THE FLEA

TRANSCRIBER’S NOTES:—Obvious print and punctuation errors were corrected.—The decorated front fage has been retained as illustration.The Cambridge Manuals of Science and LiteratureTHE FLEACAMBRIDGE UNIVERSITY PRESSLondon: FETTER LANE, E.C.C. F. CLAY,ManagerEdinburgh: 100, PRINCES STREETLondon: H. K. LEWIS, 136, GOWER STREET, W.C.WILLIAM WESLEY & SON, 28, ESSEX STREET, STRANDBerlin: A. ASHER AND CO.Leipzig: F. A. BROCKHAUSNew York: G. P. PUTNAM’S SONSBombay and Calcutta: MACMILLAN AND CO., LTD.All rights reservedAfter a drawing by Dr JordanOriental rat-flea (Xenopsylla cheopisRothsch.). Male.CambridgePRINTED BY JOHN CLAY, M.A.AT THE UNIVERSITY PRESSWith the exception of the coat of arms at the foot, the design on the title page is a reproduction of one used by the earliest known Cambridge printer, John Siberch, 1521cpPREFACETHE aim of this book is to give in plain language some account of a small, but noteworthy, group of insects. I have avoided, whenever I could, using the technical terms of zoology. To avoid doing so entirely is impossible in a book which describes insects in some detail. No technical term has, I hope, been used without an explanation.Over thirty years have elapsed since Taschenberg’s German book,Die Flöhe, appeared. Our knowledge has made enormous strides since then. More species of flea are now known from the British Islands alone than were then known from the whole world. So far as I am aware, no book, devoted to what is known about fleas, has ever been published in English. The statements about these insects in the general text-books of entomology are frequently antiquated and inaccurate. But there is a fairly extensive literature on theSiphonapterascattered through scientific periodicals mostly in English, German, Italian, Dutch and Russian. I have given some references in the Bibliography.The naturalists now living who have devoted any time to the special study of fleas may almost be counted on one’s fingers. In England there are Mr Charles Rothschild and Dr Jordan; in the Shetland Islands, the Rev. James Waterston; in Germany, Taschenberg of Halle and Dampf of Königsberg; in Russia, Wagner of Kieff; in Holland, Oudemans of Arnhem; in Italy, Tiraboschi of Rome; in the United States, Carl Baker and a few others. I have not mentioned medical men who have investigated fleas in connection with plague.There are small collections of fleas in the Natural History Museums at South Kensington (London), Paris, Berlin, Königsberg, Vienna, Budapest, S. Petersburg and Washington. Of private collections Mr Charles Rothschild’s at Tring is by far the best in the world. It contains something like a hundred thousand specimens and is most admirably kept. I must express profound and sincere gratitude to Mr Rothschild for having helped me in numberless ways and advised me in many difficulties.It is well known that the mere mention of fleas is not only considered a subject for merriment, but in some people produces, by subjective suggestion, violent irritation of the skin. The scientific study of fleas has, however, received a great impetus since it has been ascertained that they are the active agentsin spreading plague. Rat-fleas are of various kinds, and not all fleas will bite man. A knowledge of the different species has suddenly become useful. The humble, but ridiculous, systematist with his glass tubes of alcohol for collecting fleas, his microscopic distinctions, and Latin nomenclature has become a benefactor of humanity. Some people seem to be practically immune to the bites of fleas, but even to such persons their visits are unwelcome. A famous Frenchwoman once declared: “Quant à moi ce n’est pas la morsure, c’est la promenade.”H. R.London,September, 1913.CONTENTSCHAP.PAGEPrefacevI.Introductory1II.The external structure of a flea21III.The mouth-parts and sense-organs38IV.The internal organs of a flea52V.The Human flea and other species62VI.The Chigoes and their allies74VII.Fleas and Plague83VIII.Rat-fleas and Bat-fleas97AppendixA.Systematic view of the orderSiphonaptera108”B.A list of British fleas and their hosts110”C.On collecting and preserving fleas113”D.Bibliography118Index122LIST OF ILLUSTRATIONSMale Oriental rat–fleafrontispieceFIGUREPAGE1.The larva of a flea62.Types of genal and thoracic combs of a flea263.The hind leg of a flea304.The mouth-parts of a flea435.The antenna of a flea476.The alimentary canal of a flea537.The head of a female dog-flea and a female cat-flea718.Pregnant female ofDermatophilus cæcata81c1CHAPTER IINTRODUCTORYFleasform a group of insects that have, until recently, been little studied by zoologists. We call them insects because they are jointed animals, or Arthropods, with three pairs of legs in the adult condition. The reader will best understand the position which fleas occupy in the general classification of animals by remembering that the arthropods, or jointed animals, are one of a dozen subkingdoms, or phyla, to which the various members of the great animal kingdom have been assigned. There is good ground for believing that all the animals included in each phylum trace their ancestry back to a common primitive form which lived in more or less remote ages. Besides (1)Insects, the arthropods, or jointed animals, include (2)Crustaceans, such as crabs, lobsters, shrimps, wood-lice, water-fleas and barnacles; (3)Myriapods, such as centipedes and millipedes; and (4)Arachnids, such as spiders, scorpions, mites and ticks. To all these varied forms of animal life fleas, and other insects, are therefore more or less nearly related.The animals belonging to this large and important collection, which compose the arthropod phylum, have certain common characteristic features. We find a body made up of a series of more or less completely similar segments placed one behind the other. In this they resemble certain worms which are far less highly organised. The body is elongated, symmetrical on either side, and the mouth and anus are at opposite ends. There is, however, an important advance on the segmented worms. Each typical segment carries a pair of appendages which are very different from the foot-stumps that are found on certain worms. These appendages of arthropods are divisible into distinct limb-segments, separated from one another by moveable joints, and acted upon by special muscles.The common ancestor of all the various arthropods which are found living on the earth to-day, was probably composed of a series of segments each very similar to the last and each bearing a pair of very similar appendages. In the course of ages, these appendages have been astoundingly modified in form and in function. So it happens that we find in the arthropods of the present day pairs of antennæ, of mandibles and other mouth-parts, of pincers, of legs, of swimming-feet and of tail pieces which on close examination can all be traced back to a common structure. The body-segments, also, have beenstrangely fused together and modified. All that has been so far said applies equally to fleas and to other insects.It is of great interest, when one comes to make a minute study of the form and external structure of a flea, to try and trace the modifications that must have taken place in the course of descent from the ancestral arthropod; but the relationship of fleas to other insects living at the present day is of more immediate concern. Insects are highly specialized arthropods and fleas are highly specialized insects. This means that they have become vastly modified from the primitive ancestral type and fitted thereby for a life among certain defined and peculiar surroundings.It will be unnecessary to remind the reader who knows anything of zoology or of botany that all classification is now based on descent. Since naturalists have abandoned a belief in the special creation of the various species of animals now living on the earth and have conclusively shown that they have arisen by descent and modification from other forms, the problem is to reconstruct a vast genealogical tree. What then were the ancestors of the fleas and to what other insects, in consequence, do they appear to be related?It is probable that the ancestors of the fleas were winged insects, and that the organs of flight weregradually lost, as they became useless, when a partially parasitic life was adopted. At one time entomologists regarded fleas as wingless flies and placed them in the order Diptera. Certain supposed scaly plates on their bodies were regarded as the atrophied relics of wings. It is, however, more than doubtful whether this view is correct; and all modern entomologists who have given any special study to fleas are agreed that they are sufficiently unlike any other living insects to deserve a place in an order by themselves. To this order the nameSiphonapterahas been given: which means that the insects comprised in it are provided with sucking mouths and are destitute of wings. Another name for the order is Aphaniptera, but this is gradually falling into disuse. Linnæus (1758) only mentions two species of flea: the human flea which he appropriately namedPulex irritans, and the chigoe of hot countries which he calledPulex penetrans, from the habit which the female has of burrowing under the skin of her victims. At the time of writing, about 460 species of flea have been described and named; but some of the names are doubtless synonymous, and the actual number of separable species that have been discovered is somewhere about four hundred. The vast majority of these have been described within the last few years, which shows what can be done when attention is turned to any neglected group of animals. Therecan be no doubt that many undiscovered species still remain, and will now, in due course, be collected, described and named.The position which should be assigned to the order Siphonaptera in the general scheme of insect classification is a question on which the most learned modern entomologists have disputed with considerable vigour. Some see the nearest relatives among the beetles, others among the flies. The majority, as we shall see later on, would place them near the Diptera: but since no convincing arguments have been produced on either side it may be wisest to regard the question as still at present unsolved.Fleas belong to one of the groups