The extermination of rats has become one of the serious prob

TREASURY DEPARTMENT

Public Health and Marine-Hospital Service of the United States

THE RAT AND ITS RELATION TO THE PUBLIC HEALTH

BY VARIOUS AUTHORS

PREPARED BY DIRECTION OF THE SURGEON-GENERAL

[Illustration: [Logo]]

WASHINGTON

GOVERNMENT PRINTING OFFICE

1910

TABLE OF CONTENTS.

Page.

_Introduction_ (Walter Wyman) 9

_Natural History of the Rat_ (David E. Lantz) 15

Classification of rats 15

Distribution of the genus _Mus_ in America 17

History of the brown rat 19

General description of the species in America and key to the

species 20

Habits of rats 22

Breeding habits 22

Abundance 23

Migrations and invasions 24

Food 26

Feeding habits 26

Ferocity 27

_Plague Infection in Rats_ (George W. McCoy) 29

Mode of examination 30

Gross lesions of natural rat plague, acute 32

Subcutaneous injection 32

The bubo 33

The granular liver 34

The spleen 35

Pleural effusion 35

Gross lesions of natural rat plague, chronic 36

Rat plague without gross lesions 37

Microscopical examinations 37

Bacteriological diagnosis of rat plague 38

Pest-like bacteria found in rats 41

Artificial infection of rats with plague 41

Modes of infection 42

Local reaction 43

The bubo 44

The liver and spleen 44

Chronic plague due to artificial inoculation 44

The histology of rat plague 45

Natural rat plague 46

Immunity of rats to plague 46

References 48

_Rat Leprosy_ (Walter R. Brinckerhoff) 49

Introduction 49

Review of literature 49

Description of disease 50

Etiology 52

Summary 52

Bibliography 53

_Bacterial Diseases of the Rat other than Plague_ (Donald H.

Currie) 55

Danysz bacillus or bacillus typhi Murium of Loeffler 55

Pneumonia 55

Staphylococcus abscesses 56

Bacillus pseudo-tuberculosis rodentium (Pfeiffer) 57

Toyama’s bacillus 57

Infections of mice 57

_Organic Diseases of the Rat, Including Tumors_ (George W. McCoy) 59

Usefulness of wild rats for laboratory purposes 59

Circulatory apparatus 60

Pulmonary apparatus 60

Digestive tract 61

Cirrhosis of the liver 61

Fatty degeneration of the liver 61

Hernia 61

Genito-urinary tract 62

Nephritis 62

Abscess of the kidney 62

Atrophy of the kidney 63

Vesical calculi 63

Tumors 64

Metastases 67

Histological structure 67

Lipomata 67

Fibromata 67

Sarcomata 67

Adenomata and Carcinomata 67

_Ectoparasites of the Rat_ (Nathan Banks) 69

Fleas—Siphonaptera 69

Lice—Anoplura 77

Mites—Acarina 80

_Internal Parasites of Rats and Mice in Their Relation to

Diseases of Man_ (Ch. Wardell Stiles and Charles G. Crane) 87

Summary 87

Introduction 87

Protozoa 88

Cestoda 95

Nematoda 101

Acanthocephala 108

_Compendium of Animal Parasites Reported for Rats and Mice—(Genus

Mus)_ (Ch. Wardell Stiles and Albert Hassall) 111

_The Flea and Its Relation to Plague_ (Carroll Fox) 123

Theories as to the transmission of plague 123

Insects that have been suspected in the transmission of plague 124

Experiments proving that fleas can transmit plague 125

The bacillus in the flea 126

How the flea clears itself of bacilli 127

Regional distribution of fleas on rats 127

Anatomy of the mouth parts of the Ceratophyllus Fasciatus 128

Outside the head 128

Inside the head 129

The act of biting 131

How the flea infects its host 132

Enumeration of fleas that have been found on rats 133

Results of identification of fleas in California 135

Synopsis of fleas commonly found on rats 136

Ceratophyllus Fasciatus, Bosc 136

Lœmopsylla Cheopis, Rothschild 138

Ctenopsyllus Musculi, Dugés 140

Pulex Irritans, Linnæus 142

Ctenocephalus Canis, Curtis 143

References 144

_Rodents in Relation to the Transmission of Bubonic Plague_

(Rupert Blue) 145

Epidemiological observations in San Francisco 147

Theories as to the cause of seasonal prevalence 149

The occurrence of plague in the marmot of Asia and ground

squirrel of California 150

Plague infection in ground squirrels 150

The natural habitat of plague 151

References 152

_Rodent Extermination_ (Wm. Colby Rucker) 153

Trapping 154

Poisoning 156

Natural enemies 159

Cutting off of the rat’s food supply 160

Building the rat out of existence 161

_Natural Enemies of the Rat_ (David E. Lantz) 163

Animals that destroy rats 163

Hawks 163

Owls 164

Wild mammals 166

Skunks 166

Weasels 166

Minks 167

Domestic animals 167

Dogs 167

Cats 167

Ferrets 168

Other animals 168

Mongoose 168

Alligators 168

Snakes 169

Bounties on predatory animals 169

_Rat-Proofing as an Antiplague Measure_ (Richard H. Creel) 171

Rat-proofing of primary importance 173

Rat-proofing is expensive 174

Methods of rat-proofing 175

Rat-proofing ordinances should be specific 177

Choice of architecture and building materials 178

_Inefficiency of Bacterial Viruses in the Extermination of Rats_

(Milton J. Rosenau) 179

Introduction 179

Experiments upon rat virus in the Hygienic Laboratory 183

Experiments with microorganisms for destroying rats by the U.

