CHAPTER VIII
RAT-FLEAS AND BAT-FLEAS
THE chief conclusions arrived at, as the result of the investigations, during the years 1905 to 1909, into the mode of spread of plague in India, may be briefly stated in the following fashion: The Advisory Committee, under whose direction the investigation was carried out, consider that: firstly, in nature, plague is spread among rats by the agency of rat-fleas; secondly, bubonic plague is not directly infectious from man to man; thirdly, in the great majority of cases, during an epidemic of plague, man contracts the disease from plague-infected rats through the agency of plague-infected rat-fleas; fourthly, where there are annual epidemics they occur during some part of that season when the prevalence of fleas is greatest.
That being so, it is manifest that an accurate knowledge of rat-fleas, their forms, their habits, and their life-history may prove of great importance.
Three species of the genus _Mus_ follow quickly in the wake of civilized man and establish themselves all over the globe. They may all be looked upon as more or less domestic animals. The house-mouse (_M. musculus_) is familiar everywhere. The old black rat (_M. rattus_) chiefly infests ships and seaports. The brown rat (_M. norwegicus_) is the most aggressive and distinctive. But all three, by accidental transference from port to port in ocean-going vessels, have become distributed over the world. Their fleas, to a limited extent, have become distributed with them. In connection with the spread of plague these three small rodents are of prime importance; and not less important are the fleas which are parasitic on them.
In California, the ground-squirrel (_Otospermophilus beecheyi_) has been proved to play an important part in plague infection; and a full account of its fleas, and of experiments in transferring rat-fleas to squirrels and squirrel-fleas to rats, has been published by American naturalists.
In 1903 Dr Blue, who was in charge of measures for suppressing plague at San Francisco, observed that an epidemic disease was killing the ground-squirrels in the country round San Francisco Bay. It was shown, somewhat later, that the mortality among the squirrels was caused by plague, and there can be little doubt that it was transferred from rats to squirrels. In harvest time rats migrate to the fields and use the same runs and holes as the squirrels. Under these conditions a transfer of fleas from rats to ground-squirrels is almost certain to ensue. Two species of flea have been recorded from the Californian ground-squirrel, and both are parasites of rats. The chain of evidence is really complete, for those who have made a business of hunting ground-squirrels testify to the readiness with which fleas will leave a dead squirrel and bite a human being. In the records of plague in California there are several cases in which there seems to be very little doubt that the disease resulted from handling plague-infected squirrels.
Fleas being wingless insects travel with considerable difficulty over the ground; and though their hopping powers are notorious they are unable to make any long-continued progress in this way. The methods by which they get dispersed are of interest. Some may be carried by the host in its natural wanderings. Rats appear to be constantly picking up and dropping fleas. Sick rats harbour more fleas than others and therefore more frequently drop them. A hundred fleas have been collected off one plague-sick rat; and, as we know, if this rat was moribund, some of these fleas would most likely be infected. It is obvious that a plague-sick rat may travel about leaving as it wanders a trail of infected fleas behind it. Rats, too, are frequently transported with certain kinds of merchandise and carry their fleas to the most distant parts of the globe, travelling with all the speed and luxury which modern steamships afford. Rats will dive into sacks of grain or bran and hide, so that the bag can be loaded as cargo without anyone suspecting the presence of a rat inside. _M. rattus_ and its fleas, from the habits of the host, are especially likely to be transported in this way. Besides, many fleas are now dispersed without their hosts in merchandise of various kinds. They may travel great distances in these days of rapid transport, though adult fleas, without a host to feed on, generally die in about five days. But larvæ, which eat organic rubbish, and pupæ, which do not eat at all, might arrive alive at the end of a journey of well over a month. On arrival, they would seek their true host, or the next best available animal. Not having yet fed, and being newly emerged, they might survive as long as a fortnight without a suitable host.
Fleas dislike damp breeding places, but dirty carpets, chopped straw, old sacking, paper shavings, and such-like rubbish suit them admirably.
_M. rattus_ is fond of making nests on grain bags and in such sacking the larvæ of fleas are often found. Where trade is carried on in sacks and gunny bags this means of distributing fleas and plague should be kept in mind.
Some rat-fleas, as we know, will feed on man as well as on rats; but their behaviour is rather different when they feed on rats and on man. It has been repeatedly noticed that the fleas were much more readily attracted by the rat than by man. Although the fleas jump on to a man’s hand they take some time to begin to feed. They crawl about and seem to have some doubt where best to begin their sucking operations. Also it has been observed that the fleas much more readily fall off a man’s arm, when he moves, than they do when a rat moves. It seems that they are able to get a firmer hold on the rat than on a man; and it is of interest to note in this connection the larger claws of the human flea compared with the claws of the rat-flea.