of insects which go through a complete metamorphosis. Their life-history consequently falls into four divisions: egg, larva, pupa and imago. If the climate permits, the female flea lays her eggs all the year round, and from one to five are dropped at a time. Unlike those of many other parasites they are never attached to the hairs of the hosts, but appear to be deposited indiscriminately on the floors of houses or in the nests and sleeping places of their hosts. The eggs generally hatch in a few days, and a minute, white, wormlike larva emerges (Fig. 1). The larvæ, of some, and possibly of all, fleas are provided with a wonderful adaptation in the shape of an egg-breaker or hatching-spine. This is a thin plate, like the edge of a knife, where the point of the head comes in contact with the shell. The movements of the prisoner make a slight split in the egg-shell, which then bursts asunder. This organ has vanished in later larval life, and it is probably lost after the first moult. The larva is legless and has thirteen segments. It grows rapidly, and, as it grows, moults its skin several times. It is provided with mouth-parts adapted for biting, and eats any decaying organic refuse. The larvæ may be reared on the sweepings of an ordinary room or the dirty scurf which collects at the bottom of old birds’ nests. It is hardly necessary to add that the mother takes no interest whatever in the larvæ and that the belief that she feeds them on dried blood is not based on any sound foundations.i6Fig. 1. The larva of a flea. The body consists of thirteen segments and is legless. On the fore part of the head are the antennæ and on the upper part of the head is shown the knife-like edge of the egg-breaker. The mouth-parts are adapted for biting. On the last segment of the body are the two caudal stylets.The larval stage lasts some days, and the animal spins a small cocoon before pupating. In the course of a few more days, the time probably depending on the weather, the perfect flea emerges. The larvæ generally live in places where the perfect insects will have an opportunity of finding a host as soon as they leave the pupal envelope. The nests of their hosts where the young are being reared are always favourite places. It seems possible that the comparative immunity from fleas which hoofed mammals or Ungulates enjoy may be due to the fact that the young beast follows its mother from the time of birth instead of passing its early life helpless in a nest.Observations made on the development of the dog-flea (Ctenocephalus canis) in India show that eggs laid on October 17 hatched on October 19. The larva spun its cocoon on October 25 and the mature flea emerged on November 2. In Northern Europe the human flea takes about four weeks in summer and six weeks in winter to pass through its metamorphosis.Unlike many parasitic insects, fleas do not constantly pass their time upon the bodies of their victims. The greater part of their life is probably spent on the ground, in the house, or nest, of the mammal or bird which serves them with blood. In this respect there is considerable difference in the habits of different species of flea. Some attach themselves to an animal and actually burrow into the skin. These are the most parasitic species. Some only come to feed and leave to lay their eggs. Many probably do not suck blood more than once in their lives.An animal which harbours fleas and which nourishes the adult insect with blood is called ahost. No fleas are more than what is called temporary parasites; which means that they pass but a portion of their lives on their hosts and frequently take occasion to hop on and off. All fleas, apparently, go from host to host. The labours of diligent collectors have proved that the greatmajority of mammals and birds have fleas. As a general rule, it is true to say that certain species of flea are associated with certain species of host. Thus man is the true host ofPulex irritans; the cat family are the true hosts of the cat-flea (Ctenocephalus felis); and the dog family are the true hosts of the dog-flea (Ctenocephalus canis). But the human flea is sometimes found on cats and dogs, and cat and dog-fleas occasionally bite human beings; and cat-fleas are found on dogs and dog-fleas are found on cats. All fleas, so far as we know, may occasionally pass from one species of host to another; but they do not, for the most part, seem to flourish in unaccustomed quarters. Some fleas are more catholic in their tastes than others. Some seem to be very strictly confined to one host, and even when starving only suck strange blood under protest. There is a species of flea that has only (except by accident) been found on the long-tailed field-mouse and another that has only been found on the hedgehog. Other fleas are commonly found on two absolutely distinct animals; a good instance of this is the human flea which, at all events in certain parts of England, is a regular parasite of the badger.As distinguished from true or natural hosts one must separate what may be termed casual or accidental hosts. All animals which come in contact with one another, or which live in close proximity,may exchange fleas. So even bird-fleas may be collected from mammals and typically mammalian fleas from birds. In this fashion puzzles may arise which tax the ingenuity of the collector to solve. Bird-fleas are sometimes found on bats, and this may be obviously attributed to the bats having inhabited a hole which was tenanted by starlings or an old loft infested with the fleas of pigeons. All beasts of prey are sometimes found to harbour the fleas of animals they have devoured. Rabbits’ fleas are found on wild-cats; hedgehogs’ fleas on foxes; mice fleas on weasels; and fleas characteristic of small birds on stoats. So also in the case of mice, rats and voles with holes and runs in the same hedgerow, the parasites usually peculiar to one are not uncommonly found on the others. It is sometimes difficult to determine the true host of a flea.Much more puzzling to explain are the reasons which confine a flea to a certain host and which cause closely allied hosts to have different fleas. The fleas from the house-martin and the sand-martin are quite different; those from the domestic fowl and the domestic pigeon are distinct species. The causes which have affected the evolution of the various forms of flea are too obscure to enable anyone at the present day to offer any satisfactory explanation.Speaking generally, the fleas found on birds have points in common, and they probably form a naturalgroup to themselves. What may be called true bird-fleas have been collected from almost all European birds. An unwieldy genus (Ceratophyllus) comprises many species of different flea. Some species are very abundant and infest the nests of many different birds. Others are extremely rare. One of these rarities (C. vagabundus) is found in the nests of puffins and other sea-birds. Another has been collected on antarctic petrels. Penguins have a special genus of flea to themselves. A specimen, unique at one time (Ceratophyllus borealis), in Mr N. C. Rothschild’s collection was obtained from the gannet. It has now been found on rock-pipits in the Shetland Islands.Two very rare fleas (C. farreniand C.rothschildi) are found in the nests of house-martins; yet the nests of these birds are infested with common species besides. A plague flea (Xenopsylla) has been found on an African swift.Forty-six different species of flea have been found in the British Islands, but many of these are extremely scarce.We know too little about the geographical distribution of fleas to lay down many accurate generalities. When a great deal more material has been collected and studied, it may be possible to show that certain groups are associated with certain regions of the earth or certain orders of animals. To some extent this is already seen to be the case. The fleasindigenous to the New World are distantly related to those of the Old World. Broadly speaking the geographical distribution of the parasite must follow that of the host. But sometimes the parasite is impatient of cold and cannot follow the host out of the tropics. The chigoes and their allies are fleas of hot countries. Different kinds of bats are found from the tropics to the Arctic circle, but the same bat-fleas are not found everywhere.When a flea has a cosmopolitan range it is probable that it has travelled over the world in company with its host.Monkeys have no fleas. This is an assertion that is commonly received with surprise and incredulity. Occasionally a gorilla or a chimpanzee may get a chigoe in its toe. And monkeys in zoological gardens or menageries are possibly exposed to the danger of catching an occasional human flea from the people who crowd round their cages. These are remote contingencies which may happen to anyone. Healthy wild monkeys are much too clean and active to harbour fleas. When they search one another’s fur in a fashion that must be familiar to most persons, they are clearing their coats of particles of scurf or of similar scraps of dirt and not of fleas. So, speaking generally, it may be said that no fleas have been found truly parasitic on monkeys.Bats have fleas, but not in great abundance. Allbat-fleas are rare on their hosts and extremely difficult to find and collect. The same species are not found on fruit-bats and the ordinary smaller insect-eating bats. The geographical distribution of some bat-fleas is puzzling. For instance, one species (Ischnopsylla unipectinata) is found on the greater horse-shoe bat in Europe; but it is, apparently, not found on the same bat in the British Islands. In Somaliland and in India it is found on other bats.With certain exceptions, Ungulates are remarkably free from fleas. This great order of mammals includes a variety of hoofed animals: oxen, sheep, goats, deer, pigs, camels, giraffes and antelopes.The only true fleas found on these are two species of the genusVermipsylla, which resemble the chigoes in so far that the pregnant females burrow into the host and expand there. One species has been found on camels and horses in Transcaucasia; another on roe-deer in Northern China. The female of the last is often found ensconced on the inside of the nostrils of the deer. Of course