S. Biological Survey 186

Experiments during the San Francisco plague outbreak 188

Opinions of others 190

Pathogenicity for man 193

References to the literature 201

Résumé 204

_Plague Eradication in Cities by Sectional Extermination of Rats

and General Rat-Proofing_ (Victor G. Heiser) 205

_The Rat in Relation to Shipping_ (Wm. C. Hobdy) 207

Adaptability of the rat to his surroundings 208

Damage to cargo 209

Fumigation 211

Summary 213

_The Rat as an Economic Factor_ (David E. Lantz) 215

Introduction 215

Utility of the rat 215

Destructiveness of the rat 216

Grains 216

Merchandise in stores and warehouses 218

Merchandise in transit 219

Poultry and eggs 219

Game and wild birds 220

Fruit and vegetables 221

Flowers and bulbs 221

Fires 222

Buildings and furniture 222

Miscellaneous 223

Amount of losses caused by rats 224

Indirect losses 225

_The Rat in Relation to International Sanitation_ (John W. Kerr) 227

International sanitary regulations 228

Inquiry into the crusade against rats throughout the world 230

Rat extermination in United States ports 231

Rat extermination in Chinese cities 232

Rat extermination in Madras, Bombay, and Calcutta, India 234

Rat extermination in Yokohama and Nagasaki, Japan 235

Rat extermination in East Africa 237

Rat extermination in Cape Town, South Africa 238

Rat extermination in Alexandria and Cairo, Egypt 238

Extermination of rats at the port of Constantinople 238

Rat extermination in Russian ports 239

Destruction of rats in Trieste, Austria 240

Destruction of rats in Genoa, Italy 240

Destruction of rats in Barcelona, Spain 241

Rat destruction in French ports 241

Ministerial decree relating thereto 242

Destruction of rats in German ports 243

Measures against rats in Rotterdam, Holland 245

Destruction of rats at Antwerp, Belgium 245

Destruction of rats in Denmark 245

Danish law of March 22, 1907 245

Collection and destruction 247

Destruction of rats in Swedish ports 248

Destruction of rats in English ports 249

Measures against rats in Australian ports 250

Measures against rats in South American ports 252

Measures against rats in West Indian ports 252

Destruction of rats in Panama 253

Measures against rats in Vancouver, B. C. 253

Necessity of concerted action of nations 254

LIST OF ILLUSTRATIONS.

Page.

Fig. 1a. Upper molars of the brown rat (_Mus_): tubercles in 16

three rows

Fig. 1b. Upper molars of the rice rat (_Oryzomys_): tubercles in 16

two rows

Fig. 2a. Right hind foot of brown rat, showing long sixth foot 17

pad

Fig. 2b. Right hind foot of house mouse, showing round sixth 17

foot pad

Fig. 3a. Ears of brown rat and black rat, showing relative size 21

fig. 3b.

Fig. 4. Necropsy appearance of normal rat 48

Fig. 5. Necropsy appearance of plague-infected rat 48

Fig. 6. Flea, showing the various parts 70

Fig. 7. Louse—_Polyplax spinulosus_ 78

Fig. 8. Mite—_Lælaps echidninus_ 81

Figs. 9 Internal parasites of rats and mice 90–109

to 58.

Fig. 59. Isolated plague-infected center, Manila, P. I. 206

Fig. 60. Scheme for testing rat-plague infection, Manila, P. I. 206

Plate I. Mouth parts of _Ceratophyllus fasciatus_ 130

II. _Ceratophyllus fasciatus_ 136

III. _Lœmopsylla cheopis_, Rothschild 138

IV. _Ctenopsyllus musculi_, Duges 140

V. _Pulex irritans_, Linnæus 142

VI. _Ctenocephalus canis_, Curtis 144

THE RAT AND ITS RELATION TO THE PUBLIC HEALTH.

By WALTER WYMAN,

_Surgeon-General of the Public Health and Marine-Hospital Service_.

INTRODUCTION.

The science of bacteriology has elucidated many facts with respect to

the causation of disease, and with this advance in knowledge, old

theories regarding the miasmatic and humoral origin of human ills have

been abandoned.

Epidemiological studies have likewise determined the methods of

transmission of many of the infectious and contagious diseases, thus

eliminating erroneous conceptions that they are attributable to some

mysterious condition of the atmosphere or soil, or to a visitation of

the wrath of the Almighty.

Both these sciences have contributed to our knowledge of the

relationship of living things, particularly with respect to their

influence upon each other in relation to health and disease. It is now

known, for instance, that mosquitoes are the pests of man, not only

because of their bites, but because they at times transmit malaria,

dengue, filariasis, and yellow fever. So, too, it is known that rodents

are the enemies of man, not only because of the toll exacted from him,

but because they are the principal agents in the propagation and spread

of bubonic plague.

Ancient writings abound in allusions to pestilences and their connection

with epizootics among rats and mice.

In the Book of Samuel there is reference to a pestilence having relation

to mice, and that it might be stayed the Philistines made offerings of

golden images of the mice that marred the land.

