eggs of the mature worms are deposited in long
agglutinated chains in water or damp situations.

I must conclude. In the body of this work will be found
many notices of insect parasites that are awaiting transference
to some vertebrate. I need only allude to the role of the
mosquito, to that of the louse of the dog, and especially to that
of the little myriapod (Glomeris) which, like the common
glow-worm (Lampyris), possesses phosphorescent properties. I
mention this again partly in correction of an entomological
error (at p. 296) which escaped me at the time of going to
press. Leidy has described a mature nematode (Ascaris iwfecta)
from Passalus cornutus, and numerous Filariae are known to
infest insects {Blatta, Forjicula, Phosphuga, &c, &c). From
an earwig I obtained a filaria nearly five inches in length.

We have seen that the larvae of Dracunculus, CucuUanus, as
well as those of other important nematodes, dwell in bodies of
cntomostracous Crustacea, whilst those of JUchinorhynchus attack

1,8 i

PARASITES OF ANIMALS

the Gammari and their allies. The well-known Udonell" caUgorim
attaches itself to Crustacea that are themselves parasitic.

As many of the so-called free nematodes live in the slime
of animals, Yillot is of opinion that no very distinct line of
demarcation can fairly be drawn between the parasitic and
free species. This work, however, having dealt only with
genuine parasites, I have purposely omitted any detailed account
of the so-called free nematoids. I mention this lest it should
be supposed that I had shown a studied neglect of the more or
less remarkable labours of Biitschli, Bastian, Eberth, Linstow,
Marion, Yillot, Claus, De Man, Carter, and many others.

Bibliogeaphy (No. 60). — Bastian, H. C, "Monograph on
the Anguillulidge, or free Nematoids, marine, land, and fresh-
water, with description of 100 new species," ' Linnean Trans/
for 1865, vol. xxv, p. 73. — Idem (see Bibliog. No. 2). — Idem,
" Free Nematoids," being an article in the ' Popular Science
Review' for 1868, vol. vii, p. 163. — Brady, G. S., ' Monograph
of the free and semiparasitic Copepoda/ London, 1878. —
Biitschli, 0., " Untersuchungen ueber freilebende Nematoden
und die Gattung Chsetonotus," ' Sieb. und Koll. Zeitschrift/
1876. — Carter, E. J., " On a Bisexual Nematoid Worm which
infests the common House-fly (Musca domestica)," ' Ann. Nat.
Hist./ 1861, and in the ' Bombay Med. and Phys. Soc. Trans./
new series, 1860. — Olaparede (see Panceri). — Glaus, 0., ' Beo-
bachtungen ueber d. Organis. und Fortpflanz. v. Leptodera
appendiculata,' 1869. — Oobbold, "Note on Insect Parasites," in
'Rep. of Entomological Club/ in the ' Midland Naturalist/
March, 1878, p. 80.— Dole, W., " Remarks on a Parasite of
Humble Bees," in ' Journal of the Quekett Microscopical
Club/ 187b.—Dufour, L., " Sphasrularia," 'Ann. des Sci. Nat./
1836. — Dujardin, " On Mermis," c Ann. des Sci. Nat./ 2e ser.,
torn. 18, p. 129. — Eberth (see Bibliog. No. 2). — Gamer, B.,
" Note on a Distoma," in his paper 1 On the Lamellibranchiate
Conchifera/ 'Trans. Zool. Soc./ 1841— Ghaleb, 0., "Observa-
tions and Experiments on the Migrations of Filaria rhyti-
pleurites, a Parasite of Cockroaches and Rats," ' Comptes
Rendus/ July 8, 1878, and 'Ann. Nat. Hist./ Aug., 1878.—
Idem, " Note sur l'anat. et les migrations de deux Nematoides
parasites, le Poecilogaster blatticola et Fil. rhytipl.," Paris, 1876
(quoted by O. von Linstow).— Giard, M. A., " On the parasitic
Isopoda of the genus Entoniscus (infesting Crustacea)," from
'Comptes Rendus/ Aug., 1878, in 'Ann. Nat. Hist./ Ota,

EVERTEBJIATA

485

1878. Idem, " On the Orthonectida, parasitic on Echinoder-

raata and Turbellaria (Rhopalura)," f Ann. Nat. Hist./ Feb.,
1878. — Grube, A., " On Cyclops as a new Cestoid-bearing
Host," from 'Zoologisch. Anzeiger/ Bd. i, s. 74, in ' Journ.
Royal Microsc. Soc./ Nov., 1878, p. 254.— Hunter, J., " Filaria
of the Bee," in ' Catal. (by Owen) of the contents of the Mus.
Royal Coll. Surg./ part iv, fasc. i, p. 37, 1830.— Kynston,
" Worms attached to a Grasshopper," ' Proc. Ashm. Soc./ in
' Corbyn's India Review/ and in ' Journ. of Foreign Sci./ 1837,
p. 172. — Lima, J. F. da S., " Remarks on the Filaria medi-
nensis or Guinea-worm ; on the occurrence of this parasite in
the Province of Bahia; and on its entrance into the human
body by drinking water trans, from the Portuguese by Dr
J. L. Paterson, and pub. in the ' Veterinarian ' for Feb., 1879
et seq. — Linstow, " Helminthologische Beobachtungen," in
'Archiv fur Naturgeschichte/ 1876. — Lubbock., Sir J., "On
Sphterularia bombi," ' Nat. Hist. Rev./ 1861. — Idem, "Notes,"
&c, ibid., 1864, p. 265. — Mason, J. W., " Note on the Geo-
graphical Distribution of the Temnocejohala chilensis (parasitic
upon a freshwater crayfish, Paranephrops setosus, in New
Zealand)," ' Annals Nat. Hist./ 1875, p. 336.— Marion, A. F.,
' Revision des Nematodes (&c.)/ Marseilles. — Maund, B., " A
description of Filaria forficulce," e Rep. Proc. Linn. Soc./ in
'Zool. Journ./ 1832-34, p. 263. — Meissner (see Thomson). —
Owen, JR. (see Hunter). — Pagenstecher (see Bibl. No. 58). —
Panceri, P. (e di E. D. Claparede), "Nota sopra une alciopide
parassito dell Cydippe densa," ' Mem. della Soc. Ital. di Sci.
Nat./ 1867. — Sars, "Intestinal Worm in an Acaleph.," from
'Wiegmann's Archiv/ in ' Ann. Nat. Hist./ 1845. — Siebold, 0.
J. von, in 'Wiegmann's Arch./ 1835. — Idem, in ' Ray Soc.
Rep/ (by Busk), 1847.— Idem, "Worms," &c, ibid., p. 503,
1847. — Idem, " Report on Helminthology, and on the Nemer-
tinae" (trans, by W. B. Macdonald, in ' Ray Soc. Rep. on Zool./
1842, p. 280), Edinburgh, 1845.— Idem (see Thomson). — Thom-
son, A. (for review of the writings of Meissner, Von Siebold,
and others, respecting the development of Mermis, Gordius, &c,
see the classical and elaborate art. " Ovum "), in ' Supp. to
Todd's Cyclop./ 1859. — Vogt, C, " On some Inhabitants
(Cercariae) of the Fresh-water Mussels," from 'Ann. des Sci.
Nat./ in 'Ann. Nat. Hist./ 1850.— Whitman, 0. 0., "The
Embryology of Clepsine (with valuable Bibliography)," f Quart.
Journ. Micr. Sci./ July, 1878.

48G

rARASITES OF ANIMALS

Appendix.— The memoirs announced by Dr T. E. Lewis in
the January issue of the 'Microscopical Journal/ and referred
to at the close of my account of Filaria Bancrofts, having
appeared, I fulfil the promise previously made (p. 202). In
the few lines at my disposal I may observe that the beautiful
brochure (quoted below) supplies fuller details of the results
already announced by Lewis in the ' Proceedings of the Asiatic
Society of Bengal.-' In respect of the nematoid hsematozoa, the
memoir is chiefly important as confirming Manson's observa-
tions regarding the changes undergone by the Filariae that have
been transferred to the stomach of the mosquito, and especially
also, as advancing some novel facts in reference to the occurrence
of bird's blood-corpuscles, associated with embryonic nematoids,
in the same viscus of the insect. The worms are regarded by
Lewis as transferred avian haematozoa, a view which gains
strength by their comparison with the similar larvae which he
had detected in the blood of Indian crows {Oorvus splendens).
In Egypt, as Sonsino had himself informed me by letter, similar
haematozoa are to be found in crows, and avian filariae of this
kind were long previously described, as Lewis and Sonsino point
out, by Borell, Herbert, Schmidt, and Yirchow. Facts of this
order undoubtedly complicate matters, and suggest that extreme
measure of caution in drawing conclusions, which Lewis himself
everywhere displays.

Respecting the final changes undergone by the mosquito-
filariae before their re-entrance into the human body, Lewis does
not appear to have gone further than Dr Manson. By rupturing
the body of the most advanced larvae, Lewis readily recognised
the oesophagus and intestine, but he remarks, significantly, "I
have not been able to distinguish any other differentiated viscus
in any of the specimens, and certainly, nothing suggestive of
differentiation of sex" (p. 83). In an earlier part of the me-
moir Dr Lewis takes objection to my view that the urinary
nematoids found by me in a case of Bilharzia are genetically
related to Filaria sanguinis hominis. His distinguished coad-
jutor, Dr D. Cunningham, also denies the possibility of such
relationship. No doubt, if the urinary maternal worm was really
oviparous my view is untenable ; but the proved presence of
imperfectly formed ovarian ova, in which no trace of embryonic
formation was discernible, has forced upon me the conviction that
prolapsus and rupture of the uterine tubes of the parent worm
had occurred, and that their rupture had occasioned the escape

APPENDIX.

487

of ova in various stages of growth. As free embryos were
also detected, the adult worm was probably viviparous. There
is an error in the representation of the oval-shaped ovum given
in the figure (p. 183). I retain drawings of eighteen perfect
nematoid ova from the Bilharzia case, and not one of these
shows any double contour of the chorional envelope. In
the case of the imperfect ova, the double contour is obviously
due to the close apposition of the yelk-membrane to the shell-
membrane, there being no true shell. As regards " a correc-
tion " which Lewis makes in respect of the question of priority
of description of the mature Filaria sanguinis hominis I can only
find space to state frankly, that Lewis is perfectly correct.
The error was quite unintentional on my part. The adult worm
was first discovered by Bancroft, and upon the strength of his
admittedly scanty record I named the worm Filaria Bancrofti.
In the matter of supplying a proper diagnosis and an anatomical
description I was completely anticipated by Lewis. No doubt,
Dr Bancroft could have furnished a fuller description of the
parasite, had he desired to do so, but here is what he says in
the letter addressed to me from Melbourne on the 20th of
April, 1877 : — " I thought it better to send you this account of
filariae than to publish it direct, as you so kindly set me on the
track of the investigation." Here I feel constrained to remark
that few, if any, of my many correspondents in helminthology,
have displayed more engaging candour. Whilst actually writing
this Appendix (April 15th, 1879) I have received a new record
of filarious cases from Dr Bancroft, who also sends me some
mosquitoes captured by a victimised patient whose blood
swarmed with filarias. In one of the captured insects Bancroft
himself detected forty-five filariae. The cases have been for-
warded to the e Lancet ' for publication. Lastly, in reference
to the closing paragraph of Bancroft's previous letter to me
(pub. in the 'Lancet/ Feb. 1st), I have received the following
interesting commentary at the hands of Dr Silva Araujo,
whose letter is dated from Bahia, March 3rd, 1879 : — " Je dois
vous communiquer que ce fait vient confirmer l'idee qui existe
chez nous, ou le peuple croit et affirme que — quand une per-
sonne qui souffrait auparavant d'erysipele a un abces cela la
preserve de nouveaux acces. La raison ne sera-t-elle pas que
dans ce cas, avec l'ouverture de 1' abces, le ver sort ? Je le
crois. Ces faits viennent demontrer que la cause de la maladie
est le ver. Cependant nous avons ici a Bahia plusieurs con-

488

PARASITES OF ANIMALS

freres qui ne le croient point ! Et a Rio- de- Janeiro aussi il y
en a, peut-etre davantage (!)." T will only add that Dr Araujo
deceives himself if he imagines that the full etiological sig-
nificance of parasites in relation to disease will receive general
professional recognition for many years to come.

Supplement to Bibliography No. 23, p. 202 (with emenda-
tions).— Araujo, ' Memoria sob re a Filariose ou a molestia pro-
dusida por uma nova especie de parasita cutaneo/ Bahia, 1875.
— Idem, " Da filariose/' ' Globo/ Jornal do Rio de Janeiro,
1876, e ' Re vista Medica do Rio de Janeiro/ 1876, anno 3o,
No. 2, 15 de Julho, p. 107. —Idem, " Caso de chyl uria, ele-
phancia do escroto, escroto lymphatico, craw- craw e erysipela
em um mesmo individuo ; descobrimento da Wuchereria filaria
na lympha do escroto. Tratamento pela electricidade com
excellentes resultados," ' Gaz. Med. da Bahia/ 2a serie, vol. 2o,
No. 11, Nov. de 1876. — Idem, "A Filaria Wuchereri no sangue,"
'Gaz. Med. da Bahia/ Mar. de 1878, p. 106, e seguintes. —
Idem, " A muricoca e as filarias Wuchereri," ' Gaz. Med. da
Bahia/ Setembro de 1878. — Idem, " La Fil. immitis," &c,
Transl. of Mem. (Z. c, Bibl., No. 45) in ' Lyon Medical/ Nov.,
1878, p. 319 et 363. — Bancroft, "Instance of a European having
taken leprosy in Queensland," in a letter to myself ; see " Case
from Bancroft," quoted at p. 203. — Ohassaniol, A. (et F. Guyot),
" Hematurie graisseuse ou chyleuse," in their " Notes de Geo-
graphie Med. recueillies a Taiti," in ' Archives de Med. Navale/
Jan., 1878, p. 65.— Cobbold, " Worms in the Heart of Dogs,"
letter in the ' Lancet/ April 5, 1879, p. 498. — Coles, "On Lymph-
scrotum," 'Brit. Med. Journ./ March 9, 1878. — Fayrer, Sir J.,
" Lecture on Elephantiasis Arabum," in the ' Lancet,' March
29, 1879, p. 433— Idem, 'Report of Pathol. Soc./ 'Lancet/
Feb. 22, 1879, p. 267. — Idem, ' Rep. of Epidemiological Soc./
ibid., p. 269. — Idem, 'Letter on Filaria/ see Hoysted. — Ghaleb,
0. (with P. Pouquier), " On Filaria hcematica," from ' Comptes
Rendus/ Feb. 5, 1877, in ' Annals Nat. Hist./ April, 1877.—
Hoysted, J., " Notes of a Case of Filaria sanguinis in a Dog ;"
see Bibliog. No. 49, p. 311. — Lewis, T. E., 'The Microscopic
Organisms found in the Blood of Man and Animals, and their
relation to Disease/ Calcutta, 1879. — Idem, " The Hasmatozoa
of Man (excerpt of the above)," 'Quart. Journ. of Microsc. Sci./
April, p. 245 (both from ' 14th Ann. Rep. of the San. Commis-
sioner with the Govt, of India — Mahuna, 'Letter respecting
Fil sang. horn, in Chyluria ' (1. c, Bibliog. No. 23).

INDEX.