chigoes may attack domestic Ungulates of all kinds; but no other members of the familyPulicidæor typical fleas except those two above mentioned have been found on hoofed mammals.Insectivora such as moles, shrews and hedgehogs are the hosts of a great variety of species. The same thing may be said of the Rodents, which includeporcupines, squirrels, rats, mice and a vast number of other small mammals whose geographical distribution includes almost the whole of the habitable globe. Probably more different species of fleas have been collected from Insectivora and Rodents than from all the other orders of mammals grouped together.The Carnivora, excluding the Pinnepedia, or seals, sea-lions and walruses, harbour numerous species.Among the Edentata a very remarkable and highly specialised genus of fleas is parasitic on armadilloes in South America. This genus (Malacopsylla) consists of two species only, which are confined to South America and are found on the armadilloes and on carnivorous animals which probably have preyed on them. The thorax of these fleas is much reduced and very small in size. Their piercing organs are slender and weak, but they possess enormous spines on the legs with which they hold on to their hosts. These two South American fleas (M. grossiventrisandM. androcli) will be referred to again later as striking examples of fleas with strongly developed legs and weakly constructed mouth-parts. The contrary combination of powerful mouths and degenerate legs is also found in other groups of fleas, as will be seen in the chapter on the chigoes.The Marsupials of Australia and South America have special fleas which were probably associated with this strange order of pouched mammals beforethey became divided into the American and Australian groups. Fleas have been collected on the spiny ant-eater (Echidna) which belongs to the lowest order of Monotremes or egg-laying mammals.On almost every form of bird, including the most aquatic kinds, fleas of various species have been obtained.Only one instance has been recorded of a flea occurring on a reptile. A female of one of the species of burrowing chigoes (Echidnophaga ambulans) from Australia was collected by Dr Woodward from the Brown Snake (Diemenia superciliosa). This reptile, which is well known in Australia, belongs to a sub-family that contains some of the most deadly poisonous snakes and is allied to the cobras. The Brown Snake is a terrestrial snake, and one must regard the presence of the flea on such a host as a rare and chance occurrence. The snake was captured at Herdman’s Lake, near Perth in West Australia. The same species of flea has also been obtained from the phalangers (Trichosurus) which live in the tops of the Australian gum-trees; from the little terrestrial and nocturnal rat kangaroos (Bettongia); and from the banded ant-eater (Myrmecobius), another Australian Marsupial. It is possible that the flea moved from some small mammal which was being devoured by the snake and managed to fix itself between the scaly plates of the reptile.When fleas are hatched in a nest they have no choice but to attach themselves to the young mammals or birds. But even in that case they frequently leave their hosts and do not for very long remain stationary. Moreover, when a host dies and becomes cold the fleas invariably leave their quarters, which explains how it may happen that Carnivora get infested with the fleas of their prey. This change of hosts which is always occurring makes it impossible to draw conclusions from material collected in zoological gardens where many animals are herded together. In menageries, too, the normal conditions of breeding are absent. A German naturalist collected 2036 fleas from theatres, concert-halls, ball-rooms, schools and barracks in the grand-duchy of Baden and found that more than fifty per cent. were dog-fleas (Ctenocephalus canis). What the proportion may be in other parts of Europe we have no materials from which to form a judgment. In zoological gardens cat-fleas (Ct. felis) are generally numerous in most of the cages.It is, of course, well known to every zoologist that species are not fixed or constant and that various forms of mammal or of bird tend to show geographical variations. When a long series of skins are laid out on a table and carefully examined it is seldom that those from the west of any great region cannot be picked out and distinguished from those obtained inthe east. So we also get northern and southern forms of the same species varying slightly. These variations are perceptible in many forms of insects, and zoologists now describe these local races as subspecies and designate them with trinomials. No one, however, knows enough as yet about all the various forms which are assumed by fleas to attempt, except in a few instances, to do so in the case of these animals.The study of Siphonaptera is still quite in its infancy. We know little or nothing of the minute geographical variation of fleas. That there is such a thing can already be seen in a few species. In the meantime the study of variation must be postponed until collectors have amassed a more plentiful amount of material; and it is best to treat all forms which are to all appearance constantly different as being specifically distinct until more is known about variation.Any classification of fleas that may now be attempted can only be tentative. It will be enough for present purposes if the reader will remember that the Order Siphonaptera can be divided into three groups or families: (1) the chigoes and their allies, which are the most parasitic fleas (Sarcopsyllidæ); (2) the typical fleas to which the majority of species belong (Pulicidæ); and (3) the bat-fleas (Ceratopsyllidæ), which have certain peculiarities that will be described in a later chapter.Of the antiquity of fleas, and of the period in geological history when the order made its appearance, little can be said. When it was thought that fleas were confined, as parasites, to warm-blooded mammals and birds, evolutionists were inclined to say that the parasites could not have appeared before their hosts. The discovery of a flea on a reptile opened the vista of possibly enormous antiquity stretching back to Permian or Carboniferous ages. The fossil record is most meagre. If we reject as too doubtful the supposed remains of a flea from the lower Oligocene strata at Aix in Provence, only one undoubted fossil has been discovered. Nor does it seem certain that fleas are entirely restricted to preying on vertebrates. Dr Dampf introduced a number of common bird-fleas (Ceratophyllus gallinæ) of both sexes to some hairy caterpillars. He observed that several of the fleas buried their heads in the hairy covering of the larvæ and remained some time in the attitude of sucking blood. While this was going on the victims made violent demonstrations of annoyance and discomfort. He also observed that a naked caterpillar was not attacked.Mr Boden has also recorded how he found in a seed-warehouse some peas that were being eaten by two species of Lepidopterous larvæ. On bringing these home and keeping them in a jar, he found among them some small larvæ which ultimatelyturned into fleas, probablyPulex irritans. These fleas, being confined without other food, were observed to prey on the Lepidopterous larvæ and to feed freely on their juices. The larvæ which were attacked by fleas pined and died. The fluid from the stomach of the fleas when they were crushed was transparent and not red like vertebrate blood which often exudes when a mammalian flea is pinched and cracked open.A French entomologist has also reported that the numerous fleas which swarm in the dwelling-houses of Corsica, for want of other nourishment turn their attention to flies that may be incapable of flight.The only fossil remains of a flea that have, so far, been discovered are a single insect in a piece of Baltic amber of Oligocene age. Many organic remains have been preserved in this fashion, but this is the first mammalian parasite that has been found. The flea is admirably protected by its semi-transparent surroundings, and the most minute details of structure, the arrangement of bristles on the body, and the number of segments to the labial palpi can be discerned. This unique object is in the collection of Professor Klebs. The first point to note is that a flea of this antiquity hardly differs from the existing insects of the present day. It has been referred to an existing genus (Palæopsylla) of which there areat least four species living. Three of these are parasites of the mole, and the fourth is found on shrews. There is good reason to suppose that the host of the fossil was some insectivorous mammal. The early specialisation of fleas is strikingly illustrated. This insect is already adapted for life on some warm-blooded animal. It has a thoracic comb, and its mouth-parts are in all respects like those of a modern flea. It belongs to a genus which is still commonly distributed over Europe. When we consider how remote are the chances that a mammalian flea should first get embedded in amber and should, subsequently, be detected and described by a naturalist, we may well understand that the owner of the fossil asked, though without success, £1200 for it.The ordinary person regards fleas as a subject for humour of an obvious and familiar kind. The utilitarian despises a man who can cheerfully spend his time in collecting fleas. Yet it seems probable that a study of their forms and habits may be of immediate benefit to the human race. The discovery that fleas are connected with the spread of plague is an instance of apparently unprofitable scientific labour proving of direct advantage to mankind. An accurate knowledge of the structure and habits of fleas is now seen to be of importance to all who are engaged in fighting one of the most dreaded infectious diseases. When plague breaks out men of science now at onceturn their attention to the fleas. This is likely to prove more directly efficacious than the mediæval custom of marking the house with a red cross and inscribing the legend, “God have mercy on us.”c2CHAPTER IITHE EXTERNAL STRUCTURE OF A FLEAIncomparing the structure of a flea with that of a man, or any other of the higher animals, it is of the utmost importance to