During the centuries that have intervened rats have migrated to

practically every quarter of the earth, causing untold losses on account

of their depredations. They have also, in all probability, been the

primary agents of transmission in the pandemics of plague which have

visited the earth. The fact that plague is due to a specific

microorganism, and that its presence in man is also associated with

epizootics in rats, has led to a more careful study of this animal,

particularly in relation to his habits, the diseases from which he

suffers, and the methods necessary to his control. Prior to the

beginning of the present pandemic of plague which had its origin in

China, interest in the rat was almost wholly an economic and financial

one. Since that time evidence has been rapidly accumulating which proves

that this animal and his parasites are responsible for the transmission

of plague and that plague itself is essentially a disease of the rat.

A knowledge of this animal on the part of the sanitarian therefore

becomes essential. During the enforcement of antiplague measures in

California, Hawaii, the Philippine Islands and elsewhere, observations

of great value have been made and their practical application has

resulted in better directed efforts for the elimination of the disease.

In studies of plague and leprosy with the view to their diagnosis and

control, it is not enough now to isolate the microorganisms responsible

for these diseases, but the sanitarian must be able to recognize the

pathological conditions present in animals affected, and to do so he

must have practical knowledge of this subject in order that he may

differentiate between the various diseases from which these animals

suffer.

Opportunity for observation and study of the diseases of rats and the

methods necessary to their eradication has been afforded to the officers

of the Public Health and Marine-Hospital Service who are constantly

stationed on the outposts in the warfare against exotic diseases. The

results of these observations have been utilized by officers of the

service, and some of them have been published for the benefit of others.

The rat has received much attention of late in other parts of the world.

In Denmark, for instance, a legalized warfare against rodents has been

begun, principally on account of their influence in the transmission of

trichinosis. In England there exists The Incorporated Society for the

Destruction of Vermin, and in other places rat destruction is being

agitated both from economic and public health standpoints.

In view of the great importance of the rat in relation to the public

health, it has been thought advisable to collect and publish all

pertinent information on the subject, in order that public health

officials who should be on the lookout for the appearance of plague

among rodents might have available a reliable treatise on the subject.

Studies of rodents from a biologic and economic standpoint come within

the province of other departments of the public service, and the

cooperation of the Biological Survey and Bureau of Entomology of the

Department of Agriculture was therefore requested and received.

The subjects dealt with in this publication have been prepared by those

having wide experience.

In the chapter on natural history by Mr. David E. Lantz there is given a

classification of rats as well as the distribution of the genus _Mus_ in

America. An interesting and important fact is mentioned that the

Biological Survey has no records of the presence of the brown rat in

Nevada, Utah, Wyoming, Idaho, and the greater part of Montana. Mr. Lantz

also describes the different species in America, and refers to their

habits as to breeding, feeding, migrations, invasions, and ferocity. The

facts presented by him emphasize the great difficulty of ridding cities

of these pests.

Passed Assistant Surgeon McCoy discusses plague infections in rats and

describes the methods of examination. He also describes the gross

lesions found in plague rats, gives the bacteriologic diagnosis of rat

plague and the cultural characteristics of the plague bacillus on

various media. He gives the methods of artificial infection of rats with

plague, and reviews the recent work of Ledingham in relation to the

histology of rat plague. Finally, he presents results of his own

investigations to show that the wild rat is not especially susceptible

to plague infection, and that a certain percentage of such animals enjoy

a natural immunity to plague.

Doctor Brinckerhoff discusses rat leprosy; states that it is very

similar to human leprosy, and that it is caused by a bacillus which

closely resembles the bacillus of Hansen. He describes the pathological

changes found, and expresses the hope that the disease will receive

further earnest study, in order that additional information may throw

light on the problems presented by leprosy in man.

Passed Assistant Surgeon Currie briefly outlines the bacterial diseases

of the rat, other than plague and leprosy. He mentions the great utility

that would follow the discovery of a rat destroying bacterium, but

states that it appears now more than probable that few such natural

diseases of rats exist.

In a chapter on organic diseases of the rat, Doctor McCoy summarizes the

results of his observations made during examinations of these animals in

the Federal laboratory of the service at San Francisco. These

observations are of interest, and will assist those engaged in such work

to further classify the pathological changes noted as well as

differentiate them from plague.

The ectoparasites of the rat are classified and described by Mr. Nathan

Banks, and he has presented in condensed form information of much

practical value upon the subject.

Dr. Ch. Wardell Stiles discusses the internal parasites of rats and mice

in relation to the diseases of man. He regards the rat as a permanent

reservoir for trichinosis, and states that this disease will probably

never be eradicated from man until rats and mice are practically

eradicated, and a national campaign directed against trichinosis must

take the rat into consideration.

A compendium of animal parasites reported for rats and mice is presented

in a chapter by Ch. Wardell Stiles and Albert Hassall. While, as the

authors state, no list of this kind can ever lay claim to being

complete, it represents the present knowledge of the subject.

In a discussion of the flea and its relation to plague, Passed Assistant

Surgeon Fox summarizes the theories as to the transmission of this

disease. He also mentions the insects that have been suspected of

transmitting plague and presents accumulated evidence that fleas

actually convey the infection. He then gives the anatomy of the mouth

parts of the _Ceratophyllus Fasciatus_, the common rat flea of North

America. He also enumerates the fleas that have been found on rats, and

gives the results of identifications of 19,768 fleas in San Francisco

and Oakland, Cal. The plates accompanying this article, and their

description should be of great value to those engaged in antiplague

measures.