PAGE

Acanthia lectularia, or the common bed bug .... 273

Acanthocephala in birds 446
„ include but one family . . • .5

„ in the Hunterian Museum . . . .413

Acanthotceniadcp., a family of tapeworms . • • .4

Acaridce, a family of Arachnida . * . • .5

Acephalocysts in cattle, Hunterian specimens of . . . • 140

Agricultural societies, their indifference to helminthology . . .412

Aguti, Spiroptera mediospiralis from the .... 403

Alligator, entozoa from an . . . . • .455

Amphistoma and Polystoma in the frog .... 452

„ hominis, notice of Lewis's and McConnell's . . .36

„ subclavatum, ciliated embryo of . . .49

Amphistomes infesting deer . . . . . . 332

,, „ elephants ..... 395

„ „ the horse ..... 357

Amphistomidce, a family of flukes . . . . .4

Anaemia, tropical, due to Anchylostomata . . ... 203

Anchylostomum-helminthiasis, Wucherer's account of the . . 213

Anchylostomum (Dochmius) duodenale, description of . . 212

Aneurism of the horse, Bollinger's account of. . . . 368

Anguillula (Leptodera) stercoralis, description of 234
AnguillulidoR, a family of round worms . . . .5

Annelid parasites, ectoparasitic character of the . . . .5

Anoplocephala perfoliata as a synonym .... 401

Ant-eaters, internal parasites of » . . . . 321

Antelopes and deer, Pentastomes found in . . . 350

Anthomyia canalicularis as a human parasite .... 271

Arachnid parasites, various families of . . . .5

Argulidm, a family of parasitic crustaceans . . . .6

Arhynchotania critica of the hyrax ..... 403

Arhynchotceniada, considered as a group . . . .4

Armadillo, parasites found in the ..... 321

Ascaridce, a family of round worms . . . . .4

Ascarides infesting cetaceans . . . . , 426

„ (Oxyurides) in relation to cleanliness .... 232

Ascaris alata, Bellingham's so-called ..... 237

„ Cornelyi from the vulturine pintado . . . . 447

„ halicoris from the Indian dugong .... 429

490

INDEX

PAGE

Ascaris infecta from Passalus cornutus .... 483

„ lumhricoides as a human parasite .... 243

» „ Davaine's experiments with . . . 244;

» }, development of the .... 244

« „ Heller's " find " respecting the young of . . 244

»j >, remarkable cases caused by the . . 248

„ maculosa causing avian epizooty .... 411

n >y Unterberger's experiments with . . . 245

„ megalocephala and A. lumhricoides not identical . . . 243
» „ experiments with the eggs of . . .246

m » of solipeds ..... 378

„ mystax considered as a human parasite .... 237

» „ Leuckart's experiments with .... 241

„ „ researches of Nelson respecting .... 240

„ nigrovenosa, parthogenesis of . . . . , 452

„ suilla considered as a synonym .... 243

» >, of the hog, a synonym ..... 405

„ tentaculata of opossums ..... 433

„ tetraptera, development of the embryos of . 215

„ vesicularis from the ring-necked pheasant . . , 412

Aspidocephalus scoleciformis of opossums .... 433

Aspidocotylus mutabilis, a parasite of fish .... 360
Ass, frequency of aneurism in the . . . . .367

„ liver-fluke frequent in the ..... 356

„ strongyles from the stomach of the .... 383

Australian entozoa, Mr Krefft's brochure on . . . . 431
„ hedgehog, tapeworm from the . . . .433

Avian hasmatozoa, observations by Lewis on . . . . 486

„ parasites in the British Museum .... 448

„ „ in the Hunterian Museum . . , 448

Bacteria, the best known forms of . . . 277

BacteridcB, as a family of protozoa . . . . 7

Balantidium {Paramecium) coli of man . . . 282

„ semiparasitic character of . . . .7

Balana, flukes occurring in the genus . . . .421

„ rostrata infested by Ascaris angulivalvis . . . 426

Balanoptera rostrata, filarise from ..... 425
„ „ fluke from . . . . .421

Bats, entozoa and ectozoa infesting ..... 294
Bear, Cysticerci in the heart of a . . . . 140

Beef, cyst-infected, quantity destroyed in India . . . .76

„ in India, prevalence of " measled " . . . .61

„ tapeworm, description of the so-called . . . .56

Bcc, parasite of the, known to John Hunter .... 480
Beetles (Passalus) infested by nematodes .... 4S3

Beluga leucas, worm from the ear of . . . . . 427

Bilharzia hcematobia, history and development of the . . .38

Birds, frequency of entozoa in . . . . .434

„ Spiroptera helicina in the feet of ... 440

INDEX

491

PAGE

Birds, the gape disease in . .• • • • • 443

Blackbirds, tapeworms destructive to young .... 440

Blackcap, flukes reared by Zeller in the .... 436

Blackfish or tursio, cestode from the ..... 422

Bladderworms and measles in cattle . . . . .62

Blaptida as parasitic insects . . . . .7

Blaps mortisaga as a human parasite ..... 270

Bleak, the (Leuciscus), a possible source of tapeworm . . . 301

„ the, in relation to Bothriocephalus lotus .... 470

Bonito, parasites from the ...... 458

Bopyridce, a family of parasitic crustaceans . . . .6

Bothrocephali of water-birds obtained from fish .... 468

„ researches of Bertolus and Duchamp on . . 110

Bothriocephalus cordatus, brief description of . . . Ill

„ cristatus, brief description of . . . . Ill

„ in Holland, Dr. Fock's remarks on . . 109

„ latus, distribution and characters of . . 106

„ „ source and proscolices of . . , , 107

„ proboscideus of the salmon .... 468

„ tropicus, note on the so-called . . . .96

Bot infesting the stomach of the rhinoceros .... 401

Bottle-head {Hyperoodon), flukes from the .... 421

Brain, Cysticeroi occurring in the human . . . .92

Bronchi of whales, worms in the ..... 425

Bucephalus probably a larval state of Gasterostoma . . . 462

„ regarded as a germ -sac ..... 453

Bug, the common, as a human parasite .... 273

Cachalot, cestode from the high-finned .... 422

Calceostoma, hooks of the suckers of . . . . . 464

CaligidcB, a family of crustaceans . . . . .6

Campula oblonga regarded as a synonym .... 419

Capercaillie, entozoa of the ...... 44Q-

„ Trichosoma and Ligula from a . . . . 447

Carnivora, internal parasites of the ..... 29'/

Cataphractus infested by Aspidocotylus .... 360

Cat, Australian, Bothriocephalus from .... 30f>

„ flukes and tapeworms infesting the .... 308

Cattle diseased by parasites, natural cure of . . .83

„ measles and bladderworms in . . . . .61

„ of the Upper Punjab infested by cysts, percentage of . .75

„ plague bodies, nature of the so-called .... 280

„ species of lice infesting ..... 35a

„ tapeworms found in . . . . . , 332

Cercaria diplocotylea of water-snaila ..... 452

„ of the common fluke ..... 325

Cercarice and Redice, Pagenstecher's remarks on ... 452

Cercomonas hominis in cholera dejections «... 282

Cestoda, families of the order . , . . .4

Cestode (larval) in Delphinus delphis ..... 422

■!<J2

INDEX

PAOB

Cetacea, external parasites of . . . . . . 428

,, pai-asites of ..... 416

„ „ in Hunterian collection .... 427

„ worms in the lungs of .... 423

„ „ in the cranial sinuses of ... 425

Chacma, strongyles found in the ..... 291

Charbon, an anthracoid disease associated with bacterids . . . 278

Cheetah, tapeworm found in the ..... 300

Cheiracanthidce, a family of round worms . . . .4

Cheiracanthus robustus in Indian dogs .... 305

„ „ mistaken for JSchinorhynchus . . .257

Cheiroptera or bats, parasites of . . . . 293

Chigoe or jigger as a human parasite ..... 274

Chinese missionary, flukes in the family of a . . .21

Cirrhipeds, parasitic, upon whales ..... 428

Civets, parasites found in . . . . . 299

Classification of Schneider referred to . . . . .4

Clepsinidce, a family of suctorial annelids . . . .5

Coati, intestinal worms of the ..... 298

Cochin-China diarrhoea due to Leptodera .... 235

Cockchafers harbor young Echinorhynchi . . . .413

Cockroaches (Blatta) infested by nematodes .... 483

Codfish, frequency of filarise in the ..... 472

Caenuri from rabbits in Guy's Museum and at Oxford . . . 140

„ researches of Rose and Numan respecting . . . 334

Ccenurus cuniculi, specimen of, from Ayrshire .... 140

„ serialis of the hyrax ..... 403

Conocephalus typicus from a dolphin ..... 426

Conopidce, the larvse of, as parasitic insects . . . .6

Conorhinus nigrovarious or pampas benchucha .... 273

Cotylogaster cochlear if or me not an equine parasite . . . 360

Crane, entozoa from a ..... 447

Crossophorus collaris of the hyrax ..... 403

Crocodile, worm from beneath the skin of a . . . 456

Crows, hsematozoa in Indian ..... 486

Crustacea, parasitic, upon whales ..... 428

„ the, as alleged human parasites .... 269

Crustacean parasites, families of . . . .6

Crustaceans, entozoa infesting ..... 480

CucullanidcB, a family of round worms . . . .4

„ development of the ..... 474

Culex, various species of, attacking man .... 273

Curlew, entozoa from the ...... 447

Cuterebra noxialis, or Macaco worm ..... 271

Cuttle-fishes, tapeworm larvse found in .... 46S

Cyamidce included with Pycnogonidse . . . . .5

Cyamus balcenarum or whale-louse ..... 428

Cyclobdella lumbricoides, the, of Brazil .... 259

Cyclops considered as an intermediate host . . . . 828

Cymothoidce, a family of parasitic crustaceans . . . .6

INDEX 493

PAGE

"Cyst' -affected meat in the Punjab, quantity of . . .76

Cystic disease in cattle, Dr J. Fleming on . • .76

„ entozoa are larval tapeworms . . .• • .4

Cysticerci, common situation of, in ration meat . . . .78

„ destruction of, by calcareous degeneration . . .83

„ detected in the living animal . . . .78

„ from sheep, Dr Maddox on . . . . .98

„ from the nasal sinuses of a porpoise . . . .421

„ from the skin of Physeter ..... 421
„ in the heart of a bear ..... 140
„ in the human body, seat of . . . .91

„ in the psoas muscles, Dr Joseph Fleming on . .75

„ in meat, Pelizzari's and Tommasi's researches on . .62

„ investigations of Lewis respecting . . . .66

„ monstrous varieties of . . . . 105

„ of the sheep alleged to contain eggs . . . .98

„ prophylactic measures respecting . . . .83

„ researches by Perroncito on . . . .67

„ said to be capable of swimming . . . .97

„ temperature necessary to destroy . . . .69

„ voluntarily swallowed by Italian gentlemen . . .71

Cysticercus bovis found in the liver and lungs . . . .59

„ „ human feeding experiments by Oliver with . .72

,, delphini, nature of the so-called .... 422
,, found in the dog ..... 302

„ from the Eed River hog ..... 405
„ in the brain, cases of . . . .92

,, of the sheep, discovery of the . . . .97

„ report by Dr Neill respecting . . . .80

„ (tela) celluloses, or pork measle . . . .89

„ tenuicollis found in man ..... 101

„ „ , the self -feeding experiment by Moller with . . 72

Cysts at the root of the tongue, diagnostic value of . .80

„ of the liver containing psorospermise .... 282

Dactylogyrus, allusion to the eggs of . . . . .42

Dasse (Hyrax), parasites of the ..... 403

Dasyprocta aguti, stomach excrescences in . . . 403

Deaths from worms, Registrar General's returns of 285

Deer, abundance of amphistomes in . . . . 332

,, filarise found in various kinds of .... 350

Delphinus Forsteri, tapeworm found in ... 422

„ phoccena, cestode (Diphyllobothrium) from . . . 422

„ rostratus, tapeworm from ..... 422

„ tacuschi, flukes found by Natterer in . . . . 417

Demodex folliculornm of man and dog .... 266

Dermatophagoides Scheremetewsky, the so-called . . . 266

Dibothridce and Bothriocephalidce as synonyms ... 4

Dibothrium hians, supposed monstrous state of . . . . 105
Dichelestidae, a family of crustacean parasites . . . .6

494

INDEX

PAGE

Dioolyles (Peccary), parasites of the genus .... 404

Dicrocaelium Buskii as a synonym . . . . .20

Didelphys philander, acanthocephalous worm of ... 434

Diphyllooothridce, a family of tapeworms . . . .4

Diphyllobothrium stemmacephalum from a porpoise . . . 422

Diplodiscus suhclavatus, water-vessels of the larvae of . .51

Diplozoon in the Diporpa condition ..... 464

Diseased meat from " rot " . . . . . . 331

Distoma Andersoni from Flatanista ..... 420

„ campula of the porpoise ..... 418

„ clavatum, particulars respecting .... 458

„ conjunctum as a human parasite . . . .30

„ crassum or Busk's fluke . . . . .20

„ cygnoides, ciliated emhryo of . . . .49

„ gigas of Nardo, description of the . . . 460

„ hepaticum a synonym of Fasciola . . . .15

„ heterophyes, description of the . . . .34

„ lancea infesting dolphins ..... 416

„ lanceolatum as a human parasite . . . .17

„ „ ciliated embryo of . . .49

„ „ description of the . . . .18

„ leptosomum and D. bracJiysomum, source of . . 438

„ macrostomum, development of . . . .435

„ megastoma, ciliated embryo of . . . .49

„ militare, development of . . . . 436

„ neuronaii Munroii, the so-called . . . .52

„ ophthalmioiium,, notice of the so-called . . . .36

„ sinense, or the Chinese fluke . . . . .29

„ species of, in Delphinus Forsteri .... 421

Distomata confounded with proglottides . . . .16

Distomidce, a family of flukes . . . . .4

Distomum spatulatum as a synonym . . . . .28

Dochmius duodenalis, discovery of, by Dubiui . . . .211

„ Sangeri from the elephant ..... 399

Dogs, destruction of dogs by heart-worms .... 304

Dog, ectozoa and follicle mites of the ..... 307

„ internal parasites of the ..... 300

Dolphin, remarkable worm from the stomach of a . . . 426

Dolphins, fluke parasites of . . . . . .416

Dorado, Fasciola fusca from the ..... 459

Dracunculus, description of the embryos of . . . 221

„ Fedschenko's discovery respecting .... 223

„ medinensis, description of the . . . .217

Duck, worms from a dusky . . . . . .417

Dugong, parasites of the ...... 429

Ear of the porpoise, worms found in the . . . .427

Earth-wolf, remarkable parasite from the .... 299

Earth-worms as intermediate bearers . . . . .846

Earwig, Filarial found in the ..... 483

INDEX

495

PAGE

Echidna, tapeworm from the ..... 433

Echinococci described by Leuckart and Naunyn . . . .117

Echinococcus brood-capsules known to Wilson and Busk . . . 117

Echinococcus hominis, synonyms of . . . . 112

„ multilocularis found in a calf. .... 116

Echinorhynchidm abound in fishes and reptiles . . . .5

Echinorhynchi found by John Hunter ..... 427

„ infesting monkeys ..... 292

„ in trout, frequency of . . . . . 475

„ of water newts ..... 455

Echinorhynchus angustatus and E. hominis .... 413

„ gigas as a human parasite .... 256

„ „ of the hog ..... 412

„ hominis, Lambl's so-called .... 256

„ in man, Welch's supposed case of 256

„ Leuckart on the development of 476

,, microcephalus from an opossum .... 434

„ Muriei from a whale ..... 427

„ pellucidus and E. brevicollis .... 428

„ porrigens from whales .... 427

„ spirula of monkeys ..... 413

„ transversus in birds ..... 446

Echinostoma hispidum from a sturgeon .... 462

Ectozoa of swine ....... 414

of the elephant ...... 399

„ of whales ....... 428

Edentata, parasites of the ...... 321

Eel, Echinorhynchi in the ...... 475

Eggs, entozoa found in the interior of . . . . . 440

Elephants destroyed by " rot " ;n Burmah . . . . 394

„ earth-eating habits of worm-infected .... 395

„ ectozoa of ..... 399

„ in England destroyed by worms .... 399

„ parasites infesting ..... 393

„ parasitic diseases of .... 397

Entozoa do not result from diseased states ... 1
„ of game birds, list of .... _ 440

„ their relation to the helmintha ... 8

Epidemics of rot disease ..... 327

Epizoa, haustellated crustacean parasites ... 6

Epizooty amongst elephants from flukes . . . 394

„ amongst pigeons ...... 441

„ in birds, very destructive to life . . . 435

„ in cats, Dr Romano's account of . . 393

„ in swine from Echinorhynchus . . . 4^3

„ in tenches from LigulcB .... 469

„ in the hog from Stephanurus . . . 4^^

,. in the Mauritius from worms . . . 332

„ in the pig from Strongyli ... 412
„ in young blackbirds from tapeworms ... . 44Q

496

INDEX

PAGE

Euatrongyli within the skull of water-turkeys .... 446

Eustrongylus gigas, description of . . . . 207

n >, in dogs and wolves .... 305

» „ source of the . . '. . . 208

„ „ wild animals infested by . . . 207

„ papillosus from a crane ..... 447

Eye-worms in cattle ...... 349

Eye of the fowl infested by filarife ..... 440

Fasciola clavata, description of the so-called .... 459
„ hepatica as a human parasite . . . .14

„ „ as a " zoological " individual .... 325
„ „ ciliated embryo of . . . . .48

„ Jacksoni infesting elephants .... 397
Fauna, parasites form a peculiar . . . . .2

Ferret, ectozoon infesting the ears of the .... 307

Filaria acuta in the limbs of birds ..... 447

„ Bancrofti, history of the discovery of . . . . 180
„ gracilis infesting monkeys . . . . .291