understand that the one has an internal and the other an external skeleton. In either case the skeleton serves as an attachment for the muscles by which the animal moves itself. Everyone is familiar with the external skeleton of a lobster and can see for himself how the muscles are attached. The structure of a flea, though so much smaller, is somewhat similar, except that the skeleton is composed of a horny substance known aschitininstead of being calcified. The chitinous cuticle entirely covers the flea, but it varies in hardness and thickness on different parts of the body. The epidermis, or true skin, lies immediately beneath. On those parts of the body which are to the rear of the head the chitin forms a series of plates or shields which overlap one another somewhat like thetiles of a house. The segmented structure of a flea is there most clearly seen; this we may suppose is an inheritance from the segmented worms.The chitin which forms the external skeleton of a flea is secreted by an outer layer of cells on the insect’s body. The deposit being thin at the joints, and thick on the plates, which serve for protection, the flea is encased in a suit of flexible armour. It is made of a fairly solid and dense substance, but, owing to the absence of carbonate and phosphate of lime, is much lighter than the familiar external skeleton of the lobster. Chitin is a very peculiar and durable substance which resists boiling in acids or alkalies. It is a structureless substance, in the sense that it does not consist of cells. Though horny in appearance it is, of course, in no sense true horn like that of the nails, hoofs, claws, and horns of vertebrates.The different species of flea vary considerably in size. Some are smaller than the familiar human flea. Others are much larger. A very large flea (Hystrichopsylla talpæ) is that found on the mole. The largest known flea (Dolichopsyllus stylosus) is found on small rodents in the United States where, as we know, all things are on a bigger scale than in the Old World. It is seven millimetres long.The colour of the horny integument varies from a pale or light yellow to a ruddy or dark brown.It is plentifully sprinkled over with spines, bristles, or hairs, directed backwards so as not to impede progress. The presence or absence, the arrangement on the body, and the size of these serve, along with other features, to distinguish different species. They seem materially to help a flea in those wriggles to escape with which we are all familiar. The bristles are not always the same in size and arrangement in the two sexes of the same species of flea. As a rule the males are more bristly than the females.These appendages of the flea’s integument are called by various writers either spines, bristles, or hairs. There is, however, no real distinction in the structure or nature of the appendages, and it is a question of degree which name is most appropriate.In distinguishing species, very little reliance can be placed on the colour of the flea. An insect newly emerged from the pupa is always lighter in colour; and the difference between the appearance of an empty stomach and a stomach gorged with red blood is surprising.The general external appearance of a typical flea belonging to the main familyPulicidæis fairly familiar to most persons. This is well seen in the figure of the oriental rat-flea. The body is compressed or flattened from side to side, and this is a feature which is extremely rare among insects. It doubtlessenables the animal to glide with greater facility through the hairs of its host. Like other insects, a flea is readily divided into a head, a thorax, and an abdomen. The head is rounded on the top and front and shows no obvious trace of segmentation; but what is known of the development of other insects leads one to think that it must properly be regarded as a number of segments closely fused together. On the under side of the front part of the head is a beak or proboscis for piercing and sucking, composed of the mouth-parts, whose structure is worthy of minute study. It will be best to examine them in detail in a subsequent chapter.Some fleas have eyes, others have none. The common mouse-flea (Leptopsylla musculi) is blind. The bat-fleas are also destitute of eyes. The nocturnal habits of their hosts would render eyes of little or no use. If eyes are present they are large and placed on either side of the head. Each is a simple eye or ocellus; the compound eyes, divided into a great number of hexagonal facets, which are characteristic of many insects, are never found in fleas.Nothing is known about the flea’s powers of vision, but there is no reason to suppose that they are at all acute. The eyes are marked with pigment. Ocelli appear to be primitive types of insect eye which are, perhaps, an inheritance from a wormlikeancestor. Presumably all the fleas of long ago had eyes and those that are now blind have lost their organs of sight from disuse. In their simplest condition, the eyes of the lower invertebrates only enable the creature, so far as one can judge, to distinguish light from darkness. Entomologists believe that the power of vision of ocelli is probably confined to very near objects and that this simple form of eye is more useful in dark places than the compound eyes. There is no reason for believing that fleas can distinguish colours or can discern any object which is more than a few inches away. It is enough for their purpose to perceive from which point light comes upon them and to make all despatch to escape in the opposite direction.In blind fleas there is often a spine where the eye should be. In one species the spine is rudimentary and there is some black pigment beneath it. It is not impossible that this is the vestige of a once functional eye. In one genus, however, the eye and the spine are both present. Of the fleas belonging to this genus one species is South American and the other European. The latter (Typhloceras poppei) is confined to the long-tailed field-mouse.The organs by which fleas keep in touch with the outward world, and with other fleas, are their antennæ. All fleas have antennæ; but unlike those of a moth, a beetle, or a grasshopper, each fits neatlyinto a groove at the side of the head and can be protruded when desired. This is another adaptation to enable the insect to creep swiftly through a forest of hairs.i26Fig. 2. Showing a type of (a)genaland (b)thoraciccombs of a flea, on the under part of the head and on the thorax respectively. Analagous combs are found in several other parasitic insects and on the abdominal segments of certain fleas.The combs which are found on the heads of many fleas are organs of exceptional interest (Fig. 2). They are toothed and horny appendages, which are connected with parasitic habits, for somewhat similar combs are found on several unrelated groups of parasitic insects, as, for instance, on parasitic beetles (Platypsyllus) found on the beaver, on insects allied to bed-bugs (Polyctenes) found on bats, and on wingless flies (Nycteribia) which infest Egyptian and South American bats. The majority of thePulicidæhave one or more combs with comparatively long teeth. These combs reach their maximumdevelopment in the bat-fleas which have no less than eight. Some Australian and South American fleas (Stephanocircus) have a helmet-like comb extending all round their heads. These combs are by some supposed to be of service in holding on to the hairs of the host; and, if one may judge from experiments made on live fleas in cotton wool, they are also used in moving forwards through the fur.All the chigoes (Sarcopsyllidæ) have a large triangular post-oral process which is more or less curved and probably prevents the flea slipping back as it pushes forward. Bat-fleas (Ceratopsyllidæ) have lobes or flaps placed two on each side of the head, which may possibly serve an identical purpose, but whether they do so is not known.These combs may be divided into three groups according to the part of the flea’s body on which they are found. Those found on the head are called genal combs and take the form shown inFig. 2. There are also combs found on the thorax. The fossil flea (Palæopsylla klebsi) described in the previous chapter has one of these thoracic combs. A certain number of fleas also have combs on the abdominal segments.There are really two types of toothed organs on fleas to both of which the name of comb may be given. One is composed of a sheet of chitin with a number of slits and teeth and resembles a true comb.The other consists merely of a number of highly chitinised bristles arranged in a row. They probably both serve the same function. Apart from their use as organs to assist movement onwards, they may also serve ashair-tightjoints and protect the flea from the inconvenience of getting the tips of the host’s fur into the joints of its horny armour.The size of the head compared to the thorax and abdomen varies considerably in different species. Some fleas have what may be called by comparison large heads and others very small ones.A small head is never found in a flea with powerful mouth-parts. The head being the bearer of piercing and sucking organs, which require strong muscles, there must be room not only for the organs but for their extensors and retractors.There are normally three rows of bristles on a flea’s head which divide the head into four sections. It is possible that these correspond to the four segments of the ancestral insect which are now fused together.The head of a flea is closely applied by the whole of its back surface to the body and that slender and conspicuous neck which is characteristic of the Diptera, or flies, is not to be found in any fleas. For this reason a flea cannot turn its head in any direction without at the same time following it round with its body.The thorax of a flea consists of three segments called respectively the prothorax, mesothorax and metathorax. The chitinous external skeleton which covers each of these three segments is primarily a hoop but each hoop is further subdivided into a number of complicated plates. Attached to the thorax are the three pairs of legs which are characteristic of all adult insects. The hind pair are very much the strongest (Fig. 3). They are the organs of hopping. It has