Surgeon Blue briefly discusses the subject of rodents in relation to the

transmission of bubonic plague. He discusses the theories as to the

cause of seasonal prevalence of this disease and presents a table

showing the number of rats examined during the different months of the

year, the number found infected, the average temperature and rainfall

for those months and the character of the days, as to the number clear,

partly cloudy, or cloudy. He refers to plague infection in ground

squirrels in California and warns against the possibility that this

animal may become responsible for the establishment of a permanent focus

of plague on the Pacific coast of the United States, as the marmots are

so concerned with regard to India.

The all-important subject of rodent extermination is considered in

detail, various phases of the subject being dealt with by different

authors.

Passed Assistant Surgeon Rucker discusses the destruction of these

animals by trapping, poisoning, cutting off of food supply, and

destroying of existing nests and at the same time preventing the making

of new ones. He describes the methods of use of the various mineral

poisons, but finally states that rodents must be builded out of

existence; in other words, habitations must be rendered rat proof.

Mr. Lantz, in discussing the natural enemies of the rat, mentions the

animals that destroy these pests. He concludes that on account of this

function bounties for the destruction of small animals that prey on

rodents can not be justified and that they should in the future be

protected in every way possible.

Passed Assistant Surgeon Creel discusses rat proofing as an antiplague

measure, and gives in detail the principles of construction necessary.

He concludes that rat proofing is the most valuable antiplague measure,

and that it should precede auxiliary measures such as trapping and

placing of poisons.

Surgeon Rosenau discusses the bacterial viruses in relation to rat

destruction. As a result of his investigations in the hygienic

laboratory and the reports of investigations and practical use

elsewhere, he concludes that the bacterial viruses have signally failed

to accomplish the mission for which they were intended, and that they

are not entirely harmless to man, as has been stated.

Passed Assistant Surgeon Heiser briefly outlines the measures

recommended for the eradication of plague in cities by means of

sectional extermination of rats and general rat proofing. He gives

results following this method of procedure in Manila, and presents

charts showing how to deal with infected city districts.

Passed Assistant Surgeon Hobdy, in a chapter on the rat in relation to

shipping, refers to the voyage-making tendencies of the rodent, its

destructiveness aboard ship, and its power of adapting itself to unusual

conditions and surroundings. In one small lumber vessel fumigated by

Doctor Hobdy at the Angel Island quarantine station there were collected

525 dead rats. Mention is also made of another vessel on which were

collected 1,700 rats after fumigation. He discusses the methods by which

it gains access to vessels, and outlines the practices that should be

observed to keep it off. He also describes in some detail the measures

to be adopted for its destruction after it has gotten aboard ship, and

mentions the different methods of fumigation.

Mr. Lantz, in a third paper, discusses the rat as an economic factor,

and states in his paper that they do not serve any useful purpose. On

the other hand, they cause enormous loss through damage to grain,

merchandise, poultry and eggs, game and wild birds, fruit and

vegetables, and flowers and bulbs. They also cause damage by setting

fire to buildings and destroying furniture. He refers to various

estimates made of the losses in the United States from rats, and they

vary from $35,000,000 to $50,000,000 a year; but at the same time he

states that, with present information, any attempt to state the amount

of loss from rats would be largely guesswork.

Assistant Surgeon-General Kerr refers to the rat as a factor in

international sanitation, and briefly outlines the provisions contained

in international sanitary agreements for their eradication. He reviews

the efforts being made at the more important seaports to exterminate

rats, as well as the methods being employed to that end. The information

presented is, in part, compiled from consular reports received through

the Department of State. There are given, so far as obtainable, copies

of laws and ordinances enacted for the destruction of rats and the

different methods practiced in ports where plague has prevailed, and the

facts presented indicate that a more or less widespread crusade against

rats is being carried on. He expresses the belief that it is too much to

expect that the rat population can ever be exterminated from any city,

but that it is not too much to expect that ocean carriers can be freed

from rodents and kept so, which action would confine plague within

continental boundaries.

Epidemiological studies made of plague since the adoption of the

International Sanitary Convention of Paris and the International

Sanitary Convention of Washington have proven that the rat and its

parasite, the flea, are the agents of transmission of the disease. In

other words, where rats go plague will go. I believe, therefore, that in

order to stop the further progress of plague, radical measures should be

adopted, and in a communication of February 26, 1909, addressed to the

Secretary of State, I suggested the advisability of submitting the

question of a systematic destruction of rodents aboard ship to an

international sanitary conference, with the view to the adoption of an

international sanitary regulation on the subject. The adoption of such a

regulation would undoubtedly lessen quarantine restrictions, prevent the

destruction of cargo by rodents, and obviate the danger of the further

spread of plague.

Until ships are freed from rats, each country must take all necessary

precautions, consistent with international agreements, to destroy rats;

and the sanitary authorities of infected localities must, at great

expense, determine the extent of infection among rodents, with the view

to its elimination. This problem when it presents itself in a community

is of great magnitude, and those responsible for its solution should be

familiar with all its phases.

It is with the view to supplying the necessary information in one

treatise that this publication is issued. In its preparation the bureau

has had the cooperation of the Department of Agriculture and

acknowledgements are due, and here made to, the officers of that

department for their hearty cooperation in contributing some of the

chapters which follow.