It has been shown, in various parts of India, that the number of rat-fleas found on rats varies with the seasons. This seasonal variation of rat-fleas corresponds in a general way with the plague mortality. During the season when plague is bad the average number of fleas per rat is above the mean. During the non-epidemic season it is below the mean. The height of the epidemic corresponds fairly closely with the season of maximum flea prevalence.
Nineteen species of _Pulicidæ_ are more or less habitually obtained by collectors on rats and mice. But the great majority of these may be called casual visitors. Six species of _Sarcopsyllidæ_ are also occasionally found on rats. These are the burrowing chigoes and their allies which usually attack the head and ears of rats.
The species of flea commonly found on rats are five in number, and the readiness with which they bite human beings has been carefully studied.
1. _Xenopsylla cheopis._ This is the oriental rat-flea first described by Mr Charles Rothschild from specimens collected in Egypt. The true home of this flea appears to be the Nile valley, where it may be found in plenty on various hosts. Many of these are desert animals and the flea shows a preference for rodents. Having been distributed all over the world by rats, it now occurs, occasionally, in all warm climates. It is the common rat-flea of the tropical and sub-tropical world. In India it often happens that the whole of the fleas collected from rats prove to be of this species. But it cannot, apparently, flourish in cold countries. In the warmer temperate zones, such as the Mediterranean and Australian seaports, it occurs in varying proportions according to the time of year. The numbers decline with cold weather. It readily bites man and is more active than any other flea in the transmission of plague. For this reason it is sometimes spoken of as “the Plague-flea.” It is a smaller and a lighter coloured insect than the human flea.
2. _Ceratophyllus fasciatus._ This is the common European rat-flea. It is the rat-flea of the temperate as opposed to the hot countries of the world. It is commonly found on black and brown rats in the British Islands and the other countries of Northern and Central Europe. It readily bites man, and there is no reason to suppose that, other conditions being equal, it would not be as efficient an agent in spreading plague as the last species has been shown to be in India.
3. _Ceratophyllus anisus._ This is a closely allied species of rat-flea which replaces the last in China and Japan.
4. _Leptopsylla musculi._ This is the mouse-flea and it is as widely distributed over the globe as its host. From mice it frequently moves to rats, and it has been found on them in various parts of Europe, America, Australia, and Japan. It occasionally bites man, but evinces little inclination to do so.
5. _Ctenophthalmus agyrtes._ This flea is commonly found as a parasite of voles and field-mice. When farm-rats take to an open life in the fields they pick up this species from the rustic rodents. In Hertfordshire, Hampshire and Suffolk one half the fleas from rats, collected in farmyards and hedgerows, were found to belong to this species; but whether it is as common on rats all over England is unknown. It appears not to bite man. A closely allied flea (_Ct. assimilis_) is found in central Europe on field-mice and equally on rats which live under the same conditions. It has not been found in England.
The principal occasional parasites of rats are dog-fleas, cat-fleas, fowl-fleas, and human fleas. The proportions in which they and rat-fleas are found vary greatly in different parts of the world. For instance, in San Francisco nine per cent. of the fleas collected from rats have sometimes been found to be human fleas; whilst in Italy as many as twenty-five per cent. have been identified as cat-and dog-fleas.
It must be borne in mind that when new countries are opened up by man the rats, which follow in his rear, exterminate numbers of the weakly native small mammals and take on their fleas. A change of habitat may be followed by an exchange of fleas.
Some interesting work has been done in testing the appetite of different kinds of flea for human blood. The oriental rat-flea (_X. cheopis_) has been kept alive for three weeks on that diet. Other species show repulsion for mankind and refuse to suck. The experiments confirm the popular belief that fleas have a marked preference for certain individuals. When the flea has refused to bite the human arm, it becomes necessary to check the experiment by trying whether the refusal is merely due to want of hunger. For this purpose a rat must be at hand. It can be secured on a board by two bandages fixed at each end by drawing pins. The rat lies, of course, on its back with its head comfortably supported by a little pillow of cotton wool. A portion of the rat’s abdominal wall is left exposed and shaved. The flea, in an inverted test-tube, can then be put on the hairless patch of the abdomen and given an opportunity of biting, which it may or may not accept.
When fleas are being collected from rats it has been noticed that the true rat-fleas are usually on the hind-quarters of the host, whilst the mouse-flea prefers the region of the head and neck.