often been pointed out that if men had the leaping powers of some fleas they would bound with ease backwards and forwards over the cross on the top of St. Paul’s Cathedral. Each leg consists of four segments beautifully articulated and plentifully supplied with bristles. At the end comes the foot with five very short segments. The last segment is provided with a pair of more or less formidable claws. Fleas use their legs for leaping, for running, and for clinging to their hosts. They also use their mouth-parts for the last purpose and it is worthy of note, as we shall see later on, that in those fleas in which the mouth-parts are shortest and weakest the legs are most liberally supplied with bristles and possess the stoutest claws. The legs of a flea are unique in the insect world owing to the enormous development of the segment nearest the body called the coxa. Most leaping insects rely for their activity on the muscles of the lower joints. In a grasshopper it is the third joint from the body (femur) which is so immensely enlarged. The three pairs of legs are each attached to a different thoracic segment.i30Fig. 3. The hind leg of a flea. The segment or joint nearest the body is thecoxawhich is unusually developed. Next come the smalltrochanterand the largerfemur. Thetibiawhich is long and slender follows. Then come the fivetarsiwith the sixth and ultimate segment provided with claws.When fleas walk, they are so to speak plantigrades walking on the sole of the foot, and all the tarsal or foot joints are applied to the surface of the ground. The claws serve as grips so as to make the most of any unevenness; and thus the insect drags itself along with surprising rapidity when it moves through the hairy coat of a mammal. But on an open surface fleas are not really rapid movers compared with many other insects.The two claws on the end of each ultimate foot segment are freely moveable and are in fact highly modified bristles or setæ.In all fleas one of the plates of the metathorax (or hindmost thoracic segment) called the epimeron, is large and prolonged towards the rear. It invariably bears a stigma. The epimeron is placed laterally to the first abdominal tergite. The older naturalists jumped to the conclusion that this was the remains of a wing. The best judges have, however, formed a decided opinion that no trace of the relic of a flying organ can be detected on the thorax of a flea. Heymons, a German entomologist, has also failed to detect any sign in dissections which he has made of the larvæ and the pupæ.The epimeron is in fact neither a scale nor awing but a portion of the thorax present in all insects. It is of no special service to the flea except as a portion of the thoracic armature which covers the body.The larva of a flea has no legs; the adult insect has six. A study of other embryo insects shows that the ancestors of insects had many legs. It is an interesting problem why insects lost the legs on their abdomens, why legs should now invariably be restricted to the thorax, and why there should never be more than three pairs. In the earliest known insects which lived on the earth, before winged forms were evolved, the number of legs was already six. But our knowledge of fleas is too small to attempt, at present, to trace their exact line of ancestral descent.The abdomen of a flea consists of ten segments. The horny plates which cover the dorsal side are called tergites; those on the ventral side sternites. In fleas, as in all holometabolous insects, that is those which pass through a complete metamorphosis, the sternite of the first abdominal segment is suppressed and has completely disappeared. The tergite which covers the dorsal part of the first abdominal segment nearest to the thorax is, however, always present.The ultimate segments of the male and female flea are modified for reproductive purposes and of these segments more must be said later.Having now given a rough outline of the external skeleton of a flea, it only remains to say something about the muscular system. Attached to the inside of the chitinous armature are an enormous number of muscles, whitish and almost transparent. They act as extensors, retractors, flexors, elevators and depressors. The joints and hinges of the skeleton allow of considerable, but not perfect, freedom. The muscles of locomotion are partly in the thorax and partly in the several joints of the legs. Our knowledge of the muscular system of fleas is very imperfect. But, as in other insects, the general arrangement of the muscles is based on the segmented structure of the body.For the reader who can accurately picture to himself the external structure of a flea and of the typical insects belonging to other orders, a few words may be said on the probable ancestry of fleas and their relationship to other living insects. This vexed and much debated question is still, as the older naturalists would have said,tremendum mysterium. Very little light has yet been thrown upon it, and the most divergent views have been expressed by learned and competent entomologists. A historic survey of the various opinions that have been held since the days of Linnæus would fill many pages; but a short summary of the different orders to which fleas have been referred by different zoologists will suffice.The older authors, Linnæus, Geoffroy, Cuvier and Duméril, and Gervais placed them among theApterabecause they were wingless. Kircher regarded them asOrthoptera, an order which includes grasshoppers and crickets; but he has had few followers. By Fabricius and by Illiger they were treated asHemipteraor bugs. Lameere, a Belgian, has recently expressed a decided view that fleas are really a family ofColeopteraor beetles. Those who have held the once orthodox opinion that they belonged to theDipteraor flies are Roesel, Oken, Straus-Dürkheim, Burmeister, Newman, Walker, von Siebold and Wagner.The structure of an adult flea, however, differs from that of an adult fly in the following noteworthy respects: the mouth-parts are differently constructed, the head of the flea is closely joined to its thorax, the three divisions of the thorax are not joined and fused, the flea is wingless, the eyes of fleas are simple ocelli, and there are differences of lesser importance in the stigmata, which give access to the tracheal system by which all insects breathe.The number of those who have regarded fleas as belonging to a distinct order of insects is considerable: they are Lamarck, De Geer, Latreille, Kirby and Spence, MacLeay, Leach, Dugès, Bouché, van der Hoeven, Westwood, Landois, Brauer, Kraepelin,and Taschenberg. Modern opinion is all but unanimous on this point.There remains, however, a second question. Even if it be agreed that there must be a distinct order forSuctoria, Aphaniptera, Siphonaptera, or fleas; where ought that order to be placed? In which other order of insects must we look for the nearest relations of fleas? For a time after the acceptance of the fact that insect forms have been evolved, and not separately created, the ancestors of fleas were searched for among some species of fly.Then Kraepelin rejected the view that flies were as closely related to fleas as most entomologists thought and his followers could only find points of difference and no points of resemblance. Dahl (1899), a German, then took up the cudgels for the fly theory. Dahl pointed out the resemblance between fleas and a group of flies calledPhoridæalso parasitic on warm-blooded animals. During the ensuing years the debate was resumed afresh with much liveliness and sometimes with a little acrimony.The fleas were placed by MacLeay and by Balbiani between theDipteraandHemiptera; by Leach between theHemipteraandLepidoptera; by Dugès between theHymenopteraandDiptera; by Brauer between theDipteraandColeoptera. Handlirsch thinks that fleas have no connection atall with beetles and Gross can find no signs of relationship with eitherColeopteraorDiptera.Embryology and the study of larval forms have thrown so much light on the ancestry of many animals, that it was hoped that a microscopic examination of the larvæ of fleas, in various stages of development, would produce some facts of importance. In this hope entomologists have, to a great extent, been disappointed. There seems to be much similarity between the embryos of beetles, moths, flies, wasps and fleas. Those who have dwelt on the likeness of the larval flea to the maggot of a fly seem to forget that the resemblance to an embryo beetle is nearly as strong.The young larva of the flea is very transparent and the digestive canal, heart and nervous system are easily recognised. The egg-shell breaker is an interesting example of the development of a temporary larval structure and it is the only known instance of such a structure in an insect. There are no traces of eyes. The antennæ are three-jointed. They are rather long and slender, being about one-third as long as the head. The head is well-developed and the larva has no feet.The biting mandibles are broad and triangular. Compared with those of other larvæ they are said to be more like the mandibles of coleopterous than of dipterous larvæ. The maxillæ, or second pair ofjaws, are somewhat reduced and rudimentary. The absence of eyes and of legs are points of similarity between the larvæ of fleas and flies. The maggot of a fly has also two pairs of jaws, and a pair of antennæ.At the tail end of the larval flea’s abdomen are two small projections called caudal stylets (Fig. 1). They are strong, recurved, chitinous, structures which prop up the body of the larva when it creeps and wriggles. There are similar props in the larvæ of certain beetles and no exactly similar organs are known in dipterous larvæ. But caudal stylets are of small taxonomic importance.In one respect the mature flea is certainly nearer to a beetle than to a fly: the three joints of the thorax are free as in a beetle and not fused as in a fly; but when one studies the mouth-parts, the true view seems to be that the mouth-parts of a flea are equally unlike those of a fly and those of a beetle. Such being the present state of our knowledge, one must wait for fresh light to be thrown on the matter by further researches. It seems unlikely that the immediate future will produce a solution of the problem.