„ hominis oris, description of the .... 206

„ horrida from the American ostrich .... 447

„ immitis in the heart of dogs ..... 304

„ inflexicaudata from Balcenoptera .... 425

„ „ of the porpoise .... 425

„ labialis, description of the ..... 206

„ lacrymalis as a synonym ..... 206

„ lentis, notice of the so-called ..... 205

„ loa, notice of the so-called ..... 205

„ macropodis gigantei in the Hunterian Museum . . . 433

„ Mansoni in the eye of a fowl ..... 441

„ medinensis considered as a synonym .... 217

„ microstoma and F. megastoma, temperature necessary to kill . 70

„ oculi considered as a synonym . . . . . 383

,, papillosa hcematica cards domestici .... 184

„ „ in the horse ..... 383

„ Salisburyii considered as a synonym .... 187

„ sanguinis, Bancroft's account of the .... 189

„ „ equi, the so-called ..... 384

„ „ hominis, correction respecting .... 487

„ „ „ discovery by Lewis of 183

„ spelcea from a wallaby ..... 433

„ (Strongylus) bronchialis, description of . . . . 207

„ terebra in the black-tailed deer .... 349

„ trachealis, remarks on the ..... 207

„ Websteri proposed as a synonym .... 433

Filarice as a cause of abscess and erysipelas .... 487

„ discovery of microscopic, by Wucherer .... 183

„ hajmatozoal, of Grube and Delafond .... 303

„ in human blood, Sonsino's verification of 185

,, of the mosquito, Hanson's description of 194

INDEX

497

PAGE

FilaridcB, a family of round worms . . . . . &

Fishos largely infostod by EchinorhynoU .... 477

„ of Brazil, amphistomos in tho ..... 463

„ parasites of ...... 457

Flea, the common, of man ...... 274

Fluke, description of the common liver . . . .15

„ description of the largo human . . . . .24

„ origin and meaning of the term . . . .4

Flukes abundant in the great kangaroo .... 431

„ causing " rot " in elephants ..... 394

„ found in fishes ...... 457

„ found in ruminating animals ..... 323

„ Pagenstecher's researches on . . . • 452

„ their destiny favoured by mimetism .... 436

Fox, fluke of the American red ..... 299

,, source of tapeworms infesting the .... 300

Fowl, entozoa infesting the common . 440

Free nematoids, Villot's opinion respecting the .... 483

Frog, Asoaris nigrovenosa of the • . . . . . 452

Galeodes araneoides or canal tick ..... 267

Gallinula chloropus, worms in the limbs of . . . 447
Gamasidce, a family of Arachnida . . . . .5

Game birds, list of entozoa infesting ..... 440

Gammarus pulex as an intermediate host . . . 476

Gape disease, methods of preventing and treating the . . . 445

" Gapes " in birds due to Selerostoma ..... 443

Gasterostoma gracilesoens, probable larval condition of . .52

„ peculiarities of the genus .... 462

Gastrodiscus Sonsinonis of the horse ..... 359

Gastrophilus rhinocer otitis, a bot-fly ..... 401

Gid hydatid found in various animals ..... 333

„ in the hog noticed by Florman ..... 405

Giraffe, large fluke found in the ..... 323

Glossina morsitans as a human parasite .... 273

Glow-worm (Lampyris), correction respecting the . , . 483

Gluttons, internal parasites of the . . . . . . 298

Gnathostoma hispidum of the hog ..... 412

Goat, the beef-measle discovered by Zenker in a . . . .83

Gongylonema pulchrum of the hog ..... 412

Goose, worms from the ashy-headed ..... 447

from the Sandwich Islands .... 447

Gordiidce, a family of round worms . . . . .5

Gordius survives desiccation . • . , , 433

Grebe, worms found in the legs of the lesser .... 447

GregarinidcB, as a family of protozoa .... 7

Grouse-disease, parasites producing the . . . 433
„ entozoa of tho red .....

Gubleria proposed as a generic title .... 7

Guillemot, entozoa from tho .

408

INDEX

PAGE

Gulls, entozoa found in . . , , m 447

Guinea-worm, development of the ..... 223

„ geographical distribution of the . . . .218

Oymnorhynchus horridus as a synonym . . . .471

Gynacophorus hcematolius considered as a synonym . . .39

Gyrodactylidce, a family of flukes . , . . .4

Gyrodactylus, conclusions of Wed! respecting .... 465

„ elegans, development of .... 466

Haddock, frequency of filarise in the . . . . . 472

Habnaturus Derbyensis, nematodes of .... 434

Hams, Cysticerci in Westphalian . . . . 405

Hares and rabbits, entozoa of . . . . .318

Hawk, entozoa from the sparrow ..... 447

Hcennatomyzus (Idocoloris) elephantis ..... 399

Scematopinus of the hog ...... 414

Hcematopota pluvialis or the clegg ..... 273

Haematozoa in dogs, discovery by Grube and Delafond of . . . 184

„ in Egyptian and Indian crows . . . . . 486

„ of the dog ...... 303

Hematuria caused by Bilharzia hcematobia . . . .54

Hcementeria, American leeches of the genus .... 259

Ecemopis sanguisorba attacking French soldiers .... 258

Heart of whales, worms in the . . . . . 425

Hedgehog, parasites of the common ..... 295

„ tapeworm from the so-called Australian . . . 433

Helmintha, the term in relation to " worms " . . . .8

Helminthology, practical results from the study of . .3

Melophilus, larva of, found in man ..... 273

„ larvae found in the horse ..... 387

Herring, frequency of filaria in the ..... 472

Eesslingia proposed as a generic title . . . . .7

Heterocheilus tunicatus from Natterer's manatee . . . 429

Heteroura androphora, sexual peculiarities of . . . 445

Hexathyridium pinguicola and H. venarum . . . .36

Eippoboscidce contain parasitic insects . . . .6

Hippopotamus, parasites of the ..... 402

Histiocephalus, a genus of nematodes ..... 433

Hog, absence of tapeworms in the ..... 404

„ cholera, supposed connection with Stephantirus . . . 410

„ skin affection from parasites in the wild . . . .414

Hog's flesh, supposed injuriousness of . . . . . 401

„ intestines perforated by parasites .... 413

Hogs, Stephanurus discovered in a Chinese race of 408
Horse, aneurism in the, from parasites .... 368

„ a thousand amphistomes found in the .... 358

„ bladderworms found in the ..... 366

„ bots or larval gadflies of the ..... 385

„ earth-eating habits of, in relation to worms . . . 357

„ ectozoa luiblo to attack the ..... 388

IN DUX

499

PAOE

Horso, epizooty from strongyles in the .... 374

,, eye-worms from the 383
„ giant strongyle found in tho ..... 385

„ htematozoa discovered hy Sonsino in the . . . . 384

„ hydatids from the 366
„ internal parasites of the ..... 357

„ plague, parasites not the cause of the .... 360

„ poultry lousiness of the ..... 388

„ psorospermial sac in the mitral valve of a . . . 281

„ scab or mange mites of the ..... 389

„ Sonsino's amphistomatoid fluke from the . . . 359

„ stomach-worms of the ..... 380

„ tapeworm epizooty in the ..... 361

„ tick, the so-called ...... 388

„ worm-aneurism of the ..... 368

Husk in young pigs caused by strongyles .... 412

„ or parasitic bronchitis in cattle ..... 335

„ or verminous bronchitis in dogs ..... 304

Hydatid disease, deaths in Victoria from .... 287

„ possibility of stamping out .... 141

Hydatids as the source of Tcenia echinococcus . . . .113

„ exogenous, endogenous, and multilocular . . .115

„ frequently infest the head of the tibia .... 129

„ hygienic considerations respecting . . . .126

„ in animals, Dr Cleghorn's statistics of . . . . 124

,, in Guy's Hospital Museum ..... 135

„ in King's College Anatomical Museum . . . 130

„ in Royal Veterinary College Museum .... 139

„ in St Bartholomew's Hospital Museum . . . 128

„ in St George's Hospital Museum .... 133

„ in St Mary's Hospital Museum .... 129

,, in St Thomas's Hospital Museum .... 134

„ in the Brighton and Sussex Hospital Museum . . . 136

„ in the Calcutta Medical College Museum . . . 137

„ in the Cambridge Anatomical Museum . . . 136

„ in the Charing Cross Hospital Museum . . . 131

„ in the London Hospital Museum . . . .133

,, in the Melbourne Hospital Museum . . . .141

„ in the Middlesex Hospital Museum . . . .130

„ in the Norfolk and Norwich Hospital Museum . . . 137

„ in the Oxford Anatomical and Pathological Museum . . 136

„ in the Royal College of Surgeons Museum . . .132

„ in the Westminster Hospital Museum .... 128

„ in University College Museum . . . .131

„ in Victoria, number of deaths from .... 123

„ mortality in England from . . . . , 285

„ occupying the bones of cattle, specimens of . . 140

„ occupying the heart of a sow .... 139

„ occupying the nasal sinuses of a porpoise . . , 421

„ of animals in metropolitan museums .... 139

500

INDEX

Hydatids of tho tibia in the Nottingham Hospital Museum . . 130

„ preserved in museums, their value . . . 128

» prevalence of, in Australia .... 123

„ probable percentage of fatal cases of . . . j 22

„ successful treatment of, by MacGillivray . . 128

„ their distribution in the organs of the body . . 122

,, the so-called daughter vesicles of . . 121

Hydrachnidce, a family of Arachnida ... 5

Symenolepis, as a new genus of cestodes . . . 102

Syperoodon, Echinorhynchus found by Hyrtl in ... 427

Hyperemia and Anosmia due to parasites . . 213

Hyrax, parasites of the Cape .... 403

Ichneumon, the Indian, fluke from the . . . 299

Insectivora, entozoa found in the .... 295

Insect parasites, various families of . . , g

Insects, entozoa of .... 480

Intestinal worms, popular ideas respecting ... 8

Itch or scab insect, description of the .... 266

Ixodes, species of, attacking man .... 267

IxodidcB, a family of Arachnida .... 5

Jigger or chigoe as a human parasite ..... 274

Kaleege, worms from the black-backed .... 447

Kangaroo, tapeworm in the gall-bladder of a . . . 432

„ worms from the knee-joint of a . . , 433

Kestrel, entozoa from the ...... 447

Kite, entozoa from the ...... 447

Klipdas {Hyrax), parasites of the ..... 403

Koala (Phascolarctos), tapeworm from a 432

KolliJceria filicollis, habitat of .... 462

Lagenorhynchus Esohrichti, worm found in . . . , 421

Lambs, tapeworm epizooty in .... 333

Lampyris, correction of an error respecting .... 483

Larvae of flukes, Pagenstecher's researches on . . . 452

Leeches considered as human parasites .... 258

„ of India, Ceylon, Japan, and Java .... 258
„ semiparasitic character of . . . .5

Lemurs, ccenuri or polycephalous hydatids in . . . 290
Leporines, acarine parasites of . . . . .319

Leptodera (Anguillula) stercoralis, description of 234

„ intestinalis, description of the .... 235

Leptus autumnalis a human parasite ..... 268

Lernsean parasites found on whales ..... 428
Lernceidce, a family of crustacean parasites . . . .6

Leuciscus, the bleak, a possible source of tapeworm , . . 108

Leucochloridium, experiments by Zcller with . . . , 486
Lice, various species found on man . . . . .271
Ligula nodosa, as an immature Bothriocephalus . . . .110

INDEX

501

PAGE

Ligula nodosa, the young state of Bothriocephalus . . . 470

„ simplicissima, researches by Duchamp on 469

Lindermannia proposed as a generic title . . • 7

Ling, Echinorhynchi in the ...... 475

Lion, hydatids from the, in the Hunterian Museum . . . 139
Liotheidce, a family of parasitic insects . . . .6

Liver fluke, the common, in man . . . . » 14

Louse of the elephant ...... 400

Lucilia hominivora, habits of the ..... 272

Lumbrici, geographical distribution of ... 247

„ in the Mauritius, remarks by Dyce on 247

„ remarkable cases of, in children .... 248

„ violent symptoms caused by ... 249

Lumbricoid worms, poisonous effects produced by 250

Lungs of the porpoise, worms in the ..... 425

Maggots, rat-tailed, found in the horse .... 387

„ various species of, found in man .... 272

MalacobdellidcB, a family of suctorial annelids . . . . 5

Manatees, parasites of the ...... 429

Marmoset monkeys, parasites of . . . . 291

Marsupial animals, parasites of . . . 430

Martens, internal parasites of .... 298

Masuri and soorti, parasites of elephants .... 395

„ or amphistomes occur in man . . . .38

Measle-cysts in the Punjab, report respecting . . . .75

Measled cattle cured by removal from sources of infection . . 83

„ meat, Giacomini's observations on . . .65

Measles and bladderworms in cattle . . . . .61

„ and Tcenice, observations of Marchi on . . . .73

„ found in mutton brought to table . . . .97

„ of pork reared by experiment . . . .93

Meat question, the, in relation to " rot " 331
„ served as rations, Cysticerci in the . . . .78

Melolontha considered as an intermediate host .... 257

„ vulgaris as an intermediate " host " . . . . 413

Melophagus, a genus of parasitic insects . . . .6

Mermis albicans, development of . . . . 482

Messmates and parasites of mollusks ..... 480

Miana bug of Persia, or Argas ..... 267

Mice, internal parasites of . . . . . . 315

Micrococci considered as human parasites .... 277

Micropteron (Delphinus), flukes in . . . . 421

Mimetism favoring fluke development .... 436

Mite found upon Balcena australis ..... 428

Mole, parasites of the common ..... 296

Mollusits, entozoa of . . . . . 480

Monads, parasitic forms of . . . . . . 282

Monkeys, hydatids obtained from ..... 141

„ parasites of ..... 289

502

INDEX

PAOB

Monodon monoceros, nematode worms of . . . 423

Monostoma plicatum from a northern whale .... 421
„ verrucosum, allusion to the eggs of . . . .41

Monostomidce, a family of flukes . . . . .4

Mosquito, avian hsernatozoa in the ..... 486
Mosquitoes of Queensland, filaria3 in the .... 4S7

Mules, fatal epidemic affecting ..... 382

Musca, larvae of, as human parasites ..... 272
Musk-rat, cestode parasite found in the .... 29G

Mutton measles, their existence denied . . . .97

Narwhal, Prosthecosacter infesting the .... 423

Nematobothrium filarina, habitat of . . . . . 4G2

Nematoda, families of the order of . . . . .4

Nematodes abundant in marsupials ..... 433
Neinatoids, opinion of Villot respecting the " free " . . 484

Nestlings destroyed by tapeworms ..... 440
Newts, Echinorhynchus anthuris of water . . . .455

Nycteribiidw as parasitic insects . . . . .6

Octobothrium lanceolatum, allusion to the eggs of . . .41

Odontobius ceti from Balcena australis .... 427

(Estridce, the, a family of parasitic insects . . . .6

„ infesting the ox and sheep ..... 351

(Estrus (Gastrophilus) equi of the horse .... 385

„ hominis, Mr Higginson's case of . . . 271

Olulaniasis of the cat . . . . . .310

Onchocerca reticulata of the horse ..... 385

OncTiotyle appendiculata from a dog-fish .... 464

Opossum, Echinorhynchus in a Brazilian . . . . 434

Opossums, flukes infesting American ..... 432

„ much infested by nematodes .... 433

Orcella brevirostris, fluke found by Dr Anderson in . . 417

Orthagoriscus mola as a " host " . . . . .471

Ostrich, entozoa from the American ..... 417

Otters, flukes and other entozoa of . . . . 298

Ox, insect parasites and tormentors of the .... 351

„ species of Strongylus infesting the .... 347

Oxyuridce, a family of round worms . . . . .5

Oxyurides, sanitary precautions in respect of . . . 232

„ source and mode of development of . . . 230

„ voluntary swallowing of human . . . .72

Oxyuris curvula of the horse ..... 380

„ vermicularis, description of the .... 227

Oysters as bearers of fluke^larva3 . . . • .26

Pachydermata, parasites of the ..... 393

Palisade worms in relation to aneurism .... 368

Paramecium (Balantidium) coli of man .... 282

Paramecium, semiparasitic character of . • -7

Parasites from birds at the Zoological Gardens .... -117

INDEX

503

PAGE

Parasites from cetacea in the Hunterian Museum . . . 427

„ in grouse from the Earl of Cawdor's estate . . . 439

„ of savages, little known respecting . . . .27

„ popular delusions concerning . . . .1

„ their etiological significance not fully recognised . . 488

Partridge, entozoa of the ...... 440

Peccaries (Dicotyles), parasites of the .... 404

PedictdidcB, a family of parasitic insects . . . .6

Pediculus, various forms of, infesting man .... 274

Pentastoma constrictum, description of the .... 263

„ tcenioides as a human parasite .... 260

n „ found in the horse .... 389

„ ,, in the dog ..... 306

Pentastomes infesting monkeys ..... 293

Pentastomidce, a family of Arachnida . . . . .5

Perch, flukes from the eye of the ..... 458

Peritrachelius insignis from "a dolphin .... 427

Pheasant, entozoa of the ...... 440

„ parasites from various species of . . . 447

Philopteridce, a family of parasitic insects . . . .6

Phoecena communis, flukes found in . . . . 418

Phyllohothrium delphini, source of . . . . 422

Pigeon, two hundred ascarides in a . . . . 442

Pig, hydatids from the, in the Hunterian Museum . . . 139

„ occurrence of Cysticerci in the brain of the . . . 405

Pigs, the cause of " husk " in young ..... 412

Pike-whale, large fluke from the . . . . .421

Pike, worms from the urinary bladder of the .... 458

Pinnipedia, parasites of . . . . ,314

Pintado, ascarides in the vulturine ..... 447

Physaloptera turgida infesting opossums . . . .433

Plagiotcenia gigantea of the rhinoceros .... 401

Plagues, parasites considered as a cause of . . .2

Platanista gangetica, fluke from ..... 420

Podiceps and Oallinula, legs of, infested by filarial . . . 447

Polecats, filarial in the cranial sinuses of ... 298

Polystoma and Amphistoma of the frog .... 452

„ appendiculata, allusion to the eggs of . . .41

Polystomes, general characters of the ..... 463

PolystomidcB, a family of flukes . . . . .4

Ponies, Welsh mountain, epizooty affecting .... 362

Porcupine ant-eater, tapeworm from the .... 433

Pork tapeworm, general description of the . . . .87

„ sources of infection by the . . . .94

Porpoise, flukes found in the common ..... 418

„ remarkable cestode from the common .... 422

,, worm from the tympanic cavity of the . . . 427

Porrigo, mortality in England from ..... 285

Poultry lousiness not a human disorder .... 274

Proglottides mistaken for Dislomata . . . . .16

504

INDEX

Prosthecosacter, several species of

» specimens of, in Hunter's Museum

Proteles or earth-wolf, parasites of
Protozoa, parasitic families belonging to the

n - parasitic forms of, and gregarinsB
Psorospermies as a family of Protozoa .