As regards the tastes and habits of oriental rat-fleas in the matter of food a long series of experiments may be summarized in this way:
(1) When many rat-fleas are present some will attack man, even when a rat is available for their food-supply. (2) When the number of rat-fleas is small, and when their true host is present, they will not attack man. (3) When rat-fleas are starved they will readily attack all animals, not being particular in the choice of a host. (4) Rat-fleas deprived of their food for from 72 to 96 hours attack and feed on man more readily than at other times. (5) Rat-fleas, even when starved, prefer their true host to man. (6) Rat-fleas may be attracted to man, jump on him, but take some time to feed on him. Plague-infected fleas might in this way be carried from one place to another without infecting the man; but they would, when brought near a rat, attack it in preference to man.
The fleas found on bats possess certain peculiarities which have led to their being grouped together. They form a family to which the name _Ceratopsyllidæ_ has been given. They are recognized by two flaps, one on each side of the head. What these are and what service, if any, they render to their possessors is unknown. Bat-fleas also, as a rule, have maxillæ shaped like dumb-bells; but in one genus (_Thaumopsylla_), found on fruit-bats, they are triangular as in other fleas. The maxillæ as the reader may remember, are parts of the insect’s mouth, and, though placed like jaws on each side of the aperture, they are not used in piercing the skin and sucking blood (Fig. 4). They bear feelers called the maxillary palpi. The flea (_Thaumopsylla breviceps_) which is found on South African fruit-bats and which has triangular maxillæ, seems to be a connecting link between this peculiar group of fleas and the main family _Pulicidæ_.
Bat-fleas are commonly well supplied with combs. They usually have them on the abdomen, as well as the head, and the maximum number of eight combs is found in bat-fleas. Their structure and life-history agree generally with that of other fleas. They breed in hollow trees, caves, ruins, church-towers and lofts where bats hibernate or spend the hours of daylight. The larvæ feed on the droppings of the bats, and the mature insect, after emerging from the pupa case, takes the first opportunity that comes of getting on to its host. Bats are seldom found to be much infested with fleas; for this reason, bat-fleas are somewhat difficult to obtain and many of the species that are known are extremely rare.
The hosts of bat-fleas, obviously, vary more as to the surroundings which they inhabit than almost any other animals. They are found from the equator north and south to the Arctic circle and the straits of Magellan, in the densest tropical forests and flitting round the barest northern buildings. Some pillage the rich fruit gardens of India, whilst other smaller bats work hard for a precarious diet of gnats round a Siberian village. Two sharply divided groups of bats exist: (1) The fruit-bats (Macrochiroptera) with flat molar teeth adapted for a vegetable diet. These are found in the warmer parts of the Old World but not in America. (2) The insectivorous bats (Microchiroptera) whose molar teeth are equipped with sharp cusps for biting their animal food. These have an almost world-wide distribution, and one species at least ranges within the Arctic circle. The same fleas are not as a rule found on the large fruit-bats as on the small ordinary bats. But some bat-fleas have an extensive range. The same species has been taken from different bats of various kinds in Sierra Leone, in Madagascar and in Java.
All bat-fleas are blind. This absence of eyes, in fleas which are parasites of strictly nocturnal animals, lends colour to the suggestion that fleas which are blind have lost their eyes because they had no need of them. Disuse is speedily followed by degeneration.
APPENDIX A
SYSTEMATIC VIEW OF THE ORDER _SIPHONAPTERA_
Order SIPHONAPTERA. Latreille (1825).
Insects with body laterally compressed. Head rounded and fixed by the whole posterior part to the thorax. Mouth-parts for piercing and sucking, consisting of paired mandibles with serrate margins and unpaired labrum. These are sheathed by the labium and labial palpi. Maxillæ usually triangular with four-jointed palpi. Eyes simple, placed in front of the antennæ, occasionally rudimentary or absent. Antennæ of three main segments which lie when at rest in a groove. Three thoracic segments, always free, each consisting of a notum and a sternum. The sterna of the second and third segments are further divided into a sternum, an episternum and an epimeron, the two latter constituting the pleura. Wings and rudiments of wings entirely absent. Abdomen of ten segments of which the sternite of the first segment is suppressed. Abdomen enormously swollen in pregnant females of certain species. Combs frequently present on head, thorax, and abdomen. Legs developed for leaping. Coxa powerful; femur thickened; tarsi of five segments, ending in two claws on the distal segment. Metamorphosis complete. Larva of thirteen segments. Pupa enveloped in silken cocoon. Imago a temporary parasite (usually) on warm-blooded vertebrates.