TRANSCRIBER’S NOTES:—Obvious print and punctuation errors were corrected.—The decorated front fage has been retained as illustration.

TRANSCRIBER’S NOTES:—Obvious print and punctuation errors were corrected.—The decorated front fage has been retained as illustration.

TRANSCRIBER’S NOTES:

—Obvious print and punctuation errors were corrected.

—The decorated front fage has been retained as illustration.

The Cambridge Manuals of Science and LiteratureTHE FLEACAMBRIDGE UNIVERSITY PRESSLondon: FETTER LANE, E.C.C. F. CLAY,ManagerEdinburgh: 100, PRINCES STREETLondon: H. K. LEWIS, 136, GOWER STREET, W.C.WILLIAM WESLEY & SON, 28, ESSEX STREET, STRANDBerlin: A. ASHER AND CO.Leipzig: F. A. BROCKHAUSNew York: G. P. PUTNAM’S SONSBombay and Calcutta: MACMILLAN AND CO., LTD.All rights reservedAfter a drawing by Dr JordanOriental rat-flea (Xenopsylla cheopisRothsch.). Male.CambridgePRINTED BY JOHN CLAY, M.A.AT THE UNIVERSITY PRESSWith the exception of the coat of arms at the foot, the design on the title page is a reproduction of one used by the earliest known Cambridge printer, John Siberch, 1521

The Cambridge Manuals of Science and Literature

CAMBRIDGE UNIVERSITY PRESS

London: FETTER LANE, E.C.C. F. CLAY,Manager

Edinburgh: 100, PRINCES STREETLondon: H. K. LEWIS, 136, GOWER STREET, W.C.WILLIAM WESLEY & SON, 28, ESSEX STREET, STRANDBerlin: A. ASHER AND CO.Leipzig: F. A. BROCKHAUSNew York: G. P. PUTNAM’S SONSBombay and Calcutta: MACMILLAN AND CO., LTD.

All rights reserved

After a drawing by Dr JordanOriental rat-flea (Xenopsylla cheopisRothsch.). Male.

After a drawing by Dr JordanOriental rat-flea (Xenopsylla cheopisRothsch.). Male.

After a drawing by Dr Jordan

Oriental rat-flea (Xenopsylla cheopisRothsch.). Male.

CambridgePRINTED BY JOHN CLAY, M.A.AT THE UNIVERSITY PRESS

With the exception of the coat of arms at the foot, the design on the title page is a reproduction of one used by the earliest known Cambridge printer, John Siberch, 1521

cpPREFACETHE aim of this book is to give in plain language some account of a small, but noteworthy, group of insects. I have avoided, whenever I could, using the technical terms of zoology. To avoid doing so entirely is impossible in a book which describes insects in some detail. No technical term has, I hope, been used without an explanation.Over thirty years have elapsed since Taschenberg’s German book,Die Flöhe, appeared. Our knowledge has made enormous strides since then. More species of flea are now known from the British Islands alone than were then known from the whole world. So far as I am aware, no book, devoted to what is known about fleas, has ever been published in English. The statements about these insects in the general text-books of entomology are frequently antiquated and inaccurate. But there is a fairly extensive literature on theSiphonapterascattered through scientific periodicals mostly in English, German, Italian, Dutch and Russian. I have given some references in the Bibliography.The naturalists now living who have devoted any time to the special study of fleas may almost be counted on one’s fingers. In England there are Mr Charles Rothschild and Dr Jordan; in the Shetland Islands, the Rev. James Waterston; in Germany, Taschenberg of Halle and Dampf of Königsberg; in Russia, Wagner of Kieff; in Holland, Oudemans of Arnhem; in Italy, Tiraboschi of Rome; in the United States, Carl Baker and a few others. I have not mentioned medical men who have investigated fleas in connection with plague.There are small collections of fleas in the Natural History Museums at South Kensington (London), Paris, Berlin, Königsberg, Vienna, Budapest, S. Petersburg and Washington. Of private collections Mr Charles Rothschild’s at Tring is by far the best in the world. It contains something like a hundred thousand specimens and is most admirably kept. I must express profound and sincere gratitude to Mr Rothschild for having helped me in numberless ways and advised me in many difficulties.It is well known that the mere mention of fleas is not only considered a subject for merriment, but in some people produces, by subjective suggestion, violent irritation of the skin. The scientific study of fleas has, however, received a great impetus since it has been ascertained that they are the active agentsin spreading plague. Rat-fleas are of various kinds, and not all fleas will bite man. A knowledge of the different species has suddenly become useful. The humble, but ridiculous, systematist with his glass tubes of alcohol for collecting fleas, his microscopic distinctions, and Latin nomenclature has become a benefactor of humanity. Some people seem to be practically immune to the bites of fleas, but even to such persons their visits are unwelcome. A famous Frenchwoman once declared: “Quant à moi ce n’est pas la morsure, c’est la promenade.”H. R.London,September, 1913.

cp

THE aim of this book is to give in plain language some account of a small, but noteworthy, group of insects. I have avoided, whenever I could, using the technical terms of zoology. To avoid doing so entirely is impossible in a book which describes insects in some detail. No technical term has, I hope, been used without an explanation.