», considered as human parasites

Psorosperms abundant in the hog

„ in the flesh of animals
Pulex penetrans, gigger or chigoe
Pulmonary vessels of whales, worms in the
Pycnogonidce, a family of Arachnida .

Quadrumana, entozoa and ectozoa of .
Quail, entozoa of the

Queensland, Bancroft's collection of entozoa in

Rabbits and hares, internal parasites of
„ attacked by the autumnal spider
„ Cosnuri found in the soft parts of

Racoons, parasites infesting .

Rats, internal parasites of .

Redim and Cercarice, Pagenstecher's remarks on

Redshank, entozoa from the

Reduvius, or fly bug, various species of

Redwing, EchinorhyncM from a

Reptiles, internal parasites of

Rhea Americana, filarise obtained by Darwin from

Rhinoceroses, parasites of

Rhopalophorus, a genus of flukes

Rhyncliotceniada considered as a group

Rhytina stelleri, parasites of

Rodentia, parasites of

" Rot " affecting the liver of porpoises
„ disease, conclusions respecting the

Round worms, various families of

Ruminating animals, ectozoa found on
entozoa of

Salmon, pit-headed tapeworms of the .
Salmo umbla, worms in the kidney of
Sanguisuga medicinalis or the grey leech
Sanitation in connection with " rot " .
Sarcoptes of the hog

„ scabiei, description of the .
Savages, little known about the parasites of
Scabies, mortality in England from
Scab or itch insect of man .
Sealops canadensis, nematode found in
Schistosoma, considered as a synonym of Bilharzia
Sclerostoma and Anchylostoma as synonyms

INDEX

505

PAGE
4)12

Sclerostoma dentatum of the nog .

„ pinguicola of Vcrrill, the so-called .... 409

• i • 3 443
„ syngamus in birds .

Seals and walruses, parasites of

Sheep, hydatids from cattle and . 139
„ of New South Wales, fatal epizooty in . . . .347

o o

„ parasitic bronchitis affecting .

„ whip-worms occasioning " scour " iu • • • 349

Shrews, parasites known to infest . 29f>
Simondsia paradoxa, description of . . • • • 406

Simulium reptans or the creeping gnat .
Skunk, tapeworm infesting the

Sloths, nematode worms found in . • • • 322

Snake-birds, parasites in the cranium of 446
Snake, larval nematode from the heart of a . • • 455

„ worms from an Egyptian hooded . 456

Solidungula, parasites of the .

Solpugidce, a family of Arachnida . . • • .5

Soorti and masuri parasites of elephants . 395

Spharularia bombi, Sir J. Lubbock's description of . • 481

„ found in various bees . 480
„ Schneider's account of the development of . • 482

Spirilla, found in the blood of fever patients .... 277
Spiroptera helicina in the feet of birds . 440

„ megastoma of the horse . 3^0
„ sanguinolenta, Lewis's remarks on . . • • 303

„ sexalata infesting Dicotyles .... 407

„ strongylina of the hog . 406
Sporocysts or unorganised germ-sacs . 453
Squirrels, internal parasites of 315
Stephanurus dentatus discovered by Natterer .... 407
„ Diesing's description of . . • • • 408

„ in Australia, announcement respecting . . • 409

„ in relation to porcine epizooty . 411
„ Nattereri, proposed synonym .... 412
,, Professor Fletcher's account of . • • 411

„ rediscovered by Dr. White .... 409

Sticklebacks, Qyrodactylms from the tail of . • • 466

Stoat, parasites found in the common . . . .299

Stomoxys calcitrans, or the leg-sticker .... 273

Strongyle, meaning and origin of the term . . • .4

Strongyles, Welsh equine epizooty from .... 377

Strongylidce, a family of round worms . . • .5

Strongylus acuticaudatus from a Tinamou .... 447
„ armatus of the horse ..... 367
„ dentatus confounded with Stephanurus . . . 412

„ {JEustrongylus) gigas, found in man .... 208
„ filaria, experiments by Leuckart with . . . 337

„ „ temperature necessary to kill the . . .70

„ micrurus, or cattle lung-worm .... 335

506

INDEX

PAGE

Strongylus micrurus, original experiments with . . . 337

„ minutissimus in Algerian sheep . . . 34,5

,, Pallasii from Beluga leucas . . , ' ' 427

„ paradoxus a cause of husk . . , , 4] 2

„ pergracilis and the grouse disease . . . 439

„ tetr acanthus, larvse of . . ' . , 374

Sturgeon, Echinostoma Mapidum from a . ■ ' .- ■ • 462

Suidce, rarity of flukes in the . , . . , 404

Sun-fish, Tetrarliynchus infesting the .... 471

Swine Breeders' Association, U.S. National . . . .412

„ external parasites of . . , , . 414

„ internal parasites of . . . , m 404

Sword-fish, parasites of the ..... 453

Sydney, entozoa collected hy Krefft and Masters near . . . 431

Syngamus (Sclerostoma) trachealis in birds . . . 443

Syrphida include rat-tailed maggots .... 7

Tachyglossus setosus, tapeworm found in . . 433

Tacuschi, parasites of the so-called ..... 417

Tapeworm, Cullingworth's monstrous variety of 103
„ from the vulpine opossum . . , . . 432

„ in the Baikal, statements of Kaschin respecting . . 82

,, mortality in England from .... 285

„ the broad, derived from fish .... 107

Tapeworms infesting marsupials ..... 432

„ . injurious to avian bearers ..... 438

„ (Ligulce) causing piscine epizooty .... 469

„ of the dog ...... 300

„ symptoms occasioned by . . . . .90

„ various families of . . . .4

Tapir, parasites of the American ..... 402

Tcenia abietina, a variety of T. mediocanellata .... 102

„ acanthotrias, remarks on the so-called .... 103

„ and cysticercus, coexistence of . . . . .94

„ angulata in nestling blackbirds ..... 440

,, bipapillosa from the wombat ..... 432

„ calva and the grouse disease ..... 459

„ canina identical with T. elliptica .... 101

„ crassicollis, its source suggested by Goeze . . .57

„ „ possessing two heads .... 105

„ cyathiformis, allusion to the eggs of . . .41

„ . echinococcus, its prevalence in Iceland .... 125

,, „ reared from hydatids . . . .113

„ elliptica as a human parasite ..... 101

„ f estiva in Macropus and Halmaturus . . . .432

,, Jlavopuncta, brief description of the .... 101

„ from the Cape of Good Hope, variety of . . . . 104

,, geophiloides a new tapeworm ..... 432

„ lata of Pruner, probable synonym of . . .96

„ lophosoma, considered as u species . . . .99

INDEX

507

PAGE

Tcenia Madagascariensis, brief description of the . . • 100

„ magna of the rhinoceros . • • • • "^01

„ marginata, as a human parasite .... 100

„ tnediocanellata, description of the • • .56

„ „ Cullingworth's specimen of . .99

„ „ feeding experiments with . . .58

„ „ Mussulmans infested by . . . .78

„ megaloon, doubtful character of the .... 105

„ nana, brief description of the ..... 100

„ perfoliata, temperature necessary to kill the . . .70

„ phalangistce from the vulpine opossum . . . 432

„ phoptica, a new tapeworm . . . • • 433

„ remarkable Hunterian specimen of . • • 102

„ saginata, considered as a synonym . . . .57

„ serrata, temperature necessary to kill . . . .69

„ solium, general description of the . . • .87

„ sources of infection by . . . . .94

„ tenella, probable source of the . . . . .95

„ variabilis, allusion to the eggs of . . .41

Tetrarhynchi of sharks and rays ..... 470

Tetraphyllobothridce, a family of tapeworms . . . .4

Tetrarhynchus reptans of the sun -fish .... 472

Tenches destroyed by Ligulce ..... 469

Tetrastoma renale, notice of the so-called . . . .36

Tetrabothrium triangulare, from Delphinus .... 422

Thecosoma considered as a synonym of Bilharzia . . .39

Thorn-headed worms resemble nematodes . . . .5

Thread-worm, description of the common .... 227

Tinamou, worms from the . . . . . . 447

Tongues of living animals inspected for " cyst " . . .80

Tope, or penny dog-fish, Tetrarhynchus of the .... 471

Trachearian arachnida, parasitic families of . . .5

Trematoda, families of the order of . . . . .4

Trematode-larvsB, Pagenstecher's reseaches on . . . . 452

Trematodes infesting opossums ..... 432

Tricenophorus nodulosus with two heads .... 105

Trichina qjffinis, the so-called ..... 406

,, spiralis, discovery of, in man .... 194

„ „ description of the mature .... 153

„ „ experiments conducted in England with . . 157

,, „ in relation to diseased meat .... 166

„ „ temperature necessary to kill the . . .70

Trichinidce form a family of round worms . . . .5

Trichinosis in animals, symptoms of . . . . , 163

„ in England, an outbreak of ... 169

„ in Germany, outbreaks of .... 168

Trichocephalidse, a family of round worms . . . .5

Trichocephalus affinis in sheep ..... 349

„ crenatus of the hog ..... 406

„ dispar, description of the .... 178

508

INDEX

PAGE

Trichosoma, new species found by Solger . . . 456

Tricuspidaria nodulosa, with two heads .... 105

„ (Tricenophorus) nodulosa infesting fishes . . . 470

Tristomcs not strictly entozoa ..... 463

Tristomidce, a family of flukos . . . . .4

Trout, Echinorhynchi in the ..... 475

„ the, a probable source of Bothriocephahis . . . 470

Tsetse fly fatal to the horse ...... 388

Turbellarians, remark concerning the . . . . .8

Turbot, Echinorhynchi in the ..... 475

Turkeys, water, parasites found by Wyman in . . . . 446

Turtle, worms from a snapping ..... 455

Tympanum of whales, worms in the cavity of the . . . 425

Tyroglyphus Merieourti of Laboulbene .... 266

Urobales palustris in connection with the guinea-worm . . . 219

Udonella caligorum, peculiar habit of . . . . . 484

„ mode of development of . . . . 463

Wagtail, flukes reared by Zeller in the .... 436

Wallaby, Eilaria from the abdomen of a . . . 433

„ tapeworms observed in the ..... 432

Warblers, development of flukes in . . . . 435

Warbles formed by the larva? of (Estrus .... 351

Wasps ( Vespa), Sphcerularia in .... 481

Water-hen, worms found in the legs of the .... 447

Water- turkeys, worms in the cranium of ... 446

Weasles, internal parasites of .... 298

„ source of tapeworms in . . . . 296

Whale-lice (Cyami and Pycnogonida) ..... 428

„ (Cyamidft), zoological position of . . . .5

Whale-worms found by John Hunter ..... 427

Whales, ectozoa of ...... 428

„ internal parasites of .... 416

Whip- worm (Trichocephalus) of the hog .... 406

White-throat, flukes reared by Zeller in the .... 436

Wombat, tapeworm found in the ..... 432

Worm-aneurism, Bollinger's researches on . . . 368

Worm-like sac in the mitral valve of a horse .... 2S1

Worms are not a criterion of previous disease .... 435

„ from the heart of a guillemot ..... 447

„ infesting the Virginian opossum .... 433

„ in the knee-joint of a kangaroo .... 433

„ in the tympanic cavity of whales .... 425

„ in whales, found by John Huuter .... 427

„ mortality in England from . . • 285

„ of the kangaroo in the Hunterian Museum . . . 433

Zebra, hydatids from the liver of the ..... 366

„ „ from a, described by Huxley .... 117

PltlNTED BY J. E. ADLAED, BAETHOLOMKW CLOSE.

WORKS BY THE SAME AUTHOR.

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CATALOGUE OF ENTOZOA IN THE MUSEUM OF THE
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London, New Burlington Street.
May, 1879. .

SELECTION

peom

MESSRS J. & A. CHURCHILL'S

(lateral Catakpe

COMPRISING

ALL EE CENT WOEKS PUBLISHED BY THEM

ON THE

AET AND SCIENCE

MEDICINE

INDEX

PAGK

Acton on the Reproductive Organs . 8
Adums (W.) on Clubfoot . . .6
— (R.) on Rheumatic Gout . 19
Allingham's Diseases of the Rectum 7
Anatomical Remembrancer . . 11
Anderson (McC) on Eczema . . 19

— (McC.) on Parasitic Affec-

tions . . . .19
Aveling's Influence of Posture . . 14
Balfour's Diseases of the Heart . 16
Bantock's Rupture of Perineum . 14
Barclay's Medical Diagnosis . . 12
Barker's Puerperal Diseases . . 13
Barnes' Obstetric Operations . .14

— Diseases of Women . . 14
Basham on Diseases of the Kidneys . 8
Beale on Kidney Diseases . . .8

— on Microscope in Medicine . 11
Bellamy's Guide to Surgical Anatomy 10
Bennet's Winter and Spring on the

Mediterranean . . .17

— Pulmonary Consumption . 17

— Nutrition . . . .18
Bennett (J. R.") on Cancerous Growths 19
Berkart's Asthma . . . .15
Bigg's Orthopraxy . . .6
Biuz's Elements of Therapeutics . 12
Black on the Urinary Organs . . 8
Blakiston's Clinical Reminiscences . 11
Bose's Rational Therapeutics . . 11

— Recognisant Medicine . . 11
Braune's Topographical Anatomy . 11
Brodhurst's Orthopaedic Surgery . 6
Bryant's Practice of Surgery . . 4
Burdett's Cottage Hospital . . 15
Burnett on the Ear . . . .6
Buzzard on Syphilitic Nervous Affec-
tions . . . . . .8

Carpenter's Human Physiology . . 10
Carter (H. V.) on Mycetoma . . 20
Carter (W.) on Renal and Urinary

Diseases . . . . .8
Cauty on Diseases of the Skin . . 20
Chapman on Neuralgia . . .19
Charteris' Practice of Medicine . 11
Clark's Outlines of Surgery . . 4
— Surgical Diagnosis . . 5
Clay's Obstetric Surgery . . .13
Coles' Dental Mechanics . . .23
Cormack's Clinical Studies . . 12
Cottle's Hair in Health and Disease . 20
Cullingworth's Nurse's Companion . 14
Curling's Diseases of the Rectum . 7

— Diseases of the Testis . 7
Dalby on the Ear . . .6
Dalton's Human Physiology . . 9
Day on Children's Diseases . . 13

— on Headaches . . . .18
Dobell's Lectures on Winter Cough . 15

— Loss of Weight, &c. . .15

PAGK

Domville'sManual for Hospital Nurses ] 5
Druitt's Surgeon's Vade-Mecum . 4
Duncan on the Female Perineum . 14
Dunglison's Medical Dictionary . 22
Ellis's Manual of Diseases of Children 12
Emmet's Gynaecology . . .14
Eulenburg and Guttmann's Sympa-
thetic System of Nerves . . 19
Fayrer's Observations in India . . 4
Fergusson's Practical Surgery . . 4
Fenwick's Guide to Medical Diagnosis 12
Flint on Phthisis . . . .16
— on Percussion and Auscultation 16
Foster's Clinical Medicine . . 11
Fox (C. B.) Sanitary Examinations . 21
Fox (T.) Atlas of Skin Diseases . 20
Frey's Histology . . .9
Gamgee on Fractures of the Limbs 4