I. Family _Sarcopsyllidæ_. Taschenberg (1880).
Rostrum (= labium + labial palpi) rather long but very weak and pale, consisting of two or three segments inclusive of the unpaired basal segment. Genal edge of head always produced downwards into a triangular process situated behind the insertion of the maxillæ at the ventral oral angle. Thoracical tergites together shorter than first abdominal tergite.
To this group belong the chigoes and their allies, the most truly parasitic fleas. About fourteen species have been described, which can be grouped into three genera, viz. _Dermatophilus, Hectopsylla, Echidnophaga_.
II. Family _Pulicidæ_. Taschenberg (1880).
Rostrum (= labium + labial palpi) more or less strongly chitinized, consisting, except in a few cases, of five, or more, segments inclusive of the unpaired basal one. Thoracical tergites together longer than first abdominal tergite.
Here belong the majority of _Siphonaptera_.
III. Family _Ceratopsyllidæ_. Baker (1905).
Head on each side with two flaps situated at the front oral corner. Here belong the bat-fleas only. There are several genera, and about twenty-five species have been described. In most of the bat-fleas the maxillæ are shaped like a dumb-bell, but in the genus _Thaumopsylla_ they are triangular as in the _Pulicidæ_.
* * * * *
Oudemans (1909) has put forward an alternative classification of the order _Siphonaptera_ based on the morphology of the head:—I. _Integricipita_, II. _Fracticipita_.
APPENDIX B
A LIST OF BRITISH FLEAS AND THEIR HOSTS
A list of the British Fleas (_Siphonaptera_) revised to March 1913:
Names Usual Hosts
PULEX, _L._ P. irritans Man, Badger
XENOPSYLLA, _Glink_. X. cheopis, _Rothsch._ Rat
ARCHÆOPSYLLA, _Dampf_ A. erinacei, _Bouché_ Hedgehog
CTENOCEPHALUS, _Kolen_ C. canis, _Curt._ Dog C. felis, _Bouché_ Cat
SPILOPSYLLUS, _Baker_ S. cuniculi, _Dale_ Rabbit
ORNITHOPSYLLA, _Rothsch._ O. lætitiæ, _Rothsch._ Puffin and Manx shearwater (Scilly Is. only)
CERATOPHYLLUS, _Kolen_ C. fasciatus, _Bosc._ Rat C. londiniensis, _Rothsch._ House-mouse, Rat C. sciurorum, _Schrk._ Squirrel, Dormouse C. melis, _Wlk._ Badger C. mustelæ, _Wagner_ Bank-vole and Field-mice C. penicilliger, _Grube_ Bank-vole and Field-mice C. walkeri, _Rothsch._ Stoats, Voles, and Field-mice C. gallinæ, _Schrk._ Chickens and many birds C. fringillæ, _Wlk._ Sparrow C. garei, _Rothsch._ Many birds C. rusticus, _Wagner_ House-martin C. farreni, _Rothsch._ House-martin C. hirundinis, _Curt._ House-martin C. columbæ, _Gerv._ Pigeon C. styx, _Rothsch._ Sand-martin C. gallinulæ, _Dale_ Many birds: especially freshwater breeders C. vagabundus, _Bokeman_ Nests of Sea-fowl C. borealis, _Rothsch._ Gannet and Rock-pipit C. rothschildi, _Waterst._ House-martin
CTENOPHTHALMUS, _Kolen_ C. agyrtes, _Heller_ Field-mice and Voles C. agyrtes nobilis, _Rothsch._ Water-rat C. bisoctodentatus, _Kolen_ Mole
DORATOPSYLLA, _Jord. and Rothsch._ D. dasycnemus, _Rothsch._ Shrew
RHADINOPSYLLA, _Jord. and Rothsch._ R. pentacanthus, _Rothsch._ Weasel R. isacanthus, _Rothsch._ Bank-vole
PALÆOPSYLLA, _Wagner_ P. sorecis, _Dale_ Shrew P. minor, _Dale_ Mole P. kohauti, _Dampf_ Mole
LEPTOPSYLLA, _Rothsch._ L. musculi, _Duges_ Mouse L. spectabilis, _Rothsch._ Bank-vole
TYPHLOCERAS, _Wagner_ T. poppei, _Wagner_ Long-tailed Field-mouse
HYSTRICHOPSYLLA, _Taschb._ H. talpæ, _Curt._ Mole
ISCHNOPSYLLA, _Westw._ I. elongatus, _Curt._ Noctule Bat I. intermedius, _Rothsch._ Serotine Bat I. simplex, _Rothsch._ Natterer’s Bat I. octactenus, _Kolen_ Pipistrelle Bat I. hexactenus, _Kolen_ Long-eared Bat
NYCTERIDOPSYLLA, _Oudemans_ N. longiceps, _Rothsch._ Pipistrelle Bat N. eusarca major, _Rothsch._ Noctule Bat
APPENDIX C
ON COLLECTING AND PRESERVING FLEAS
There are two methods by which fleas may be preserved for study when they have been collected. The first is by keeping the specimens in small tubes of alcohol; the second is by mounting each in Canada balsam on a slide for the microscope. The advantage of the former method is that the material can be used for dissection. The student can do nothing without a microscope, though some of the commoner species can be identified with tolerable certainty by a practised eye which is assisted by a pocket-lens.