NATURAL HISTORY OF THE RAT.

By DAVID E. LANTZ.

_Assistant, U. S. Biological Survey, Department of Agriculture_.

INTRODUCTION.

The extermination of rats has become one of the serious problems of

modern times. That such noxious animals should have flourished so long

is not creditable to our civilization. While no kind of rat can be

regarded as harmless, the various species differ greatly in harmfulness.

In comparison with the cosmopolitan species that have reached our shores

from the Old World, our native rats do little damage. It is important,

therefore, to be able to recognize the introduced forms, to understand

their habits, and to concentrate efforts for their extirpation.

CLASSIFICATION OF RATS.

Rats and mice belong to the _Rodentia_, an order which comprises more

than a third of all living species of mammals. Also, it exceeds any

other mammalian order in the number of its individuals.

Rodents are mainly herbivorous mammals, mostly of small size, having a

furry, sometimes a spiny, integument, clawed digits, and usually

plantigrade feet. The most important distinguishing character of the

order is its dentition. This is marked by the absence of canine teeth

and the presence of strongly developed incisors growing from permanent

pulps. The incisors are never more than two in the lower jaw and usually

but two in the upper. They are elongated, curved, chisel-like in shape,

and continue to grow throughout the life of the animal. Only the front

of these teeth is covered with enamel, a provision which keeps them

sharp by the more rapid wearing away of the softer dentine in the body

of the tooth, as the upper and lower pairs meet in gnawing. Between the

incisors and the cheek, or molar, teeth of rodents there is a wide,

vacant space, marking the entire absence of canines.

The most extensive family of rodents is the _Muridæ_, a name which

applies to rats and mice in the widest sense of those terms. It is

difficult to characterize the family, since its members differ widely.

However, most of them are rat-like in form and light and active in

movements. None of the family have premolars; and, except in a single

genus (_Hydromys_), the number of molars is three. Oldfield Thomas, the

eminent English zoologist, includes in this family no less than 77

genera, or almost half the total of 159 which he ascribes to the whole

order _Rodentia_.[A] He further subdivides the _Muridæ_ into a dozen

subfamilies, of which the _Murinæ_ and the _Sigmodontinæ_ are the most

extensive. The name _Cricetinæ_ is now generally used instead of

_Sigmodontinæ_, though not always with the same limitations.

Footnote A:

Proc. Zool. Soc., pp. 1012–1028, 1896.

The _Murinæ_ comprise only Old World rats and mice, while the

_Cricetinæ_ are, in the main, American forms. In the _Murinæ_ the cusps,

or tubercles, of the unworn upper molars are arranged triserially, or in

three longitudinal rows; in the _Cricetinæ_ they are arranged

biserially, one row on the outer and one on the inner margin (fig. 1).

The wearing away of these cusps leaves characteristic curved lines of

hard enamel surrounding areas of dentine. When the cusps are in pairs

the worn pattern looks somewhat like the Greek letter sigma (Σ), whence

the name sigmodont, often applied to native American rats and mice.

[Illustration:

Fig. 1a. Fig. 1b.

FIG. 1a.—Upper molars of the brown rat (_Mus_): tubercles in three

rows.

FIG. 1b.—Upper molars of the rice rat (_Oryzomys_): tubercles in two

rows.

]

The _Murinæ_ are the true rats and mice, typified by the genus _Mus_,

which contains by far the largest number of species. Trouessart, in his

Catalogus Mammalium, enumerates 260 species of _Mus_ described before

1905. Since that date a number of new forms have been described.

The genus _Mus_ is characterized by narrow, ungrooved incisors; three

small, rooted molars; soft fur mixed with hairs, sometimes with spines;

a rudimentary pollex having a short nail instead of a claw; a long tail

bearing rings of overlapping scales and often naked or nearly so. The

ears are rather large, the eyes bright and prominent, and the muzzle

somewhat pointed. The members of the genus are natives of the Old World,

throughout which, with the exception of Madagascar, they are quite

generally distributed. Nearly seven-eighths of the whole number of

species are commonly called rats.

The distinction between rats and mice is arbitrary and based on size.

Exclusive of the tail, rats may be said to vary in length from 4½ to 10

inches or more, while mice measure from 2 to 4 inches. With few

exceptions, rats have six well-defined footpads (plantar tubercles), the

last on the hind foot being elongated in shape; the last hind-foot pad

of mice is usually circular (fig. 2).

Of the many species of _Mus_ only three or four have developed the

ability to adapt themselves to such a variety of conditions as to become

cosmopolitan. Four have found lodgment in America: The common house

mouse (_Mus musculus_); the old English black rat (_Mus rattus_); the

Egyptian, or roof, rat (_Mus alexandrinus_); and the brown rat (_Mus

norvegicus_), known also as the gray rat, barn rat, wharf rat, sewer

rat, and Norway rat. The black rat and the roof rat differ from each

other chiefly in color. Indeed some zoologists regard them as races of

the same species, and the trinomial _Mus rattus alexandrinus_ for the

roof rat is now in use among zoologists.

[Illustration:

FIG. 2a.—Right hind foot of brown rat, showing long sixth foot pad.

FIG. 2b.—Right hind foot of house mouse, showing round sixth foot pad.

]

DISTRIBUTION OF THE GENUS MUS IN AMERICA.