— on Treatment of Wounds . 4
Gant's Science and Practice of Sur-
gery 4

— Diseases of the Bladder . . 8

Gaskoin on Psoriasis or Lepra . . 20
Glenn's Laws affecting Medical Men . 20
Godlee's Atlas of Human Anatomy . 11
Gowan on Consumption . . .15
Habershon on Diseases of the Liver . 18

— on Diseases of the Abdo-

men . . . .18

— on Diseases of the Stomach 18

— on the Pneumogastric Nerve 18
Hamilton's Nervous Diseases . . 18
Hancock's Surgery of Foot and Ankle 6
Harris on Lithotomy . . .7
Harrison's Stricture of Urethra . 7
Hayden on the Heart . . .16
Heath's Minor Surgery and Bandaging 5

— Diseases and Injuries of Jaws 5

— Operative Surgery . . 5

— Practical Anatomy . . 10
Higgens' Ophthalmic Practice . . 22
Holden's Landmarks . . .10

— Human Osteology . . 10
Hood on Gout, Rheumatism, &c. . 19
Hooper's Physician's Vade-Mecum . 11
Horton's Tropical Diseases . . 17
Hutchinson's Clinical Surgery . . 5
Huth's Marriage of Near Kin . . 9
Ireland's Idiocy and Imbecility . 21
James' Sore Throat . . . .16
Jones (C. H.) and Sieveking's Patho-
logical Anatomy . . .10

— (C. H.) on Functional Nervous

Disorders . . 18

Jones (H. McN.) Aural Surgery . 6
— Atlas of Diseases of

Membrana Tympani 6
Jones (Wharton) Ophthalmic Medi-
cine and Surgery . . 23
Jordan's Surgical Inllammations . 6

INDEX

PAG*

Jordan's Surgical Inquiries . . 6
Leber and Rottenstcin's Dental Caries 23
Lee (H.) on Syphilis . . .8
Leared on Imperfect Digestion . . 18
Liebreich's Atlas of Ophthalmoscopy 22
Liveing on Megrim, &c. • . .18
Macdonald's (A.) Disease of the heart 16
Macdonald's (J. D.) Examination of

Water . . . _ . . .21
Mackenzie on Diphtheria . . .16
Macnamara on Diseases of the Eye . 22
Madden's Health Resorts . . .17
Marsden on certain Forms of Cancer 19
Mason on Harelip and Cleft Palate • 5

— Surgery of the Face . . 5
Maunder's Operative Surgery . . 4

— Surgery of Arteries . . 4
Mayne's Medical Vocabulary • . 22
Morris (H.) Anatomy of the Joints • 10
Ogston's Medical Jurisprudence . 20
Osborn on Hydrocele . . .7
Parkes' Manual of Practical Hygiene 21
Parkin's Epidemiology . . .23
Pavy on Food and Dietetics . . 18

— on Diabetes . . . .18

Peacock's Valvular Disease . . 16
Phillips' Materia Medica . . .12
Pirrie's Surgery . ... .4

Pollock's Rheumatism . . .19
Ramsbotham's Obstetrics . . .13
Reynolds' Uses of Electricity . . 22
Roberts' (C.) Manual of Anthro-
pometry . . . . .9

Roberts' (D. Lloyd) Practice of Mid-
wifery . . . . . .13

Roussel's Transfusion of Blood . . 5
Routh's Infant Feeding . . .13
Roy's Burdwan Fever . . .17
Royle and Harley's Materia Medica . 12
Rutherford's Practical Histology . 9
Salt's Medico-Electric Apparatus . 22
Sanderson's Physiological Handbook . 9
Sansom's Diseases of the Heart . 16
Savage on the Female Pelvic Organs 4
Savory's Domestic Medicine . . 15
Sayre's Orthopaedic Surgery . . 6
Schroeder's Manual of Midwifery . 13
Semple on the Heart . . .15
Sewill's Dental Anatomy . . .23
Shapter's Diseases of the Heart . 16
Sheppard on Madness . . .21
Sibson's Medical Anatomy . . .10
Sieveking's Life Assurance . . 21
Smith(E.) Wasting Diseasesof Children 13
— Clinical Studies . . 13
Smith (Henry) Surgery of the Rectum 8
Smith (Hey wood) Gynaecology . . 14
Smith (J.) Dental Anatomy . . 23
Smith (W. R.) Nursing . . .15
Spender's Bath Waters . . .17

PAGE

Steincr's Diseases of Children . . 13
Stille and Maisch's Dispensatory . 12
Stocken's Dental Materia Medica . 12
Stowe's Toxicological Chart . . 20
Sullivan's Tropical Diseases . . 17
Swain's Surgical Emergencies . . 5
Swayne's Obstetric Aphorisms . . 14
Taft's Operative Dentistry . . 23
Tait's Hospital Mortality . . .15
Taylor's Principles of Medical Juris-
prudence . . . .20

— Manual of Medical Juris-

prudence . . . .20

— Poisons in relation to Medical

Jurisprudence . . .20
Teale's Dangers to Health . . 21
Thomas on Ear and Throat Diseases 6
Thompson's Stricture of Urethra . 7

— Practical Lithotomy and

Lithotrity . . .7

— Diseases of Urinary Organs 7

— Diseases of the Prostate . 7

— Calculous Disease . . 7
Thornton on Tracheotomy . . 16
Thorowgood on Asthma . . .15

— on Materia Medica . 12
Thudichum's Pathology of Urine . 8
Tibbits' Medical Electricity . . 22

— Map of Motor Points . . 22
Tilt's Uterine Therapeutics . . 13

— Change of Life . . .13

— Health in India . . .17
Tomes' (C. S.) Dental Anatomy . 23

— (J. and C. S.) Dental Surgery 23
Tuke on the Influence of the Mind

upon the Body . . . .21
Van Buren on Diseases of the Genito-
urinary Organs . . .8
Veitch's Handbook for Nurses . . 15
Virchow's Post-mortem Examinations 10
Wagstaffe's Human Osteology . 9
Walton's Diseases of the Eye . . 22
Ward on Affections of the Liver , 17
Waring's Practical Therapeutics . 12
— Bazaar Medicines of India . 17
Wells (Soelberg) on Diseases of the Eye 23

— Long, Short, and Weak Sight . 23
Wells (Spencer) on Diseases of the

Ovaries . . . .14
West's Diseases of Women . . 14
Whistler's Syphilis of Larynx . . 17
Wilks' Diseases of Nervous System . 18

— Pathological Anatomy . . 10
Wilson's(E.)Anatomist'sVade-Mecum 11

— Diseases of the Skin . .19

— Lectures on Ekzema . . 19

— Lectures on Dermatology . 19
Wilson's (G.) Handbook of Hygiene . 21
Woodman & Tidy's Forensic Medicine 21

CATALOGUE OF RECENT WORKS

THE PRACTICE OF SURGERY :

a Manual by Thomas Bryant, F.R.C.S., Surgeon to Guy's Hospital.

Third Edition, 2 vols., crown 8vo, with 672 Engravings, 28s. [1878]
THE PRINCIPLES AND PRACTICE OF SURGERY,

by William Pirrie, F.R.S.E., Professor of Surgery in the University

of Aberdeen. Third Edition, 8vo, with 490 Engravings, 28s. [1873]
A SYSTEM OF PRACTICAL SURGERY,

by Sir William Fergusson, Bart., F.R.C.S., F.R.S. Fifth Edition,

8vo, with 463 Engravings, 21s. [1870]
OPERATIVE SURGERY,

by C. F. Maunder, F.R.C.S., Surgeon to the London Hospital.

Second Edition, post 8vo, with 164 Engravings, 6s. [1872]

BY THE SAME AUTHOB.

SURGERY OF THE ARTERIES :

Lettsomian Lectures for 1875, on Aneurisms, Wounds, Haemorrhages,
&c. Post 8vo, with 18 Engravings, 5s. [1875]

THE SURGEON'S YADE-MECUM,

a Manual of Modern Surgery, by Robert Drttitt. Eleventh Edition,
f cap. 8vo, with 369 Engravings, 14s. [1878]

THE SCIENCE AND PRACTICE OF SURGERY :

a complete System and Textbook by F. J. Gant, F.R.C.S., Senior Sur-
geon to the Royal Free Hospital. 8vo, with 470 Engravings, 24s. [1871]

OUTLINES OF SURGERY AND SURGICAL PATHOLOGY,

including the Diagnosis and Treatment of Obscure and Urgent
Cases, and the Surgical Anatomy of some Important Structures and
Regions, by F. Le Gros Clark, F.R.S. , Consulting Surgeon to St.
Thomas's Hospital. Second Edition, Revised and Expanded by the
Author, assisted by W. W. Wagstaefe, F.R.C.S., Assistant-Surgeon
to St. Thomas's Hospital. 8vo, 10s. 6d. [1872]

CLINICAL AND PATHOLOGICAL OBSERVATIONS IN INDIA,
by Sir J. Fayrer, K.C.S.I., M.D., F.R.C.P. Lond., F.R.S.E., Honorary
Physician to the Queen. 8vo, with Engravings, 20s.

TREATMENT OF WOUNDS :

Clinical Lectures, by Sampson Gamgee, F.R.S.E., Surgeon to the
Queen's Hospital, Birmingham. Crown 8vo, with Engravings, 5s. [1878]

BY THE SAME ATJTHOB,

FRACTURES OF THE LIMBS

and their Treatment. 8vo, with Plates, 10s. 6d. P8™

THE FEMALE PELVIC ORGANS,

their Surgery, Surgical Pathology, and Surgical Anatomy, in a
Series of Coloured Plates taken from Nature : with Commentaries,
Notes, and Cases, by Henry Savage, M.D. Lond., F.R.C.S., Consulting
Officer of the Samaritan Free Hospital. Third Edition, 4to, £1 15s.

ris75]

PUBLISHED BY J. AND A. CHURCHILL

SURGICAL EMERGENCIES

together with the Emergencies attendant on Parturition and the
Treatment of Poisoning : a Manual for the use of General Practi-
tioners, by William P. Swain, F.R.C.S., Surgeon to the Royal Albert
Hospital, Devonport. Second Edition, post 8vo, with 104- Engravings,
6s. 6d.

TRANSFUSION OF HUMAN BLOOD :

with Table of 50 cases, by Dr. Roussel, of Geneva. Translated by
Claude Guinness, B.A. With a Preface by Sir James Paget, Bart.
Crown 8vo, 2s. 6d. [1877]

ILLUSTRATIONS OF CLINICAL SURGERY,

consisting of Coloured Plates, Photographs, Woodcuts, Diagrams, &c,
illustrating Surgical Diseases, Symptoms and Accidents ; also Opera-
tions and other methods of Treatment. By Jonathan Hutchinson,
F.R.C.S., Senior Surgeon to the London Hospital. In Quarterly
Fasciculi, 6s. 6d. each. Fasciculi I to X bound, with Appendix and
Index, £3 10s. [1876-9]

PRINCIPLES OF SURGICAL DIAGNOSIS

especially in Relation to Shock and Visceral Lesions, by F. Le Gros
Clark, F.R.C.S., Consulting Surgeon to St. Thomas's Hospital.
8vo, 10s. 6d. [1870]

MINOR SURGERY AND BANDAGING :

a Manual for the Use of House-Surgeons, Dressers, and Junior
Practitioners, by Christopher Heath, F.R.C.S., Surgeon to Uni-
versity College Hospital, and Holme Professor of Surgery in University
College. Fifth Edition, fcap 8vo, with 86 Engravings, 5s. 6d. [1875]

BY THE SAME AUTHOR,

INJURIES AND DISEASES OF THE JAWS :

Jacksonian Prize Essay. Second Edition, 8vo, with 164> Engrav-
ings, 12s. [1872]

BY THE SAME AUTHOR.

A COURSE OF OPERATIVE SURGERY :

with 20 Plates drawn from Nature by M. Leveille, and coloured
by hand under his direction. Large Svo. 40s. [1877]

HARE-LIP AND CLEFT PALATE,

by Francis Mason, F.R.C.S., Surgeon and Lecturer on Anatomy at
St. Thomas's Hospital. With 66 Engravings, Svo, 6s. [1877]

BY THE SAME AUTHOR,

THE SURGERY OF THE FACE :

with 100 Engravings. 8vo, 7s. 6d. [1878]

CATALOGUE OP RECENT WORKS

DISEASES AND INJURIES OF THE EAR,

by W. B. Dalby, F.R.O.S., M.B., Aural Surgeon and Lecturer on
Aural Surgery at St. George's Hospital. Crown 8vo, with 21 Engrav-
ings, 6s. 6d. [1873]

AURAL SURGERY ;

A Practical Treatise, by H. Macnaughton Jones, M.D., Professor
of the Queen's University in Ireland, Surgeon to the Cork Ophthalmic
and Aural Hospital. With 46 Engravings, crown 8vo, 5s. [1878]

BY THE SAME ATTTHOE,

ATLAS OP DISEASES OP THE ME MB RAN A TYMPANT.

In Coloured Plates, containing 62 Figures, with Text, crown 4to, 21s.

[1878]

THE EAR :

its Anatomy, Physiology, and Diseases. A Practical Treatise, by
Charles H. Burnett, A.M., M.D., Aural Surgeon to the Presby-
terian Hospital, and Surgeon in Charge of the Infirmary for Diseases
of the Ear, Philadelphia. With 87 Engravings, 8vo, 18s. Q877]

EAR AND THROAT DISEASES.

Essays by Llewellyn Thomas, M.D., Surgeon to the Central
London Throat and Ear Hospital. Post 8vo, 2s. 6d. [1878]

CLUBFOOT :

its Causes, Pathology, and Treatment : J acksonian Prize Essay by Wm-
Adams, F.R.C.S., Surgeon to the Great Northern Hospital. Second
Edition, 8vo, with 106 Engravings and 6 Lithographic Plates, 15s. [1873]

ORTHOPAEDIC SURGERY :

Lectures delivered at St. George's Hospital, by Bernard E. Brod-
httrst, F.R.C.S., Surgeon to the Royal Orthopaedic Hospital. Second
Edition,8vo, with Engravings, 12s. 6d. [1876]

OPERATIVE SURGERY OF THE FOOT AND ANKLE,

by Henry Hancock, F.R.C.S., Consulting Surgeon to Charing Cross
Hospital. 8vo, with Engravings, 15s. P873]

THE TREATMENT OF SURGICAL INFLAMMATIONS

by a New Method, which greatly shortens their Duration, by Furne aux
Jordan, F.R.C.S., Professor of Surgery in Queen's College, Birming-
ham. 8vo, with Plates, 7s. 6d. [1870]

BY THE SAME ATTTHOE,

SURGICAL INQUIRIES.

With numerous Lithographic Plates. 8vo, 5s. [1873]

ORTHOPRAXY :

the Mechanical Treatment of Deformities, Debilities, and Deficiencies of
the Human Frame, by H. Heather Bigg, Associate of the Institute of
Civil Engineers. Third Edition, with 319 Engravings, 8vo, 15s. [1877]

ORTHOPiEDIC SURGERY :

and Diseases of the Joints. Lectures by Lewis A. Sayre, M.D.,
Professor of Orthopaedic Surgery, Fractures and Dislocations, and
Clinical Surgery, in Bellevue Hospital Medical College, New York.
With 274 Wood Engravings, 8vo, 20s. [1876J

PUBLISHED BY J. AND A. 0HUR0H1LL

DISEASES OP THE RECTUM,

by Thomas B. Curling, F.R.S., Consulting Surgeon to tlio London
Hospital. Fourth Edition, Revised, 8vo, 7s. 6d. [1876]

BY THE SAME ATJTHOE,

DISEASES OF THE TESTIS, SPERMATIC CORD, AND SCROTUM.
Third Edition, with Engravings, 8vo, 16s. [1878]

FISTULA, HEMORRHOIDS, PAINFUL ULCER, STRICTURE,

Prolapsus, .and other Diseases of the Rectum : their Diagnosis and
Treatment. By "William Allingham, F.R.C.S., Surgeon to St.
Mark's Hospital for Fistula'. Third Edition, 8vo, 10s. PW9]

HYDROCELE :

its several Varieties and their Treatment, by Samuel Osborn,F.R.C.S.,
late Surgical Registrar to St. Thomas's Hospital. With Engravings,
fcap. 8vo, 3s. C1878]

STRICTURE OF THE URETHRA

and Urinary Fistulse ; their Pathology and Treatment : Jacksonian
Prize Essay by Sir Henry Thompson, F.R.C.S., Emeritus Professor
of Surgery to University College. Third Edition, 8vo, with Plates,
10s. [1869]

BY THE SAME AUTHOR,

PRACTICAL LITHOTOMY AND LITHOTRITY ;

or, An Inquiry into the best Modes of removing Stone from the
Bladder. Second Edition, 8vo, with numerous Engravings. 10s. [1871]

ALSO,

DISEASES OF THE URINARY ORGANS :

(Clinical Lectures). Fifth Edition, 8vo, with 2 Plates and 71
Engravings, 10s. 6d. [1879]

ALSO,

DISEASES OF THE PROSTATE :

their Pathology and Treatment. Fourth Edition, 8vo, with numerous
Plates, 10s. [1873]

ALSO,

THE PREVENTIVE TREATMENT OF CALCULOUS DISEASE

and the Use of Solvent Remedies. Second Edition, fcap. 8vo, 2s. 6d.