Over thirty years have elapsed since Taschenberg’s German book,Die Flöhe, appeared. Our knowledge has made enormous strides since then. More species of flea are now known from the British Islands alone than were then known from the whole world. So far as I am aware, no book, devoted to what is known about fleas, has ever been published in English. The statements about these insects in the general text-books of entomology are frequently antiquated and inaccurate. But there is a fairly extensive literature on theSiphonapterascattered through scientific periodicals mostly in English, German, Italian, Dutch and Russian. I have given some references in the Bibliography.

The naturalists now living who have devoted any time to the special study of fleas may almost be counted on one’s fingers. In England there are Mr Charles Rothschild and Dr Jordan; in the Shetland Islands, the Rev. James Waterston; in Germany, Taschenberg of Halle and Dampf of Königsberg; in Russia, Wagner of Kieff; in Holland, Oudemans of Arnhem; in Italy, Tiraboschi of Rome; in the United States, Carl Baker and a few others. I have not mentioned medical men who have investigated fleas in connection with plague.

There are small collections of fleas in the Natural History Museums at South Kensington (London), Paris, Berlin, Königsberg, Vienna, Budapest, S. Petersburg and Washington. Of private collections Mr Charles Rothschild’s at Tring is by far the best in the world. It contains something like a hundred thousand specimens and is most admirably kept. I must express profound and sincere gratitude to Mr Rothschild for having helped me in numberless ways and advised me in many difficulties.

It is well known that the mere mention of fleas is not only considered a subject for merriment, but in some people produces, by subjective suggestion, violent irritation of the skin. The scientific study of fleas has, however, received a great impetus since it has been ascertained that they are the active agentsin spreading plague. Rat-fleas are of various kinds, and not all fleas will bite man. A knowledge of the different species has suddenly become useful. The humble, but ridiculous, systematist with his glass tubes of alcohol for collecting fleas, his microscopic distinctions, and Latin nomenclature has become a benefactor of humanity. Some people seem to be practically immune to the bites of fleas, but even to such persons their visits are unwelcome. A famous Frenchwoman once declared: “Quant à moi ce n’est pas la morsure, c’est la promenade.”

H. R.

London,

September, 1913.

CONTENTSCHAP.PAGEPrefacevI.Introductory1II.The external structure of a flea21III.The mouth-parts and sense-organs38IV.The internal organs of a flea52V.The Human flea and other species62VI.The Chigoes and their allies74VII.Fleas and Plague83VIII.Rat-fleas and Bat-fleas97AppendixA.Systematic view of the orderSiphonaptera108”B.A list of British fleas and their hosts110”C.On collecting and preserving fleas113”D.Bibliography118Index122

LIST OF ILLUSTRATIONSMale Oriental rat–fleafrontispieceFIGUREPAGE1.The larva of a flea62.Types of genal and thoracic combs of a flea263.The hind leg of a flea304.The mouth-parts of a flea435.The antenna of a flea476.The alimentary canal of a flea537.The head of a female dog-flea and a female cat-flea718.Pregnant female ofDermatophilus cæcata81