The tubes are best stored away in a cabinet fitted with wooden shelves and holes to take the tubes like a test-tube holder. Fleas dried and preserved loose in a box, or gummed on card, are useless for purposes of minute examination, and are soon destroyed.
Fleas may be collected from the great majority of mammals and birds in almost all parts of the globe. They can be found in the hair and under the feathers, and also in the places where the animals habitually sleep. The best places, from which a plentiful haul may often be obtained, are the holes and nests in which the young have been reared.
It is essential to remember, when an animal has been killed, that all the fleas leave as soon as the body of the host gets cold. No time, therefore, should be lost in searching for specimens.
If the animal is small enough it may be put into a cardboard box, or a white linen bag, and a few drops of chloroform or benzine can be poured on it. In a short time the fleas will be found dead in the bag or at the bottom of the box. Some may also be found in the hairs and feathers when they are turned back.
In the case of a large mammal the hair must be turned backwards shortly after death, when the live fleas may be seen running about and caught. For this purpose a small camel’s hair brush is very useful. If a flea is touched with a brush of this kind which has been dipped in chloroform, benzine, or alcohol, the insect sticks to the brush, but can be easily floated off into the tube of preservative. The best preserving liquid is 50 per cent. alcohol. Methylated spirits can be used. Acetic acid can also be used; but it is objectionable because in a short time it destroys the corks of the tubes.
Each tube should only contain the fleas collected from one host, but as many specimens as possible should be secured, because there may be several species of flea on the same host.
The tube must be securely corked and labelled, with the date, the locality, and the name of the host. In foreign countries it may not be always easy to do this. In such cases the skin of the host must be preserved with the tube for subsequent identification. A number corresponding to the label on the tube should be attached to the skin of the host.
Fleas collected without records of the host from which they were obtained are of little or no scientific value. For this reason a tube should contain the parasites of one host only.
A convenient way of preserving records temporarily is to write in _pencil_ on a small piece of paper which can be rolled up and put in the alcohol in the tube.
Small mammals generally, including bats, are good hosts; Rodents and Insectivora afford usually the most fruitful captures. In trapping mice and voles only those traps should be used in which the animals are caught alive, or the fleas will have left their hosts before they can be secured and examined. Field-mice caught in the ordinary small penny mouse-trap are often found dead in the morning. The best traps are made on the principle of the ordinary mouse-trap, but larger. A piece of bacon-rind on the hook is a good bait for almost all small mammals. Where a number of traps are put down and left out they should, of course, be visited daily.
When a live mouse, or other small mammal of similar size, has been captured it may be transferred from the trap into a small white linen or holland bag. The animal can then be killed by tapping its head or breaking its neck from the outside of the bag. After this has been done, fleas may be searched for in the fashion described above.
Most of the small mammals which act as hosts for fleas are nocturnal. The localities where they may be trapped are numberless, but only a small proportion of the captures may yield anything for the flea-collector. I have heard of a collector of small mammals who travelled through remote parts of Spain and never lost an occasion for putting down his traps when he had to change trains at a country railway junction. In England it would, however, seldom be worth doing this, as, on many lines, there is an attempt to make the arrival of one train and the departure of another correspond.
The following plan for securing bird-fleas will be found successful. The nests of birds should be taken as soon as the young are fledged and flown. If the nest is small it can be put into a glass-topped box lined with white paper. If too large, the whole nest, or the most firmly matted and dirty part, may be put into a glass globe (such as gold-fish are kept in) and a piece of paper tied over the mouth. From time to time the nest should be slightly damped with water. In every case a label should be put into the receptacle to preserve the name of the bird which built the nest. The bottom of the nest may sometimes be seen to be full of the larval fleas; but in any case fleas will probably emerge from pupæ. The fleas will continue appearing for as much as six weeks or eight weeks after the young birds have left the nest. They must be watched for and taken off the sides and top of the box with a camel’s hair brush dipped in chloroform or benzine. I have seen dozens of fleas come from the nest of a tit (_Parus major_) in the course of a few weeks. They were all of the common species (_C. gallinæ_). When the lid of the box is removed it is difficult to prevent a few escaping in the room, but I have never known them cause inconvenience to anyone.