[1876]

STRICTURE OF THE URETHRA,

and other Diseases of the Urinary Organs, by Reginald Harrison,
F.R.C.S., Surgeon to the Liverpool Royal Infirmary. With 10 plates,
8vo, 7s. 6d. [1878]

LITHOTOMY AND EXTRACTION OF STONE

from the Bladder, Urethra, and Prostate of the Male, and from the
Bladder of the Female, by W. Pou lett Harris, M.D., Surgeon-Major
H.M. Bengal Medical Service. With Engravings, 8vo, 10s. Gd. [1876]

CATALOGUE OF RECENT WORKS

THE SURGERY OF THE RECTUM :

Lettsomian Lectures by Henry Smith, F.R.C.S., Professor of Surgery
in King's College, Surgeon to King's College Hospital. Fourth
Edition, fcap. 8vo, 5s. [187C]

KIDNEY DISEASES, URINARY DEPOSITS,

and Calculous Disorders by Lionel S. Beale, M.B., F.R.S., F.R.C.P.,
Physician to King's College Hospital. Third Edition, 8vo, with
70 Plates, 25s. [1868]

DISEASES OF THE BLADDER,

Prostate Gland and Urethra, including a practical view of Urinary
Diseases, Deposits and Calculi, by F. J. Gant, F.R.C.S., Senior Sur-
geon to the Royal Free Hospital. Fourth Edition, crown 8vo, with
Engravings, 10s. 6d. [1870]

THE DIAGNOSIS OF DISEASES OF THE KIDNEYS,

with Aids thereto, by W. R. Basham, M.D., F.R.C.P., late Senior
Physician to the Westminster Hospital. 8vo, with 10 Plates, 5s.

[1872]

RENAL AND URINARY DISEASES:

Clinical Reports, by William Caetee, M.B., M.R.C.P., Physician
to the Liverpool Southern Hospital. Crown 8vo, 7s. 6d. [1878]

THE REPRODUCTIVE ORGANS

in Childhood, Youth, Adult Age, and Advanced Life (Functions and
Disorders of), considered in their Physiological, Social, and Moral
Relations, by William Acton, M.R.C.S. Sixth Edition, 8vo, 12s.

[1875]

URINARY AND REPRODUCTIVE ORGANS :

their Functional Diseases, by D. Campbell Black, M.D., L.R.C.S.
Edin. Second Edition. 8vo, 10s. 6d. [1875]

LECTURES ON SYPHILIS,

and on some forms of Local Disease, affecting principally the Organs
of Generation, by Heney Lee, F.R.C.S., Surgeon to St. George's Hos-
pital. With Engravings, 8vo, 10s. [18?5]

.SYPHILITIC NERVOUS AFFECTIONS :

Their Clinical Aspects, by Thomas Buzzaed, M.D., F.R.C.P. Lond.,
Physician to the National Hospital for Paralysis and Epilepsy. Post
8vo, 5s. £1874]

PATHOLOGY OF THE URINE,

including a Complete Guide to its Analysis, by J. L. W. Tiiudichum,
M.D., F.R.C.P. Second Edition, rewritten and enlarged, with En-
gravings, 8vo, 15s. [1877]

HJBLISIIED BY J. AND A. CHURCHILL

GENITO URINARY ORGANS, INCLUDING SYPHILIS :

A Practical Treatise on their Surgical Diseases, designed as a Manual
for Students and Practitioners, by W. H. Van Buren, M.D., Pro-
fessor of the Principles of Surgery in Bellevue Hospital Medical Col-
lege, New York, and E. L. Keyes, M.D., Professor of Dermatology in
Bellevue Hospital Medical College, New York. Royal 8vo, with 140
Engravings, 21s. [1874]

HISTOLOGY AND HISTO-CHEMISTRY OF MAN :

A Treatise on the Elements of Composition and Structure of the
Human Body, by Heinrich Fret, Professor of Medicine in Zurich.
Translated from the Fourth German Edition by Arthur E. J.
Barker, Assistant-Surgeon to University College Hospital. And
Revised by the Author. 8vo, with 608 Engravings, 21s. t1874J

HUMAN PHYSIOLOGY:

A Treatise designed for the Use of Students and Practitioners of
Medicine, by John C. Dalton, M.D., Professor of Physiology and
Hygiene in the College of Physicians and Surgeons, New York. Sixth
Edition, royal 8vo, with 316 Engravings, 20s. [1875]

HANDBOOK FOR THE PHYSIOLOGICAL LABORATORY,

by E. Klein, M.D., F.R.S., Assistant Professor in the PathologicalLabo-
ratory of the Brown Institution, London; J. Burdon-Sanderson,
M.D., F.R.S., Professor of Practical Physiology in University College,
London; Michael Foster, M.D., F.R.S., Praelector of Physiology
in Trinity College, Cambridge; and T. Lauder Brunton, M.D.,
F.R.S., Lecturer on Materia Medica at St. Bartholomew's Hospital ;
edited by J. Burdon-Sanderson. 8vo, with 123 Plates, 24s. E1873]

PRACTICAL HISTOLOGY :

By William Rutherford, M.D., Professor of the Institutes of
Medicine in the University of Edinburgh. Second Edition, with
63 Engravings. Crown 8vo (with additional leaves for notes), 6s.

[1876]

THE MARRIAGE OF NEAR KIN,

Considered with respect to the Laws of Nations, Results of Experience,
and the Teachings of Biology, by Alfred H. Huth. 8vo, 14s. [18753

MANUAL OF ANTHROPOMETRY:

A Guide to the Measurement of the Human Body, containing an
Anthropometrical Chart and Register, a Systematic Table of Mea-
urements, &c. By Charles Roberts, F.R.C.S., late Assistant
Surgeon to* the Yictoria Hospital for Children. With numerous
Illustrations and Tables. 8vo, 6s. 6d. [1878]

STUDENTS' GUIDE TO HUMAN OSTEOLOGY,

By William Warwick Wagstaffe, F.R.C.S., Assistant-Surgeon
and Lecturer on Anatomy, St. Thomas's Hospital. With 23 Plates
and 66 Engravings. Fcap. 8vo, 10s. Gd. [1875]

CATALOGUE OP EE CENT WORKS

PRINCIPLES OF HUMAN PHYSIOLOGY,

by W. B. Caepentee, O.B., M.D., F.R.S. Eighth Edition by Heney
Powee, M.B., F.R.O.S., Examiner in Natural Science, University of
Oxford, and in Natural Science and Medicine, University of Cambridge.
8vo, with 3 Steel Plates and 371 Engravings, 31s. 6d. " [i870]

LANDMARKS, MEDICAL AND SURGICAL,

By Luthee Holden, F.R.C.S., Member of the Court of Examiners of
the Royal College of Surgeons. Second Edition, 8vo, 3s. 6d. C1877]

BY THK SAME ATJTHOE.

HUMAN OSTEOLOGY:

Comprising a Description of the Bones, with Delineations of the
Attachments of the Muscles, the General and Microscopical Structure
of Bone, and its Development. Fifth Edition, with 61 Lithographic
Plates and 89 Engravings. 8vo, 16s. [1878]

PATHOLOGICAL ANATOMY :

Lectures by Samuel Wiles, M.D., F.R.S., Physician to, and Lec-
turer on Medicine at, Guy's Hospital; and Waltee Moxon, M.D.,
F.R.C.P., Physician to, and Lecturer on Materia Medica at, Guy's
Hospital. Second Edition, 8vo, with Plates, 18s. [1875]

PATHOLOGICAL ANATOMY :

A Manual by C. Handfield Jones, M.B., F.R.S., Physician to St.
Mary's Hospital, and Edwaed H. Sieveking, M.D., F.R.C.P.,
Physician to St. Mary's Hospital. Edited by J. F. Payne, M.D.,
F.R.C.P., Assistant Physician and Lecturer on General Pathology
at St. Thomas's Hospital. Second Edition, crown 8vo, with 195
Engravings, 16s. [1875]

POST-MORTEM EXAMINATIONS :

a Description and Explanation of the Method of Performing them,
with especial Reference to Medico-Legal Practice. By Professor
Rudolph Yiechow, of Berlin. Fcap 8vo, 2s. 6d. [i876]

STUDENT'S GUIDE TO SURGICAL ANATOMY :

a Text-book for the Pass Examination, by E. Bellamy, F.R.C.S.,
Surgeon and Lecturer on Anatomy at Charing Cross Hospital. Fcap
8vo: with 50 Engravings, 6s. 6d. t1873]

ANATOMY OF THE JOINTS OF MAN,

by Heney Moeeis, F.R.C.S., Senior Assistant- Surgeon to, and Lec-
turer on Anatomy and Demonstrator of Operative Surgery at, the
Middlesex Hospital. "With 44 Lithographic Plates (several being
coloured) and 13 "Wood Engravings. 8vo, 16s. [1879]

MEDICAL ANATOMY,

by Feancis Sibson, M.D., F.R.C.P., F.R.S. Imp. folio, with 21
coloured Plates, cloth, 42s., half -morocco, 503. C1809]

PRACTICAL ANATOMY :

a Manual of Dissections by Cheistophee Heath, F.R.CS., Surgeon
to University College Hospital, and Holme Professor of Surgery in
University College. Fourth Edition, crown 8vo, with 16 Coloured
Plates and 2G4 Engravings, 14s. [1S77-'

PUBLISHED BY J. AND A. CHUECHILL

AN ATLAS OF HUMAN ANATOMY :

illustrating most of the ordinary Dissections, and many not usually
practised by the Student. To be completed in 12 or 13 Bi-monthly
Parts, each containing 4 Coloured Plates, with Explanatory Text. By
Rickman J. Godlee, M.S., F.ll.C.S., Assistant Surgeon to University
College Hospital, and Senior Demonstrator of Anatomy in University
College. Parts I to VII. Imp. 4to, 7s. 6d. each Part. [1877-9]

THE ANATOMIST'S VADE-MECUM :

a System of Human Anatomy by Erasmus "Wilson, F.R.C.S., F.R.S.
Ninth Edition, by G. Buchanan, M.A., M.D., Professor of Clinical
Surgery in the University of Glasgow, and Henry E. Clark, F.F.P.S.,
Lecturer on Anatomy at the Glasgow Royal Infirmary School of
Medicine. Crown 8vo, with 371 Engravings, 14s. [1873]

ATLAS OF TOPOGRAPHICAL ANATOMY,

after Plane Sections of Frozen Bodies. By "Wilhelm Braune,
Professor of Anatomy in the University of Leipzig. Translated by
Edward Bellamy, F.R.C.S., Surgeon to, and Lecturer on Anatomy,
&c, at, Charing Cross Hospital. With 34 Photo-lithographic Plates
and 46 Woodcuts. Large Imp. 8vo, 40s. [1877]

THE ANATOMICAL REMEMBRANCER;

or, Complete Pocket Anatomist. Eighth Edition, 32mo, 3s. 6d. [i^l

THE STUDENT'S GUIDE TO THE PRACTICE OF MEDICINE,
by Matthew Charteris, M.D., Professor of Medicine in Anderson's
College, and Lecturer on Clinical Medicine in the Royal Infirmary,
Glasgow. Second Edition, with Engravings on Copper and Wood,
fcap. 8vo, 6s. 6d. [1878

THE MICROSCOPE IN MEDICINE,

by Lionel S. Beale, M.B., F.R.S., Physician to King's College
Hospital. Fourth Edition, with 86 Plates, 8vo, 21s. [1877]

HOOPER'S PHYSICIAN'S VADE-MECUM ;

or, Manual of the Principles and Practice of Physic, Ninth Edition
by W. A. Guy, M.B., F.R.S., and John Harley, M.D., F.R.C.P.
Fcap 8vo, with Engravings, 12s. 6d. [1874]

A NEW SYSTEM OF MEDICINE ;

entitled Recognisant Medicine, or the State of the Sick, by
Bholanoth Bose, M.D., Indian Medical Service. 8vo, 10s. 6d. [1877

BY THE SAME ATTTHOK.

PRINCIPLES OF RATIONAL THERAPEUTICS.

Commenced as an Inquiry into the Relative Value of Quinine and
Arsenic in Ague. 8vo, 4s. [1877]

CLINICAL MEDICINE :

Lectures and Essays by Balthazar Foster, M.D., F.R.C.P. Lond.,
Professor of Medicine in Queen's College, Birmingham. 8vo, 10s. 6d.

CLINICAL REMINISCENCES: [1874J
By Peyton Blakiston, M.D., F.R.S. Post 8vo, 3s. 6d. [1S73]

CATALOGUE OP RECENT WORKS

THE STUDENT'S GUIDE TO MEDICAL DIAGNOSIS,

by Samuel Fenwick, M.D., F.R.C.P., Physician to the London
Hospital. Fourth Edition, fcap. 8vo, with 106 Engravings, Gs. Gd. [187C]

A MANUAL OF MEDICAL DIAGNOSIS,

by A. W. Barclay, M.D., F.R.C.P., Physician to, and Lecturer on
Medicine at, St. George's Hospital. Third Edition, fcap 8vo, 10s. Gd.

CLINICAL STUDIES:

Illustrated by Cases observed in Hospital and Private Practice, by Sir
J. Rose Coemack, M.D., F.R.S.E., Physician to the Hertford British
Hospital of Paris. 2 vols., post 8vo, 20s. [1876]

ROTLE'S MANUAL OF MATERIA MEDICA AND THERAPEUTICS.
Sixth Edition by John Haeley, M.D., F.R.C.P., Assistant Physician
to, and Joint Lecturer on Physiology at, St. Thomas's Hospital. Crown
8vo, with 139 Engravings, 15s. [1&7C]

PRACTICAL THERAPEUTICS :

A Manual by E. J. Waeing, M.D., F.R.C.P. Lond. Third Edition,
fcap 8vo, 12s. 6d. Q87i]

THE ELEMENTS OF THERAPEUTICS.

A Clinical Guide to the Action of Drugs, by C. Binz, M.D., Professor
of Pharmacology in the University of Bonn. Translated and Edited
with Additions, in Conformity with the British and American Phar-
macopoeias, by Edward I. Spaeks, M.A., M.B. Oxon., formerly
Radcliffe Travelling Fellow. Crown 8vo, 8s. 6d. [1877]

THE NATIONAL DISPENSATORY;

containing the Natural History, Chemistry, Pharmacy, Actions and
Uses of Medicines, including those recognised in the Pharmacopoeias of
the United States and Great Britain. By Alfeed Stille, M.D., and
John M. Maisch, Ph. D. 1628 pp., with 201 Engravings, royal 8vo,
34s. P879J

THE STUDENT'S GUIDE TO MATERIA MEDICA,

by John C. Thoeowgood, M.D., F.R.C.P. Lond., Physician to the
City of London Hospital for Diseases of the Chest. Fcap 8vo, with
Engravings, 6s. 6d. [1874]

MATERIA MEDICA AND THERAPEUTICS :

(Vegetable Kingdom), by Chaeles D. F. Phillips, M.D., F.R.C.S.E.

DENTAL MATERIA MEDICA AND THERAPEUTICS,

Elements of, by James Stocken, L.D.S.R.C.S., Lecturer on Dental
Materia Medica and Therapeutics to the National Dental Hospital.

THE DISEASES OF CHILDREN :

A Practical Manual, with a Formulary, by Edward Ellis, M.D.,
late Senior Physician to the Victoria Hospital for Children. Third

8vo, 15s.

[1874]

Second Edition, Fcap 8vo, 6s. 6d.

[1878]

Edition, crown 8vo, 7s. 6d.

[1878]

PUBLISHED BY J. AND A. CHURCHILL

THE WASTING DISEASES OF CHILDREN,

by Eustace Smith, M.D., F.R.C.P. Lond., Physician to the King of
the Belgians, Physician to the East London Hospital for Children.
Third Edition, post 8vo, 8s. 6d. [1878]

BY ME SAME AUTIIOB,

CLINICAL STUDIES OF DISEASE IN CHILDREN.