The common house mouse (_M. musculus_) found its way to America soon

after the first settlement by Europeans. It now inhabits all settled

parts of North and South America, as well as nearly the entire Old

World; but in very cold regions it does not always survive the winters,

and is therefore comparatively scarce or local. It almost always reaches

a new settlement sooner than the rat.

The black rat (_M. rattus_) has been known in Europe since the twelfth

century. It was carried to South and Middle America about three and a

half centuries ago (1554). The time of its arrival in the English

colonies of North America is not known with certainty, but it was well

established in the settled parts by the beginning of the eighteenth

century. Soon after the arrival of the brown rat, the black species

began to decrease in numbers, and has become extinct in most localities.

At present it is not uncommon in some parts of the South, and still

occurs in scattered colonies in Canada and some of the States east of

the Mississippi, and also on some of the coastal islands. It is

occasional in many of our seaports, being apparently brought from the

Far East in merchandise. Except in a few ports like San Francisco, where

new arrivals are probably rather frequent, these introduced individuals

are often destroyed before they multiply. The history of the black rat

in Europe and its disappearance before the brown rat is an exact

parallel to its history here, and the animal is now comparatively rare

north of the Alps, except in the Channel Islands.

The Biological Survey has specimens of the black rat from Massachusetts,

New Hampshire, Georgia, Florida, Alabama, California, and Washington,

and also from Mexico, Honduras, Nicaragua, and Hawaii. There are

authentic records of its recent occurrence in Newfoundland, Quebec, Nova

Scotia, New York, North Carolina, Tennessee, West Virginia, and

Mississippi. In parts of South and Middle America it is abundant.

The roof or Alexandrian rat (_M. alexandrinus_) is similar to the black

rat in form and general habits, though not in color. Little is known of

its history, but it is thought to be a native of Egypt, where it is

still abundant. It has established itself in many parts of the world,

mainly in warm climates, and is common near the coast in the southern

parts of the United States.

The Biological Survey has specimens of the roof rat from North Carolina,

Georgia, Florida, Alabama, Mississippi, Texas, Arizona, and California.

In the last-named State it is abundant in the Sacramento Valley. It is

known also from Dismal Swamp, Virginia, and from Cuba, the Bermudas,

Trinidad, San Domingo, Costa Rica, Nicaragua, Mexico, and Hawaii. Also,

it inhabits many parts of South America, where in places it is the

dominant species.

The most destructive of the rat family is the brown rat (_M.

norvegicus_). In most parts of the United States it is the common rat

about houses and barns in the country and about markets, wharves, and

warehouses in cities. It is larger and more robust than either the black

or the roof rat, and differs from both in habits. It is more of a

burrower, and lives in excavations which it makes under buildings and in

loose soil along hedges and river banks. This habit, combined with its

greater strength and ferocity has enabled it to supplant the other

species in temperate latitudes; but in the warmer parts of America and

the Old World it has not been able to drive out the others. The house

mouse everywhere holds its own against the brown rat by its ability to

escape into retreats too small for the rat to follow.

The brown rat inhabits most of the thickly populated parts of America.

North of Panama it occurs generally except in the arid interior, from

the Isthmus to the Yukon Valley and southern Greenland. In the Great

Basin it is practically unknown, and in New Mexico and Arizona it is

confined chiefly to towns along the railroads. The Biological Survey is

without records of its presence in Nevada, Utah, Wyoming, Idaho, and the

greater part of Montana. The reason for its absence in that region is

not understood, but its ability to withstand extreme cold is proved by

the fact that it flourished in latitude 78° 37′ north on board Doctor

Kane’s ship _Advance_, in the Second Grinnell Expedition, during the two

winters when that vessel was icebound. It has also adapted itself to the

continuous low temperatures of cold-storage warehouses, in which it

appears to breed freely.

HISTORY OF THE BROWN RAT.

We know little of the history of this species. Greek and Roman writers

make no mention of rats of any kind, but possibly knew the animals and

included them in their frequent references to mice. Pallas, in 1778,

described the brown rat under the name _Mus decumanus_, and this was

generally used until it was found that Erxleben had called it _M.

norvegicus_ in 1777. Previously, the common name Norway rat had often

been used for this species.

The brown rat is generally supposed to be of Asiatic origin. Various

modern writers have asserted that it came originally from Persia or

India; but W. T. Blanford states that the species is at present unknown

in Persia, and that in India the black rat is the generally distributed

species, while the brown rat occurs only along the coast and the

navigable rivers.[B] This implies that the latter species is a

comparatively recent immigrant into India.

Footnote B:

Fauna of British India. Mammals, p. 409, 1891.

As regards the arrival of the brown rat in Europe, two facts are known.

The species reached England from some eastern port about 1728 or 1729,

and according to Pallas, a little earlier, 1727, crossed the Russian

frontier from Asia and soon spread over the greater part of that

country.[C] This statement, taken in connection with that of Blanford,

makes it highly probable that before this migration the Asiatic home of

the species was north, rather than south, of the high mountains of

northern India. This view, which has been adopted by several

naturalists, is further strengthened by the fact that the animal

flourishes better in temperate than in tropical climates.

Footnote C:

Zoographica, Rosso-Asiatica, vol. 1. p. 165, 1831.