Collectors in warm countries should give their attention to the chigoes and their allies, which are of great interest and have been little studied. They are found on mammals and birds in tropical and semi-tropical countries. The males are very difficult to find, but the females are large and very parasitic. They have the appearance of a small wart firmly fixed to the skin. Small mammals may be transferred, with their chigoes attached, to a bottle of alcohol. Many examples of these insects are often found together on the more naked portions of their hosts.
APPENDIX D
BIBLIOGRAPHY
A. General Bibliography of the chief and most recent works. Many have bibliographies of earlier works.
1. TASCHENBERG (1880), _Die Flöhe_. Halle.
Although the author only describes 30 species, his book forms the basis of all subsequent scientific work. He divides fleas into (_a_) chigoes, (_b_) non-chigoes. Plates. Bibliography. References to 73 earlier writers.
2. KARSTEN (1864), _Beitrag zur Kenntniss des Rhyncoprion penetrans._ Bull. Soc. Imp., Moscou. Vol. XXXVII., p. 72.
A full account of the life and habits of the chigoe. Many references to older writers and travellers.
3. WAGNER (1889), _Aphanipterologische Studien._ Hor. Soc. Ent. Ross. Vol. XXIII., p. 199; (1893) Vol. XXVII., p. 347; (1898) Vol. XXXI., p. 539; (1902) Vol. XXXV., p. 17; (1903) Vol. XXXVI., p. 125.
A series of learned papers by a professor at the Russian University of Kieff. He has also written in Russian.
4. DAMPF (1907), _Die Ost-und Westpreussische Flohfauna._ Schriften der Physik.-ökonom. Gesellschaft zu Königsberg-i-Pr., XLVIII. Jahrgang, p. 388.
Contains an excellent general account of our present knowledge of fleas. Short Bibliography.
5. DAMPF (1910), _Palæaeopsylla Klebsiana n. sp. ein fossiler Floh aus dem baltischen Bernstein_. Schriften der Physik.-ökonom. Gesellschaft zu Königsberg-i-Pr., LI. Jahrgang, II. 248.
Good plates of the fossil flea in amber.
6. OUDEMANS (1909), _Neue Ansichten über die Morphologie des Flohkopfes sowie über die Ontogenie, Phylogenie und Systematik der Flöhe_. Novitates Zoologicæ, Vol. 16, p. 133.
Suggests a new classification based on the morphology of the head, viz. (1) Integricipita; (2) Fracticipita. By the chief authority on fleas in the Netherlands. He has also written numerous papers in Dutch.
7. JORDAN and ROTHSCHILD (1908), _Revision of the non-combed eyed Siphonaptera_. Parasitology, Vol. I., p. 1. Plates. Bibliography.
An excellent piece of work, which includes an account of the plague fleas.
8. JORDAN and ROTHSCHILD (1906), _A Revision of the Sarcopsyllidæ_. Thompson Yates and Johnston Laboratories Reports, Vol. VII., p. 15. Plates. Bibliography.
The best modern account of the chigoes.
9. BAKER (1904), _A revision of American Siphonaptera or Fleas together with a complete list and bibliography of the group_. Proc. U.S. National Museum, Vol. XXVII., p. 365. Plates. Bibliography of special papers only.
The earlier references, beginning 1699, mostly from Taschenberg.
10. BAKER (1905), _The Classification of the American Siphonaptera_. Proc. U.S. National Museum, Vol. XXIX., p. 121.
Gives an index of hosts and their fleas. Additional bibliography.
11. ROTHSCHILD (1898), _Contributions to the knowledge of the Siphonaptera_. Novitates Zoologicæ, Vol. 5, p. 533; (1900) FURTHER CONTRIBUTIONS, _etc._, Vol. 7, p. 539; (1903) Vol. 10, p. 317; (1904) Vol. 11, p. 602; (1905) Vol. 12, pp. 153 and 479; (1907) Vol. 14, p. 329; (1909), Vol. 16, pp. 53, 57, 61, and 332; (1906) _Notes on Bat-fleas_, Vol. 13, p. 186.
These papers contain, for the most part, descriptions of new species in the writer’s collection. Many fine plates illustrating morphology and structure of the external skeleton.
B. The following references are to papers on the systematic position of the _Siphonaptera_ and their relationship to other insects.