Post Svo, 7s. 6d. [187C]

INFANT FEEDING AND ITS INFLUENCE ON LIFE ;

or, the Causes and Prevention of Infant Mortality, by Charles H. F.
Routh, M.D., Senior Physician to the Samaritan Hospital for Women
and Children. Third Edition, fcap Svo, 7s. 6d. [1876]

COMPENDIUM OF CHILDREN'S DISEASES :

A Handbook for Practitioners and Students, by Johann Steiner,
M.D., Professor in the University of Prague. Translated from the
Second German Edition by Lawson Tait, F.R.C.S., Surgeon to the
Birmingham Hospital for Women. 8vo, 12s. 6d. [1874-]

THE DISEASES OF CHILDREN:

Essays by William Henry Day, M.D., Physician to the Samaritan
Hospital for Diseases of Women and Children. SecondEdition,fcap8vo.

[In the Press. 1

PUERPERAL DISEASES: . J

Clinical Lectures by Fordyce Barker, M.D., Obstetric Physician
to Bellevue Hospital, New York. 8vo, 15s. [1874]

THE STUDENT'S GUIDE TO THE PRACTICE OF MIDWIFERY,
by D. Lloyd Roberts, M.D., F.R.C.P, Physician to St. Mary's Hos-
pital, Manchester. Second Edition, fcap. Svo, with 111 Engravings, 7s.

OBSTETRIC MEDICINE AND SURGERY, [1879J
Their Principles and Practice, by F. H. Ramsbotham, M.D., F.R.C.P.
Fifth Edition, Svo, with 120 Plates, 22s. [1867]

OBSTETRIC SURGERY:

A Complete Handbook, giving Short Rules of Practice in every Emer-
gency, from the Simplest to the most Formidable Operations connected
with the Science of Obstetricy, by Charles Clay, Ext.L.R.C.P. Lond.,
L.R.C.S.E., late Senior Surgeon and Lecturer on Midwifery, St.
Mary's Hospital, Manchester. Fcap 8vo, with 91 Engravings, 6s. 6d.

SCHROEDER'S MANUAL OF MIDWIFERY, [187i]
including the Pathology of Pregnancy and the Puerperal State.
Translated by Charles H. Carter, B.A., M.D. Svo, with En crav-
ings, 12s. 6d. [I873]

A HANDBOOK OF UTERINE THERAPEUTICS,

and of Diseases of Women, by E. J. Tilt, M.D,, M.R.C.P. Fourth
'Edition, post Svo, 10s. [1878]

BY THE SAME AUTHOB,

THE CHANGE OF LIFE

in Health and Disease : a Practical Treatise on the Nervous and other
Affections incidental to Women at the Decline of Life. Third Edition
8vo, 10s. 6d. [1870]'

CATALOGUE OF RECENT WORKS

OBSTETRIC OPERATIONS,

including the Treatment of Haemorrhage, and forming a Guide to the
Management of Difficult Labour; Lectures by Robert Barnes, M.D.,
F.R.C.P., Obstetric Physician and Lecturer on Obstetrics and the Dis-
eases of Women and Children at St. George's Hospital. Third Edition,
• 8vo, with 124 Engravings, 18s. [I875]

BY THE SAME AUTHOR,

MEDICAL AND SURGICAL DISEASES OF WOMEN:

a Clinical History. Second Edition, Svo, with 181 Engravings, 28s.

[1878]

THE DISEASES OF WOMEN.

By Charles West, M.D., F.R.C.P. Revised and in part Re-written
by the Author, with numerous Additions by J. Matthews Duncan,
M.D., Obstetric Physician to St. Bartholomew's Hospital. Fourth
Edition, Svo, 16s. [1879]

OBSTETRIC APHORISMS:

for the Use of Students commencing Midwifery Practice by J. G.
Swayne, M.D., Consulting Physician-Accoucheur to the Bristol
General Hospital, and Lecturer on Obstetric Medicine at the Bristol
Medical School. Sixth Edition, fcap Svo, with Engravings, 3s. 6d. [1876]

DISEASES OF THE OVARIES :

their Diagnosis and Treatment, by T. Spencer Wells, F.R.C.S.,
Surgeon to the Queen's Household and to the Samaritan Hospital.
8vo, with about 150 Engravings, 21s. t1872]

PRACTICAL GYNAECOLOGY:

A Handbook of the Diseases of Women, by Heywood Smith, M.D.

Oxon., Physician to the Hospital for Women and to the British Lying-
in Hospital. With Engravings, crown 8vo, 5s. 6d. [1877]

THE PRINCIPLES AND PRACTICE OF GYNAECOLOGY.

By Thomas Addis Emmet, M.D., Surgeon to the Woman's Hospital
of the State of New York. With 130 Engravings, 8vo, 24s. [1879]

RUPTURE OF THE FEMALE PERINEUM,

Its treatment, immediate and remote, by George G. Bantock, M.D.,
Surgeon (for In-patients) to the Samaritan Free Hospital for Women
and Children. With 2 plates, 8vo, 3s. 6d. [1878]

PAPERS ON THE FEMALE PERINEUM, &c,

by James Matthews Duncan, M.D., Obstetric Physician to St. Bar-
tholomew's Hospital. Svo, 6s. [1878J

INFLUENCE OF POSTURE ON WOMEN

In Gynecic and Obstetric Practice, by J. H. Aveling, M.D., Physi-
cian to the Chelsea Hospital for Women, Yice-President of the
Obstetrical Society of London. Svo, 6s. DS78}

THE NURSE'S COMPANION:

A Manual of General and Monthly Nursing, by Charles J. Culling-

worth, Surgeon to St. Mary's Hospital, Manchester. Fcap. Svo,

2s. 6d. it1370

PUBLISHED BY J. AND A. CHURCHILL

A MANUAL FOR HOSPITAL NURSES

and others engaged in Attending on the Sick by Edward J. Dom-
ville, L.R.C.P., M.R.C.S., Surgeon to the Exeter Lying-in Charity.
Third Edition, crown 8vo, 2s. 6d. [1878]

LECTURES ON NURSING,

by William Robert Smith, M.B., Honorary Medical Officer,
Hospital for Sick Children, Sheffield. Second Edition, with 26 En-
gravings. Post 8vo, 6s. [1878]

HANDBOOK FOR NURSES FOR THE SICK,

by Zepherina P. Yeitch. Second Edition, crown 8vo, 3s. 6d. P876]

A COMPENDIUM OF DOMESTIC MEDICINE

and Companion to the Medicine Chest ; intended as a Source of Easy
Reference for Clergymen, and for Families residing at a Distance
from Professional Assistance, by John Savory, M.SA. Ninth
Edition, 12mo, 5s. [1878]

HOSPITAL MORTALITY

being a Statistical Investigation of the Returns of the Hospitals of
Great Britain and Leland for fifteen years, by Lawson Tait, F.R.C.S.,
F.S.S. 8vo, 8s. 6d. [1877]

THE COTTAGE HOSPITAL:

Its Origin, Progress, Management, and Work, by Henry C. Burdett,
the Seaman's Hospital, Greenwich. With Engravings, crown 8vo,
7s. 6d. [1877]

WINTER COUGH :

(Catarrh, Bronchitis, Emphysema, Asthma), Lectures by Horace
Dobell,M.D., Consulting Physician to the Royal Hospital for Diseases
of the Chest. Third Edition, with Coloured Plates, 8vo, 10s. 6d. [1875]

BY THE SAME AUTHOR,

LOSS OF WEIGHT, BLOOD-SPITTING, AND LUNG DISEASE.
With Chromo-lithograph, 8vo, 10s. 6d. [1878]

CONSUMPTION :

Its Nature, Symptoms, Causes, Prevention, Curability, and Treatment.
By Peter Gowan, M.D., B. Sc., late Physician and Surgeon in
Ordinary to the King of Siam. Crown 8vo. 5s. [1878

NOTES ON ASTHMA ;

its Forms and Treatment, by John C. Thorowgood, M.D. Lond.,
F.R.C.P., Physician to the Hospital for Diseases of the Chest, Victoria
Park. Third Edition, crown 8vo, 4s. 6d. [1878]

ASTHMA

Its Pathology and Treatment, by J. B. Bericart, M.D., Assistant
Physician to the City of London Hospital for Diseases of the Chest.
8vo, 7s. 6d. [1878]
DISEASES OF THE HEART :

Their Pathology, Diagnosis, Prognosis, and Treatment (a Manual),
by Robert H. Semple, M.D., F.R.C.P., Physician to the Hospital for
Diseases of the Throat. 8vo, 8s. 6d. [1876]

CATALOGUE 01' RECENT WOJtKS

PROGNOSIS IN CASES OF VALVULAR DISEASE OF THE
Heart, by Thomas B. Peacock, M.D., F.R.C.P., Honorary Consult-
ing Physician to St. Thomas's Hospital. 8vo, 3s. Gd. [18753

CHRONIC DISEASE OF THE HEART :

Its Bearings upon Pregnancy, Parturition and Childbed. By Angus
Macdonald, M.D., F.R.S.E., Physician to, and Clinical Lecturer on
the Diseases of Women at, the Edinburgh Royal Infirmary. With
Engravings, Svo, 8s. Gd. [1876]

PHTHISIS :

In a series of Clinical Studies, by Austin Flint, M.D., Professor of
the Principles and Practice of Medicine and of Clinical Medicine in
the Bellevue Hospital Medical College. Svo, 16s. [1875]

BY TO SAME AUTIlOIt,

A MANUAL OF PERCUSSION AND AUSCULTATION,

of the Physical Diagnosis of Diseases of the Lungs and Heart, and of
Thoracic Aneurism. Post Svo, 6s. 6d. [187GJ

DIPHTHERIA:

its Nature and Treatment, Varieties, and Local Expressions, by
Morell Mackenzie, M.D., Physician to the Hospital for Diseases of
the Throat. Crown 8vo, 5s. [1878]

DISEASES OF THE HEART AND AORTA,

By Thomas Hayden, F.K.Q.C.P. Irel., Physician to the Mater
Misericordias Hospital, Dublin. With 80 Engravings. Svo, 25s. [1875]

DISEASES OF THE HEART

and of the Lungs in Connexion therewith — Notes and Observations
by Thomas Shapter, M.D., F.R.C.P. Lond., Senior Physician to the
Devon and Exeter Hospital. 8vo, 7s. 6d. [1874]

DISEASES OF THE HEART AND AORTA :

Clinical Lectures by George W. Baleour, M.D., F.R.C.P., Physician
to, and Lecturer on Clinical Medicine in, the Royal Infirmaiy, Edin-
burgh. Svo, with Engravings, 12s. Gd. [1876]

PHYSICAL DIAGNOSIS OF DISEASES OF THE HEART.

Lectures by Arthur E. Sansom, M.D., F.R.C.P., Assistant Physician
to the London Hospital. Second Edition, with Engravings, fcap. Svo,
4s. Gd. [1876]

TRACHEOTOMY,

especially in Relation to Diseases of the Larynx and Trachea, by
Pugin Thornton, M.R.C.S., late Surgeon to the Hospital for Diseases
of the Throat. With Photographic Plates and Woodcuts, Svo, 5s. Gd.

[1876]

SORE THROAT:

Its Nature, Varieties, and Treatment, including the Connexion
between Affections of the Throat and other Diseases. By Prosser
James, M.D., Lecturer on Materia Medica and Therapeutics at the
London Hospital, Physician to the Hospital for Diseases of the Throat.
Third Edition, with Coloured Plates, 5s. Gd. [1878]

rUBLISHED BY J. AND A. CHUEOHILL

SYPHILIS OF THE LARYNX

(Lesions of the Secondary and Intermediate Stages). Lectures
delivered at the Hospital for Diseases of the Throat and Chest,
London, by W. Macneill Whistler, M.D., M.R.C.P., Physician to
the Hospital; Honorary Physician to the National Training School
for Music. With Engravings, crown 8vo, 4s. [1879]

WINTER AND SPRING

on the Shores of the Mediterranean. By Henry Bennet, M.D.

Fifth Edition, post Svo, with numerous Plates, Maps, and Engravings,

12s. 6d. C1874]

BY THE SAME AUTHOR,

TREATMENT OF PULMONARY CONSUMPTION

by Hygiene, Climate, and Medicine. Third Edition, Svo, 7s. 6d. [1878]

PRINCIPAL HEALTH RESORTS

of Europe and Africa, and their Use in the Treatment of Chronic

Diseases. A Handbook by Thomas More Madden, M.D., M.R.I.A.,

Vice-President of the Dublin Obstetrical Society. Svo, 10s. [1876]

THE BATH THERMAL WATERS :

Historical, Social, and Medical, by John Kent Spender, M.D.,

Surgeon to the Mineral Water Hospital, Bath. With an Appendix

on the Climate of Bath by the Rev. L. Blomefield, M.A., F.L.S.,

F.G.S. Svo, 7s. 6d. [1877]

ENDEMIC DISEASES OF TROPICAL CLIMATES,

with their Treatment, by John Sullivan, M. D., M.R.C.P. Post Svo,
6s. [1877]

DISEASES OF TROPICAL CLIMATES

and their Treatment : with Hints for the Preservation of Health in the

Tropics, by James A. Horton, M.D., Surgeon-Major, Army Medical

Department. Second Edition, post; Svo, 12s. 6d. [1879]

HEALTH IN INDIA FOR BRITISH WOMEN

and on the Prevention of Disease in Tropical Climates by Edward J.
Tilt, M.D., Consulting Physician-Accoucheur to the Farringdon
General Dispensary. Fourth Edition, crown Svo, -5s. [1875]

BURDWAN FEYER,

or the Epidemic Fever of Lower Bengal (Causes, Symptoms, and
Treatment), by Gopatjl Chunder Roy, M.D., Surgeon Bengal
Establishment. New Edition, 8vo, 5s. [1876]

BAZAAR MEDICINES OF INDIA

and Common Medical Plants : Remarks on their Uses, with Full Index
of Diseases, indicating their Treatment by these and other Agents pro-
curable throughout India, &c, by Edward J. Waring, M.D., F.R.CP.
Lond., Retired Surgeon H.M. Indian Army. Third Edition. Fcap

8VO, 5s. [1875]

SOME AFFECTIONS OF THE LIYER

and Intestinal Canal ; with Remarks on Ague and its Sequela), Scurvy,
Purpura, &c, by Stephen H. Ward, M.D. Lond., F.R.C.P., Physician
to the Seamen's Hospital, Greenwich. Svo, 7s. [1872]

CATALOGUE OF RECENT WORKS

DISEASES OF THE LIVER:

Lettsomian Lectures for 1872 by S. O. Habershon, M.D., F.R.C.P.,
Senior Physician to Guy's Hospital. Post 8vo, 3s. 6d. U872]

• BY THE SAME AUTHOR,

DISEASES OF THE STOMACH : DYSPEPSIA.

Third Edition, crown 8vo, 5s. [1879]

BY THE SAME ATJTHOK,

PATHOLOGY OP THE PNEUMOGASTRIC NERVE,

being the Lumleian Lectures for 1876. Post 8vo, 3s. 6d. [1W7]

BY THE SAME AUTHOE,

DISEASES OF THE ABDOMEN,

comprising those of the Stomach and other parts of the Alimentary
Canal, (Esophagus, Caecum, Intestines, and Peritoneum. Third
Edition, with 5 Plates, 8vo, 21s. [1678]

FUNCTIONAL NERVOUS DISORDERS :

Studies by C. Handfield Jones, M.B., F.R.C.P., F.R.S., Physician
to St. Mary's Hospital. Second Edition, 8vo, 18s. P870]

LECTURES ON DISEASES OF THE NERVOUS SYSTEM,

by Samuel Wiles, M.D., F.R.S., Physician to, and Lecturer on
Medicine at, Guy's Hospital. Svo, 15s. [1878]

NERVOUS DISEASES:

their Description and Treatment, by Allen McLane Hamilton, M.D.,
Physician at the Epileptic and Paralytic Hospital, Blackwell's Island,
New York City. Roy. Svo, with 53 Illustrations, 14s. P878J

NUTRITION IN HEALTH AND DISEASE :

A Contribution to Hygiene and to Clinical Medicine. By Henry
Bennet, M.D. Third (Library) Edition. 8vo, 7s. Cheap Edition,
Fcap. 8vo, 2s. 6d.

FOOD AND DIETETICS,

Physiologically and Therapeutically Considered. By Frederick W.
Pavst, M.D., F.R.S., Physician to Guy's Hospital. Second Edition,

Svo, 15s. £1875]

BY THE SAME AUTHOR.