Possibly earlier and unrecorded westward migrations of the brown rat

took place. A few years ago Professor Waile, the archeologist, while

making excavations at Cherchell on the coast of Algeria, dug up the

skull of a rat, which he stated was contemporary with the Roman

occupation of the country under the Cæsars. The skull had but one molar,

much worn, but the cranial bones were intact, and French zoologists

pronounced the remains as undoubtedly those of the “surmulot,” or brown

rat.[D] This shows that we have little more than conjectures for the

early history of this species.

Footnote D:

Comptes Rendus des Séances de L’Académie des Sciences, Paris, vol.

116, p. 1031, 1893.

The brown rat is said to have first appeared in Paris in 1750. It was

brought to the United States, probably from England, about the beginning

of the Revolution, 1775. According to Audubon, it was unknown on the

Pacific coast of the United States in 1851; but Dr. J. S. Newberry

thought it must have arrived at San Diego, Monterey, and San Francisco

at a much earlier date.[E] Doctor Cooper recorded its arrival at Fort

Steilacoom, Washington, as occurring about 1855.

Footnote E:

Pac. R. R. Reports, Vol. 6, Zoological Report, pt. 2, p. 60, 1857.

GENERAL DESCRIPTION.

The brown rat differs considerably from the black rat and the roof rat.

It is larger, has a shorter head, a more obtuse muzzle, smaller ears,

and a relatively shorter and stouter tail. The general color is

grayish-brown above and whitish below. The over hairs of the upper parts

have black tips. The tail is usually shorter than the head and body

combined. The average measurements of adult specimens of the brown rat

in the Biological Survey collections are as follows: Total length, 415

millimeters (16.3 inches); tail, 192 millimeters (7.1 inches); hind

foot, 43 millimeters (1.7 inches). This species sometimes attains a

total length of 19 to 20 inches, and has been known to weigh 24 to 28

ounces and even more. The average weight of an adult brown rat is

considerably less than a pound.

The black rat is less robust than the brown rat. It has a longer head, a

sharper muzzle, and larger and broader ears (fig. 3). The tail is longer

than the head and body combined. The fur is of a sooty, or plumbeous

black, color, paler on the underparts. It is much softer and denser than

that of the brown rat, and the mixture of very dark and lighter over

hairs gives it a peculiar shining appearance. The average measurements

of 20 apparently adult specimens in the collection of the Biological

Survey are as follows: Total length, 379 millimeters (14.9 inches);

tail, 207.4 millimeters (8.1 inches); hind foot, 35.8 millimeters (1.4

inches).

The roof rat in general resembles the black rat, except as to color and

texture of fur. Above it does not greatly differ in color from the brown

rat, but its underparts are usually more yellowish. The fur is commonly

shorter and harsher in texture than that of the black rat, but this

difference might not always be apparent in specimens of the two forms

from the same latitude. The average measurements of 50 adult specimens

of the roof rat in the collections of the Biological Survey are as

follows: Total length, 393.3 millimeters (15.5 inches); tail, 212.8

millimeters (8.4 inches); hind foot, 36.2 millimeters (1.4 inches).

[Illustration:

FIG. 3A, FIG. 3B.—Ears of brown rat and black rat, showing relative

size.

]

Both albinism and melanism are frequent among rats, and pied forms also

are common. It has been claimed that all the white rats (albinos) of the

bird stores are _Mus rattus_, but albinism is by no means confined to

this species. Doctor Hatai found that all the colonies of white rats

maintained at the neurological laboratories of Chicago University and

the Wistar Institute of Anatomy, Philadelphia, were of the _M.

norvegicus_ species.[F] The same is true of all the albino rats in the

collections of the National Museum and the Biological Survey. These

collections contain also several spotted rats (gray and white) and

sooty-black specimens indistinguishable in color from _M. rattus_, all

being undoubtedly of the _M. norvegicus_ species.

Footnote F:

Biological Bulletin, vol. 12, pp. 266–273, March, 1907.

KEY TO THE SPECIES OF MUS IN AMERICA.

Size small. Total length of adult less than 200

millimeters _Mus musculus_

Size large. Total length of adult exceeding 300

millimeters.

Ears moderate, when laid forward barely or not

reaching eye; tail shorter than (rarely equal

to) the length of head and body, darker above

than below; color of body normally gray-brown

above, white below; hind foot 38–46 millimeters _Mus norvegicus_

Ears larger, when laid forward reaching at least

to middle of eye; tail longer than head and

body, dusky all around; hind foot 33–37

millimeters.

Color grayish-brown above, white or yellowish _M. rattus

white below. alexandrinus_

Color blue-black above, slaty below _M. rattus_

HABITS OF RATS.

BREEDING HABITS.

Both climate and food supply affect the rate of multiplication of most

rodents. The rat probably increases more rapidly in a temperate and

equable climate than in one of great variability. Extremes of heat and

cold retard multiplication, decreasing both the number of litters in a

year and the number of young at a time. In northern latitudes,

apparently, more or less interruption of breeding occurs in the winter

months.

Where the country is well settled the food supply of rats is not likely

to be deficient; and when the animals have access to stores of grain,

the young mature very quickly and probably reproduce earlier than when

grain is absent.

The brown rat is more prolific than either the roof rat or the black

rat. The female brown rat has usually 12 mammæ—3 pairs of pectoral and 3

pairs of inguinal—although these numbers are not constant, one or more

teats frequently being undeveloped. The black rat and the roof rat have

only 10 mammæ—2 pairs of pectoral and 3 pairs of inguinal—with but

little tendency to vary. Records of actual observations on the number of

young confirm the deductions that might be drawn from the above facts.