1. KRAEPELIN (1884), _Ueber die systematische Stellung der Puliciden_. Hamburg, 1884.
2. PACKARD (1894), _The Systematic Position of the Siphonaptera_. Proc. Boston Nat. Hist. Society, Vol. XXVI., p. 312.
3. DAHL (1897), _Puliciphora, eine neue flohänliche Fliegengattung_. Zoologischer Anzeiger, Vol. XX., p. 409.
4. WANDALLECK (1898), _Ist die Phylogenese der Aphaniptera entdeckt?_ Zoologischer Anzeiger, Vol. XXI., p. 180.
A humorous reply to Dahl.
5. DAHL (1899), _Die Stellung der Puliciden im System_. Archiv für Naturgeschichte, Vol. 65, I. p. 71. Plates.
6. HEYMONS (1899), Die systematische Stellung der Puliciden. Zoologischer Anzeiger, VOL. XXII., pp. 223 and 301. Three figures.
A destructive criticism of the views advanced by Dahl.
7. SEMENOV (1904), _Zur Frage der systematischen Stellung der Flöhe_. Revue Russe d’Entomologie, Vol. IV., p. 277. In Russian.
C. The following references are chiefly to works on plague and fleas.
1. ADVISORY COMMITTEE (1905-1909), _Reports on Plague Investigations in India_. Journal of Hygiene, Vols. V., VI., VII., VIII. and X.
These volumes contain the five “Extra Plague Numbers.” Many references to observations and experiments on rats and fleas.
2. HANKIN (1905), _Plague Epidemiology_. Journal of Hygiene, Vol. V., p. 48.
3. OGATA (1897), _Ueber die Pestepidemie in Formosa_. Centralbl. für Bacteriol., Vol. XXI., p. 769.
4. SIMOND (1898), _La Propagation de la Peste_. Annales de l’Institut Pasteur. Vol. XII., p. 625.
5. HANKIN (1898), _La Propagation de la Peste_. Annales de l’Institut Pasteur, Vol. XII., p. 705.
6. VERJBITSKI (1908), _The part played by insects in the epidemiology of the plague_. Journal of Hygiene, Vol. VIII., p. 162.
Experiments made 1902-3 at Cronstadt and S. Petersburg with fleas. This important research was written in Russian and not translated or published till 1908.
7. DOANE (1910), _Insects and Disease_. Constable and Co.
A popular work by an American entomologist. Contains a chapter on fleas and plague. Some good micro-photographs of fleas. Bibliography.
INDEX
Abdomen, 31
Ancestors of fleas, 3
Antennæ, 25, 47
Aphaniptera, 4
Aptera, 34
Arachnids, 1
Arctomys, 95
Armadilloes, 14
Arthropods, 1, 2
Bacillus pestis, 84
Badger, 9, 68
Baker, Mr Carl, 70
Bat-fleas, 24, 106
Bat-fleas, head-flaps of, 27
Bats, 12, 107
Bettongia, 15
Bibliography, 118
Black death, 86
Blood-system, 60
Blue, Doctor, 98
Boden, Mr, 18
Bombay, Plague in, 95
Brain, 45
British fleas, list of, 110
British fleas, number of, 11
Brown snake, 15
California, plague in, 98
Carnivora, 14, 16
Caterpillars, attacked by fleas, 18
Catesby, 75
Cat-flea and dog-flea distinct species, 70
Catholic and Protestant, 9
Caudal stylets, 37
Ceratopsyllidæ, 17, 106, 110
Ceratophyllus anisus, 103
C. farreni, 11
C. fasciatus, 89, 96, 102
C. gallinæ, 70
C. gallinulæ, 70
C. rothschildi, 11
C. vagabundus, 11
Chigoes, 74
—— on bats, 75
—— burrowing habits, 78
—— distribution of, 83
—— on parrot, 75
—— pregnant females of, 81
—— post-oral process of, 27
—— on rats, 101
—— rostrum of, 77
Chitin, 21
Classification of fleas, 17
Claws of feet, 31
Coleoptera, 34
Collection of fleas, 113
Combs, 26
Corsica, fleas attack flies in, 19
Ctenocephalus canis, 8, 9, 16, 63, 70
C. erinacei, 63
Ctenophthalmus agyrtes, 103
C. assimilis, 103
Dahl, 35
Dampf, 18
Dermatophilus cæcata, 82
D. penetrans, 69, 74, 83
Diemenia superciliosa, 15
Diptera, 4, 5