CERTAIN POINTS CONNECTED WITH DIABETES

(Croonian Lectures). 8vo, 4s. 6d. t1878!
HEADACHES :

their Causes, Nature, and Treatment. By William H. Day, M.D.,
Physician to the Samaritan Free Hospital for Women and Children.
Second Edition, crown Svo, with Engravings. 6s. 6d. P878}

IMPERFECT DIGESTION :

its Causes and Treatment by Arthur Leared, M.D., F.R.C.P.,
Senior Physician to the Great Northern Hospital. Sixth Edition,
fcap 8vo, 4s. 6d. [1875]

MEGRIM, SICK-HEADACHE,

and some Allied Disorders : a Contribution to the Pathology of Nerve-
Storms, by Edward Liveing, M.D. Cantab., F.R.C.P., Hon. Fellow
of King's College, London. 8vo, with Coloured Plate, 15s. [1878J

PUBLISHED BY J. AND A. OHUEOHILL

NEURALGIA AND KINDRED DISEASES

of the Nervous System : their Nature, Causes, and Treatment, with, a
series of Cases, by John Chapman, M.D., M.R.C.P. 8vo, 14s. [1873}

THE SYMPATHETIC SYSTEM OF NERVES :

their Physiology and Pathology, by A. Eolenburg, Professor of
Medicine, University of Greif swald, and Dr. P. Gtjttmann, Priva Docen
in Medicine, University of Berlin. Translated by A. Napier, M.D.*
F.F.P.S. 8vo, 5s. [1879]

RHEUMATIC GOUT,

or Chronic Rheumatic Arthritis of all the Joints ; a Treatise by
Robert Adams, M.D., M.R.I.A., late Surgeon to H.M. the Queen in
Ireland, and Regius Professor of Surgery in the University of Dublin.
Second Edition, 8vo, with Atlas of Plates, 21s. [1872]

GOUT, RHEUMATISM,

and the Allied Affections; with a chapter on Longevity and the
Causes Antagonistic to it, by Peter Hood, M.D. Second Edition,
crown 8vo, 10s. 6d. [1879]

RHEUMATISM :

Notes by Julius Pollock, M.D., F.R.C.P., Senior Physician to, and
Lecturer on Medicine at, Charing Cross Hospital. Second Edition,
with Engravings, fcap. 8vo, 3s. 6d. [1879]

CERTAIN FORMS OF CANCER,

with a New and successful Mode of Treating it, to which is prefixed a
Practical and Systematic Description of all the varieties of this Disease,
by Alex. Marsden, M.D., F.R.C.S.E., Consulting Surgeon to the
Royal Free Hospital, and Senior Surgeon to the Cancer Hospital.
Second Edition, with Coloured Plates, 8vo, 8s. 6d. [1873]

DISEASES OF THE SKIN :

a System of Cutaneous Medicine by Erasmus Wilson, F.R.C.S.,
F.RS. Sixth Edition, 8vo, 18s., with Coloured Plates, 36s. [1867]

BX THE SAME ATJTHOE,

LECTURES ON EKZEMA

and Ekzematous Affections: with an Introduction on the General
Pathology of the Skin, and an Appendix of Essays and Cases. 8vo>
10s. 6d. also, [1870]

LECTURES ON DERMATOLOGY:

delivered at the Royal College of Surgeons, 1870, 6s. ; 1871-3, 10s. 6d.,
1874-5, 10s. 6d. ; 1876-8, 10s. 6d.

ECZEMA :

by McCall Anderson, M.D., Professor of Clinical Medicine in the Uni-
versity of Glasgow. Third Edition, 8vo, with Engravings, 7s. 6d. [1874]

BY THE SAME ATJTITOU,

PARASITIC AFFECTIONS OF THE SKIN

Second Edition, 8vo, with Engravings, 7s. 6d. [lSCSj

CATALOGUE OP REGENT WORKS

ATLAS OP SKIN DISEASES :

a series of Illustrations, with Descriptive Text and Notes upon Treat-
ment. By Tilbury Fox, M.D., F.R.C.P., Physician to the Department
for Skin Diseases in University College Hospital. With 72 Coloured
Plates, royal 4to, half morocco, £G 6s. [1877]

PSORIASIS OR LEPRA,

by George Gaskoin, M.R.C.S., Surgeon to the British Hospital for
Diseases of the Skin. 8vo, 5s. 0875]

MYCETOMA;

or, the Fungus Disease of India, by H. Vandyke Carter, M.D., Sur-
geon-Major H.M. Indian Army. 4to, with 11 Coloured Plates, 42s.

[1874]

DISEASES OF THE SKIN,

in Twenty-four Letters on the Principles and Practice of Cutaneous
Medicine, by Henry Evans Catjty, M.R.C.S., Surgeon to the Liver-
pool Dispensary for Diseases of the Skin, 8vo, 12s. 6d. 087*]

THE HAIR IN HEALTH AND DISEASE,

by E. Wyndham Cottle, F.R.C.S., Senior Assistant Surgeon to the
Hospital for Diseases of the Skin, Blackfriars. Fcap. 8vo, 2s. 6d. 0877]

THE LAWS AFFECTING MEDICAL MEN:

a Manual by Robert G. Glenn, LL.B., Barrister-at-Law ; with a
Chapter on Medical Etiquette by Dr. A. Carpenter. 8vo, 14s. [is?]]

MEDICAL JURISPRUDENCE,

Its Principles and Practice, by Alfred S. Taylor, M.D., F.R.C.P.,
F.R.S. Second Edition, 2 vols., Svo, with 189 Engravings, £1 lis. Cd.

0873]

BY THE SAME AUTHOR,

A MANUAL OF MEDICAL JURISPRUDENCE.

Tenth Edition. Crown Svo, with Engravings, 14s. 0&79]

ALSO,

POISONS,

in Relation to Medical Jurisprudence and Medicine. Third Edition,
crown Svo, with 104 Engravings, 16s. [1875]

MEDICAL JURISPRUDENCE :

Lectures by Francis Ogston, M.D., Professor of Medical Juris-
prudence and Medical Logic in the University of Aberdeen. Edited
by Francis Ogston, Jnn., M.D., Assistant to the Professor of
Medical Jurisprudence and Lecturer on Practical Toxicology in the
University of Aberdeen. Svo, with 12 Copper Plates, 18s. t187S]

A TOXICOLOGICAL CHART,

exhibiting at one Yiew the Symptoms, Treatment, and mode of
Detecting the various Poisons— Mineral, Yegetable, and Animal :
with Concise Directions for the Treatment of Suspended Animation,
by William Stowe, M.R.C.S.E. Thirteenth Edition, 2s.; on
roller, 5s. [1S7"]

)

PUBLISHED BY J. AND A. CHU11CHILL

A HANDY-BOOK OF FORENSIC MEDICINE AND TOXICOLOGY,
by W. Bathurst Woodman, M.D., F.R.C.P., Assistant Physician
and Co-Lecturer on Physiology and Histology at the London Hospital ;
and C. Meymott Tidy, M.D., F.C.S., Professor of Chemistry and of
Medical Jurisprudence and Public Health at the London Hospital.
With 8 Lithographic Plates and 11G Engravings, 8vo, 31s. 6d. C1877J

THE MEDICAL ADVISER IN LIFE ASSURANCE,

by Edward Henry Sieveking, M.D., F.R.C.P., Physician to St.
Mary's and the Lock Hospitals; Physician-Extraordinary to the
Queen; Physician-in- Ordinary to the Prince of Wales, &c. Crown
8vo, 6s. [1874]

IDIOCY AND IMBECILITY,

by William W. Ireland, M.D., Medical Superintendent of the
Scottish National Institution for the Education of Imbecile Children
at Larbert, Stirlingshire. With Engravings, 8vo, 14s. [1877]

MADNESS :

in its Medical, Legal, and Social Aspects, Lectures by Edgar
Sheppard, M.D., M.R.C.P., Professor of Psychological Medicine in
King's College; one of the Medical Superintendents of the Colney
Hatch Lunatic Asylum. 8vo, 6s. 6d. [1873]
INFLUENCE OF THE MIND UPON THE BODY

in Health and Disease, Illustrations designed to elucidate the Action
of the Imagination, by Daniel Hack Tuke, M.D., F.R.C.P.
8vo, 14s. [1872}

A MANUAL OF PRACTICAL HYGIENE,

by E. A. Parkes, M.D., F.R.S. Fifth Edition, by F. De Chaumont,
M.D., Professor of Military Hygiene in the Army Medical School.
8vo, with 9 Plates and 112 Engravings, 18s. . [1878]

A HANDBOOK OF HYGIENE AND SANITARY SCIENCE,

by George Wilson, M.A., M.D., Medical Officer of Health for Mid-
Warwickshire. Third Edition, post 8vo, with Engravings, 10s. 6d.

[1877]

SANITARY EXAMINATIONS

of Water, Air, and Food. A Vade Mecum for the Medical Officer of
Health, by Cornelius B. Fox, M.D., Medical Officer of Health of
East, Central, and South Essex. With 94 Engravings, crown 8vo,
12s. 6d. [1878]

DANGERS TO HEALTH:

A Pictorial Guide to Domestic Sanitary Defects, by T. Pridgin
Teale, M.A., Surgeon to the Leeds General Infirmary. With 55
Lithographs, 8vo, 10s. [1878]

MICROSCOPICAL EXAMINATION OF DRINKING WATER :

A Guide, by John D. Macdonald, M.D., F.R.S. , Assistant Pro-
fessor of Naval Hygiene, Army Medical School. 8vo, with 24 Plates,
7s. 6d. [1875}

CATALOGUE OF RECENT WORKS

HANDBOOK OF MEDICAL AND SURGICAL ELECTRICITY,
by Herbert Tibbits, M.D., F.R.C.P.E., Senior Physician to the
West London Hospital for Paralysis and Epilepsy. Second Edition,
8vo, with 95 Engravings, 9s. [1877]

BY THE SAME AUTHOR.

A MAP OF ZIEMSSEN'S MOTOR POINTS OF THE HUMAN BODY :
a Guide to Localised Electrisation. Mounted on Rollers, 35 X 21.
With 20 Illustrations, 5s. [1877]
GLINICAL USES OF ELECTRICITY ;

Lectures delivered at University College Hospital by J. Russell
Reynolds, M.D. Lond., F.R.C.P., F.R.S., Professor of Medicine
in University College. Second Edition, post 8vo, 3s. 6d. £1873]
MEDICO-ELECTRIC APPARATUS :

A Practical Description of every Form in Modem Use, with Plain
Directions for Mounting, Charging, and Working, by Salt & Son,
Birmingham. Second Edition, revised and enlarged, with 33 Engrav-
ings, 8vo, 2s. 6d. [1877]
A DICTIONARY OF MEDICAL SCIENCE ;

containing a concise explanation of the various subjects and terms of
Medicine, &c. ; Notices of Climate and Mineral Waters ; Formulae for
Officinal, Empirical, and Dietetic Preparations ; with the Accentuation
and Etymology of the terms and the French and other Synonyms, by
Robley Dtjnglison, M.D., LL.D. New Edition, royal 8vo, 28s. [1874]
A MEDICAL VOCABULARY ;

being an Explanation of all Terms and Phrases used in the various
Departments of Medical Science and Practice, giving their derivation,
meaning, application, and pronunciation, by Robert G. Mayne, M.D.,
LL.D. Fourth Edition, fcap 8vo, 10s. [1875]
ATLAS OF OPHTHALMOSCOPY,

by R. Liebreich, Ophthalmic Surgeon to St. Thomas's Hospital.
Translated into English by H. Rosborotjgh Swanzy, M.B. Dub.
Second Edition, containing 59 Figures, 4to, £1 10s. [1870]
DISEASES OF THE EYE :

a Manual by C. Macnamara, F.R.C.S., Surgeon to Westminster Hos-
pital. Third Edition, fcap. 8vo, with Coloured Plates and Engravings,
12s. 6d. [1876]
DISEASES OF THE EYE :

A Practical Treatise by Haynes Walton, F.R.C.S., Surgeon to St.
Mary's Hospital and in charge of its Ophthalmological Department.
Third Edition, 8vo, with 3 Plates and nearly 300 Engravings, 25s.

Q875]

HINTS ON OPHTHALMIC OUT-PATIENT PRACTICE,

by Charles Higgens, F.R.C.S., Ophthalmic Assistant Surgeon to,
and Lecturer on Ophthalmology at, Guy's Hospital. Second Edition,
fcap. 8vo, 3s.

PUBLISHED BY J. AND A. CHURCHILL

OPHTHALMIC MEDICINE AND SURGERY :

a Manual by T. Wharton Jones, F.R.C.S., F.R.S., Professor of Oph-
thalmic Medicine and Surgery in University College. Third Edition,
fcap. 8vo, with 9 Coloured Plates and 173 Engravings, 12s. 6d. [1865]

DISEASES OP THE EYE:

A Treatise by J. Soelberg Wells, F.R.C.S., Ophthalmic Surgeon to

King's College Hospital and Surgeon to the Royal London Ophthalmic

Hospital. Third Edition, 8vo, with Coloured Plates and Engravings,

25s. i [WS]

BY THE SAME AUTHOR,

LONG-, SHORT, AND WEAK SIGHT,

and their Treatment by the Scientific use of Spectacles. Fourth
Edition, 8vo, 6s. [*873]

A SYSTEM OF DENTAL SURGERY,

by John Tomes, F.R.S., and Charles S. Tomes, M.A., F.R.S., Lec-
turer on Dental Anatomy and Physiology at the Dental Hospital of
London. Second Edition, fcap Svo, with 268 Engravings, 14s. [1873]

DENTAL ANATOMY, HUMAN AND COMPARATIVE :

A Manual, by Charles S. Tomes, M.A., F.R.S., Lectm-er on Dental

Anatomy and Physiology at the Dental Hospital of London. With

179 Engravings, crown 8vo, 10s. 6d. [1876]

A MANUAL OF DENTAL MECHANICS,

with an Account of the Materials and Appliances used in Mechanical

Dentistry, by Oaklet Coles, L.D.S., R.C.S., Surgeon-Dentist to
the Hospital for Diseases of the Throat. Second Edition, crown 8vo,
with 140 Engravings, 7s. 6d. [1876]
HANDBOOK OF DENTAL ANATOMY

and Surgery for the use of Students and Practitioners by John
Smith, M.D., F.R.S. Edin., Surgeon-Dentist to the Queen in Scotland.
Second Edition, fcap 8vo, 4s. 6d. [1871]

STUDENT'S GUIDE TO DENTAL ANATOMY AND SURGERY,
by Henry Sewill, M.R.C.S., L.D.S., late Dentist to the Webt London
Hospital. With 77 Engravings, fcap. Svo, 5s. 6d. [1876]

OPERATIVE DENTISTRY:

A Practical Treatise, by Jonathan Taft, D.D.S., Professor of Opera-
tive Dentistry in the Ohio College of Dental Surgery. Third Edition,
thoroughly revised, with many additions, and 134 Engravings, Svo,
18s. [1877]

DENTAL CARIES

and its Causes : an Investigation into the influence of Fungi in the
Destruction of the Teeth, by Drs. Leber and Rottenstein. Trans-
lated by H. Chandler, D.M.D., Professor in the Dental School of
Harvard University. With Illustrations, royal Svo, 5s. I L878]

EPIDEMIOLOGY ;

or, the Remote Cause of Epidemic Diseases in the Animal and in the
Vegetable Creation, by John Parkin, M.D., F.R.C.P.E. Part I,
Contagion— Modem Theories— Cholera— Epizootics. Svo, 5s. [1873]

The following Catalogues issued by Messrs Churchill
will be forwarded post free on application :

1. Messrs Churchill's General List of nearly GOO works on
Medicine, Surgery, Midwifery, Materia Medica, Hygiene,
Anatomy, Physiology, Chemistry, Sfc, §c, with a complete
Index to their Titles, for easy reference. N.B.- — This List
includes Nos. 2 and 3.

2. Selection from Messrs ChurchiWs General List, com-
prising all recent Works published by them on the Art and
Science of Medicine.

3. A selected and descriptive List of Messrs ChurchiWs
Works on Chemistry, Materia Medica, Pharmacy, Botany,
Photography, Zoology, the Microscope, and other branches
of Science.

4. The Medical Intelligencer, an Annual List of New
Works and New Editions published by Messrs J. Sf A.
Churchill, together with Particulars of the Periodicals issued
from their Souse.

[Sent in January of each year to every Medical Practitioner in
the United Kingdom whose name and address can be ascertained.
A large number are also sent to the United States of America,
Continental Europe, India, and the Colonies.]

Messes CHURCHILL have a special arrangement with Messes
LINDSAY & BLAKISTOJST, oe Philadelphia, in accordance with
which that Pirm act as their Agents for the United States of America,
either keeping in Stock most of Messrs Chuechill's Books, or reprint-
ing them on Terms advantageous to Authors. Many of the Works in
this Catalogue may therefore be easily obtained in America.

PRINTED BY J. E. AD I/ABB, BARTHOLOMEW CLOSE.

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