At Bombay, India, during the recent investigations made by the India

Plague Commission, 12,000 rats were trapped and examined. The average

number of embryos found in pregnant brown rats was 8.1; the highest

number, 14. The average for the black rat was 5.2; the largest number,

9.[G]

Footnote G:

Etiology and Epidemiology of Plague, p. 9, Calcutta, 1908.

In temperate latitudes the average number of young produced by the brown

rat is undoubtedly greater. Instances of very large litters observed in

England are recorded in The Field (London). In two instances 22 and 23

young, respectively, were found in a single nest, though no evidence is

offered that these were the progeny of a single female; but in two other

cases 17 and 19 embryos were found in gravid females. A dealer in

feedstuffs in Washington, D. C., relates that he found 19 young rats in

a single nest in his store. Within the past few months the writer has

examined four pregnant brown rats taken in traps. The numbers of embryos

they contained were 10, 11, 11, and 13, respectively. While we have not

enough data for definite conclusions, we may safely state that the

average litter for this latitude is not less than 10.

Frank T. Buckland, in Curiosities of Natural History, relates that a

white rat which he kept in captivity gave birth to 11 young when only

eight weeks old. As gestation in rats occupies three weeks, this animal

must have bred when only five weeks old.

The number of times rats breed in a year is not definitely known, and

probably varies considerably with local conditions. Kolazy makes the

almost incredible statement that two female white rats, kept by him in

confinement and well fed, within thirteen months gave birth to 26

litters of young, numbering 180 in all. One of them produced young

regularly at intervals of 25 days.[H]

Footnote H:

Verh. Zool. Bot. Gesel. Wien., pp. 731–734, 1871.

The writer recently kept two young female brown rats with a male in a

large open cage for several months. One of the females gave birth to

young on April 15; the other on April 17. The number in these litters

was not observed, as some were devoured soon after birth, and all within

three days, presumably by the male rat. On May 23 both females gave

birth to young, 24 in number, all in one nest.

The known facts concerning the breeding of the brown rat may be briefly

summarized as follows: The animals breed from three to five times a

year, each time bringing forth from 6 to 19 young. After a gestation

period of twenty-one days, the females give birth to their young in

nests built in underground burrows or under floors, stacks, lumber,

woodpiles, or other shelter. The young are blind and naked when born,

but grow rapidly, and young females are capable of breeding when less

than three months old.

Early spring and summer are the periods of greatest reproductive

activity among rats. Young, however, are to be found every month of the

year.

The above statements apply in the main also to the black and the roof

rat, but the number of young in a litter is somewhat smaller. The newly

born young of the black rat have not the bright pink color of those of

the brown and the roof rat, but are bluish, especially on the upper

parts. Black-and-white spotted rats are at first bluish-and-red spotted,

the red areas representing the white of the adults.

ABUNDANCE OF RATS.

From the foregoing account of the breeding habits of rats, the great

difficulty of ridding cities or large areas of the animals may be

readily understood. Ordinarily, they breed more rapidly than they are

destroyed. Although few are seen in daytime, at night they fairly swarm

along river fronts and wharves, as well as in sewers, stables,

warehouses, markets, and other places where food is abundant. Their real

numbers may sometimes be discovered when any such harbor is demolished.

An ordinary farm sometimes supports an astounding number of rats. In

1901, an estate of 2,000 acres near Chichester, England, was badly

infested with the pests. They were systematically destroyed by traps,

poisons, and ferrets, under the supervision of the proprietor. In this

way 31,981 were killed, while it was estimated that tenants at the

thrashing had destroyed fully 5,000 more. Even then the property was by

no means free from rats.[I]

Footnote I:

The Field (London), vol. 100, p. 545, 1902.

During a plague of rats on the island of Jamaica in 1833, the number of

rats killed on a single plantation in a year was 38,000. The injury to

sugar cane on the island caused by the animals was at that time

estimated at half a million dollars a year.[J]

Footnote J:

New England Farmer, vol. 12, p. 315, 1834.

The report of the Indian Famine Commission presented to the English

Parliament in 1881 affords one of the best illustrations of the number

of rats that may infest a country. An extraordinary number of the

animals at that time inhabited the southern Deccan and Mahratta

districts of India. The autumn crop of 1878 and the spring crop of 1879

were both below the average, and a large portion of each was destroyed

by rats. The resulting scarcity of food led to the payment of rewards

for the destruction of the pests, and over 12,000,000 were killed.[K]

Footnote K:

British Medical Journal, Sept. 16, 1905, p. 623.

MIGRATIONS AND INVASIONS.

Migrations of rats have often been recorded. The brown rat is known in

Europe quite generally as the migratory rat. The Germans call it the

Wanderratte. Pallas narrates that in the autumn of 1727 this species

arrived from the east at Astrakhan, southeastern Russia, in such great

numbers and so suddenly that nothing could be done to oppose them. They

crossed the Volga in immense troops. The cause of this general migration

was attributed to an earthquake; but since similar movements of the same

species often occur without earthquakes, it is probable that only the

food supply of the animals was involved in the migration which first

brought the brown rat to Europe.

A seasonal movement of rats from houses and barns to the open fields

takes place in spring when green and succulent plant food is ready for

them. The return movement takes place in the autumn. This seasonal

migration is noticeable even in large cities.

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