Dissection, 61
Distribution, 11, 12
Dog-flea, development of, 8
Dog-flea and cat-flea distinct species, 70
Dolichopsyllus stylosus, 22
Echidna, 15
Echidnophaga ambulans, 15
Edentata, 14
Eggs, 5
Epidermis, 21
Epimeron, 31
Erinaceus, 68
Excreta, 57
Eyes, 24
Field-mouse, 9, 25
Fossil fleas, 18, 19
Fowl-flea, 10
Fracticipita, 110
Frontal tubercle, 50
Fruit-Bats, 107
Ganglia, 45
Gannet, 11
Genitalia, 51
Gerbillus, 68
Gizzard, 54
Gophers, 95
Ground-squirrels, 95, 98
Guinea-pigs, 66, 93
Gullet, 54
Hankin, 88
Hatching spine, 5
Head, 28
Hearing, 50
Heart, 60
Hedgehog, 9
Hemiptera, 34
Heymons, 31
Hooke, Robert, 47
Hosts, of British fleas, 110
—— change of, 10
—— meaning of, 8
House-martin, 10
Human flea, absent from Sahara, 64
Human flea, from badgers, 68
Human flea, described, 63
Human flea, from Mexican Indians, 67
Human flea, mouth-parts, 67
Human flea, from New Guinea, 65
Hypopharynx, 39
Hystrichopsylla talpæ, 22
Insectivora, 13
Integricipita, 110
Ischnopsylla unipectinata, 13
Jehangir, Emperor, 86
Justinian, Emperor, 86
Kitasato, 84
Klebs, Professor, 19
Labial palpi, 48
Labium, 40
Labrum, 42
Larva, 5, 7, 36
Leeuwenhoek, 38
Legs, structure of, 30
Leptopsylla musculi, 89, 103
Linnæus, 4, 75
Listropsylla, 50
Macrochiroptera, 107
Malacopsylla, 80
M. androcli, 14
M. grossiventris, 14
Malpighi, 59
Malpighian tubes, 59
Manchuria, plague in, 95
Mandibles, 42, 48
Marmots, 95
Marsupials, 14
Maxillæ, 40, 48
Maxillary palpi, 40
Metamorphosis, 5
Microchiroptera, 107
Monkeys, 12
Monotremes, 15
Mouth-parts, 39
Mouth, 54
Muscles, 33
Mus, genus, 97
Myriapods, 1
Myrmecobius, 15
Needles for dissecting, 61
Nervous system, 44
Nycteribia, 26
Ogata, 87
Orthoptera, 34
Otospermophilus beecheyi, 98
Oudemans, his classification, 110
Ovaries, 51
Oviedo, 74, 76
Palæopsylla, 19
Pariodontis riggenbachi, 69
Penguins, 11
Performing fleas, 72
Petrels, 11
Pharynx, 54
Pharyngeal pump, 54
Phoridæ, 35
Pigeon-flea, 10
Plague, 84
Plague and fleas, 87
—— in Egypt, 86
—— of London, 85
—— medieval precautions, 21
—— transmission of, 44
Platypsyllus, 26
Polyctenes, 26
Protestant and Catholic, 9
Puffins, 11
Pulex irritans, 4, 19, 63, 75, 95
Pulicidæ, 17, 109
Pupa, 7
Pygidium, 46, 50
Rat-fleas, 104, 105
—— and man, 101
Rats and mice, fleas of, 97
Rectal glands, 57
Rectum, 57
Rectum, discharge of blood from, 44
Reptile, flea on a, 15
Rock-pipit, 11
Rodents, 13
Rostrum, 40
Rothschild, Mr Charles, 70, 101
Salivary glands, 58, 92
Samuel, Book of, 85
Sand-martin, 10, 11
San Francisco, plague in, 94
Sarcopsyllidæ, 17, 74, 109
Seals, 14
Segmented structure, 22
Sense organs, 49
Sexual differences, 23, 51
Sexual organs, 51
Simond, 87
Siphonaptera, description of Order, 108
Size of fleas, 22
Species, number of, 4
Spilopsyllus cuniculi, 77
Spines 23, 25
Stephanocircus, 27
Sternites, 32
Stigmata, 59
Stomach, 56
—— bacilli in, 93
St Paul’s Cathedral, 29
Sucking, method of, 43
Suctoria, 35
Swift, African, 11
Taste, sense of, 50
Tergites, 32
Testes, 51
Thaumopsylla breviceps, 106
Thorax, 29
Tiraboschi, Dr Carlo, 70
Tracheæ, 59
Trapping hosts of fleas, 113
Trichosurus, 15
Typhloceras poppei, 25
Ungulates, 13
Urinary tubules, 59
Variation, geographical, 16
Verjbitski, 88
Vermipsylla, on Ungulates, 13
Wings, relics of, 4, 31
Woodward, Dr, 15
Xenopsylla cheopis, 11, 89, 102, 104
Yersin, 84
Zoological Gardens, fleas in, 16, 66
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