c1CHAPTER IINTRODUCTORYFleasform a group of insects that have, until recently, been little studied by zoologists. We call them insects because they are jointed animals, or Arthropods, with three pairs of legs in the adult condition. The reader will best understand the position which fleas occupy in the general classification of animals by remembering that the arthropods, or jointed animals, are one of a dozen subkingdoms, or phyla, to which the various members of the great animal kingdom have been assigned. There is good ground for believing that all the animals included in each phylum trace their ancestry back to a common primitive form which lived in more or less remote ages. Besides (1)Insects, the arthropods, or jointed animals, include (2)Crustaceans, such as crabs, lobsters, shrimps, wood-lice, water-fleas and barnacles; (3)Myriapods, such as centipedes and millipedes; and (4)Arachnids, such as spiders, scorpions, mites and ticks. To all these varied forms of animal life fleas, and other insects, are therefore more or less nearly related.The animals belonging to this large and important collection, which compose the arthropod phylum, have certain common characteristic features. We find a body made up of a series of more or less completely similar segments placed one behind the other. In this they resemble certain worms which are far less highly organised. The body is elongated, symmetrical on either side, and the mouth and anus are at opposite ends. There is, however, an important advance on the segmented worms. Each typical segment carries a pair of appendages which are very different from the foot-stumps that are found on certain worms. These appendages of arthropods are divisible into distinct limb-segments, separated from one another by moveable joints, and acted upon by special muscles.The common ancestor of all the various arthropods which are found living on the earth to-day, was probably composed of a series of segments each very similar to the last and each bearing a pair of very similar appendages. In the course of ages, these appendages have been astoundingly modified in form and in function. So it happens that we find in the arthropods of the present day pairs of antennæ, of mandibles and other mouth-parts, of pincers, of legs, of swimming-feet and of tail pieces which on close examination can all be traced back to a common structure. The body-segments, also, have beenstrangely fused together and modified. All that has been so far said applies equally to fleas and to other insects.It is of great interest, when one comes to make a minute study of the form and external structure of a flea, to try and trace the modifications that must have taken place in the course of descent from the ancestral arthropod; but the relationship of fleas to other insects living at the present day is of more immediate concern. Insects are highly specialized arthropods and fleas are highly specialized insects. This means that they have become vastly modified from the primitive ancestral type and fitted thereby for a life among certain defined and peculiar surroundings.It will be unnecessary to remind the reader who knows anything of zoology or of botany that all classification is now based on descent. Since naturalists have abandoned a belief in the special creation of the various species of animals now living on the earth and have conclusively shown that they have arisen by descent and modification from other forms, the problem is to reconstruct a vast genealogical tree. What then were the ancestors of the fleas and to what other insects, in consequence, do they appear to be related?It is probable that the ancestors of the fleas were winged insects, and that the organs of flight weregradually lost, as they became useless, when a partially parasitic life was adopted. At one time entomologists regarded fleas as wingless flies and placed them in the order Diptera. Certain supposed scaly plates on their bodies were regarded as the atrophied relics of wings. It is, however, more than doubtful whether this view is correct; and all modern entomologists who have given any special study to fleas are agreed that they are sufficiently unlike any other living insects to deserve a place in an order by themselves. To this order the nameSiphonapterahas been given: which means that the insects comprised in it are provided with sucking mouths and are destitute of wings. Another name for the order is Aphaniptera, but this is gradually falling into disuse. Linnæus (1758) only mentions two species of flea: the human flea which he appropriately namedPulex irritans, and the chigoe of hot countries which he calledPulex penetrans, from the habit which the female has of burrowing under the skin of her victims. At the time of writing, about 460 species of flea have been described and named; but some of the names are doubtless synonymous, and the actual number of separable species that have been discovered is somewhere about four hundred. The vast majority of these have been described within the last few years, which shows what can be done when attention is turned to any neglected group of animals. Therecan be no doubt that many undiscovered species still remain, and will now, in due course, be collected, described and named.The position which should be assigned to the order Siphonaptera in the general scheme of insect classification is a question on which the most learned modern entomologists have disputed with considerable vigour. Some see the nearest relatives among the beetles, others among the flies. The majority, as we shall see later on, would place them near the Diptera: but since no convincing arguments have been produced on either side it may be wisest to regard the question as still at present unsolved.Fleas belong to one of the groups of insects which go through a complete metamorphosis. Their life-history consequently falls into four divisions: egg, larva, pupa and imago. If the climate permits, the female flea lays her eggs all the year round, and from one to five are dropped at a time. Unlike those of many other parasites they are never attached to the hairs of the hosts, but appear to be deposited indiscriminately on the floors of houses or in the nests and sleeping places of their hosts. The eggs generally hatch in a few days, and a minute, white, wormlike larva emerges (Fig. 1). The larvæ, of some, and possibly of all, fleas are provided with a wonderful adaptation in the shape of an egg-breaker or hatching-spine. This is a thin plate, like the edge of a knife, where the point of the head comes in contact with the shell. The movements of the prisoner make a slight split in the egg-shell, which then bursts asunder. This organ has vanished in later larval life, and it is probably lost after the first moult. The larva is legless and has thirteen segments. It grows rapidly, and, as it grows, moults its skin several times. It is provided with mouth-parts adapted for biting, and eats any decaying organic refuse. The larvæ may be reared on the sweepings of an ordinary room or the dirty scurf which collects at the bottom of old birds’ nests. It is hardly necessary to add that the mother takes no interest whatever in the larvæ and that the belief that she feeds them on dried blood is not based on any sound foundations.i6Fig. 1. The larva of a flea. The body consists of thirteen segments and is legless. On the fore part of the head are the antennæ and on the upper part of the head is shown the knife-like edge of the egg-breaker. The mouth-parts are adapted for biting. On the last segment of the body are the two caudal stylets.The larval stage lasts some days, and the animal spins a small cocoon before pupating. In the course of a few more days, the time probably depending on the weather, the perfect flea emerges. The larvæ generally live in places where the perfect insects will have an opportunity of finding a host as soon as they leave the pupal envelope. The nests of their hosts where the young are being reared are always favourite places. It seems possible that the comparative immunity from fleas which hoofed mammals or Ungulates enjoy may be due to the fact that the young beast follows its mother from the time of birth instead of passing its early life helpless in a nest.Observations made on the development of the dog-flea (Ctenocephalus canis) in India show that eggs laid on October 17 hatched on October 19. The larva spun its cocoon on October 25 and the mature flea emerged on November 2. In Northern Europe the human flea takes about four weeks in summer and six weeks in winter to pass through its metamorphosis.Unlike many parasitic insects, fleas do not constantly pass their time upon the bodies of their victims. The greater part of their life is probably spent on the ground, in the house, or nest, of the mammal or bird which serves them with blood. In this respect there is considerable difference in the habits of different species of flea. Some attach themselves to an animal and actually burrow into the skin. These are the most parasitic species. Some only come to feed and leave to lay their eggs. Many probably do not suck blood more than once in their lives.An animal which harbours fleas and which nourishes the adult insect with blood is called ahost. No fleas are more than what is called temporary parasites; which means that they pass but a portion of their lives on their hosts and frequently take occasion to hop on and off. All fleas, apparently, go from host to host. The labours of diligent collectors have proved that the greatmajority of mammals and birds have fleas. As a general rule, it is true to say that certain species of flea are associated with certain species of host. Thus man is the true host ofPulex irritans; the cat family are the true hosts of the cat-flea (Ctenocephalus felis); and the dog family are the true hosts of the dog-flea (Ctenocephalus canis). But the human flea is sometimes found on cats and dogs, and cat and dog-fleas occasionally bite human beings; and cat-fleas are found on dogs and dog-fleas are found on cats. All fleas, so far as we know, may occasionally pass from one species of host to another; but they do not, for the most part, seem to flourish in unaccustomed quarters. Some fleas are more catholic in their tastes than others. Some seem to be very strictly confined to one host, and even when starving only suck strange blood under protest. There is a species of flea that has only (except by accident) been found on the long-tailed field-mouse and another that has only been found on the hedgehog. Other fleas are commonly found on two absolutely distinct animals; a good instance of this is the human flea which, at all events in certain parts of England, is a regular parasite of the badger.As distinguished from true or natural hosts one must separate what may be termed casual or accidental hosts. All animals which come in contact with one another, or which live in close proximity,may exchange fleas. So even bird-fleas may be collected from mammals and typically mammalian fleas from birds. In this fashion puzzles may arise which tax the ingenuity of the collector to solve. Bird-fleas are sometimes found on bats, and this may be obviously attributed to the bats having inhabited a hole which was tenanted by starlings or an old loft infested with the fleas of pigeons. All beasts of prey are sometimes found to harbour the fleas of animals they have devoured. Rabbits’ fleas are found on wild-cats; hedgehogs’ fleas on foxes; mice fleas on weasels; and fleas characteristic of small birds on stoats. So also in the case of mice, rats and voles with holes and runs in the same hedgerow, the parasites usually peculiar to one are not uncommonly found on the others. It is sometimes difficult to determine the true host of a flea.Much more puzzling to explain are the reasons which confine a flea to a certain host and which cause closely allied hosts to have different fleas. The fleas from the house-martin and the sand-martin are quite different; those from the domestic fowl and the domestic pigeon are distinct species. The causes which have affected the evolution of the various forms of flea are too obscure to enable anyone at the present day to offer any satisfactory explanation.Speaking generally, the fleas found on birds have points in common, and they probably form a naturalgroup to themselves. What may be called true bird-fleas have been collected from almost all European birds. An unwieldy genus (Ceratophyllus) comprises many species of different flea. Some species are very abundant and infest the nests of many different birds. Others are extremely rare. One of these rarities (C. vagabundus) is found in the nests of puffins and other sea-birds. Another has been collected on antarctic petrels. Penguins have a special genus of flea to themselves. A specimen, unique at one time (Ceratophyllus borealis), in Mr N. C. Rothschild’s collection was obtained from the gannet. It has now been found on rock-pipits in the Shetland Islands.Two very rare fleas (C. farreniand C.rothschildi) are found in the nests of house-martins; yet the nests of these birds are infested with common species besides. A plague flea (Xenopsylla) has been found on an African swift.Forty-six different species of flea have been found in the British Islands, but many of these are extremely scarce.We know too little about the geographical distribution of fleas to lay down many accurate generalities. When a great deal more material has been collected and studied, it may be possible to show that certain groups are associated with certain regions of the earth or certain orders of animals. To some extent this is already seen to be the case. The fleasindigenous to the New World are distantly related to those of the Old World. Broadly speaking the geographical distribution of the parasite must follow that of the host. But sometimes the parasite is impatient of cold and cannot follow the host out of the tropics. The chigoes and their allies are fleas of hot countries. Different kinds of bats are found from the tropics to the Arctic circle, but the same bat-fleas are not found everywhere.When a flea has a cosmopolitan range it is probable that it has travelled over the world in company with its host.Monkeys have no fleas. This is an assertion that is commonly received with surprise and incredulity. Occasionally a gorilla or a chimpanzee may get a chigoe in its toe. And monkeys in zoological gardens or menageries are possibly exposed to the danger of catching an occasional human flea from the people who crowd round their cages. These are remote contingencies which may happen to anyone. Healthy wild monkeys are much too clean and active to harbour fleas. When they search one another’s fur in a fashion that must be familiar to most persons, they are clearing their coats of particles of scurf or of similar scraps of dirt and not of fleas. So, speaking generally, it may be said that no fleas have been found truly parasitic on monkeys.Bats have fleas, but not in great abundance. Allbat-fleas are rare on their hosts and extremely difficult to find and collect. The same species are not found on fruit-bats and the ordinary smaller insect-eating bats. The geographical distribution of some bat-fleas is puzzling. For instance, one species (Ischnopsylla unipectinata) is found on the greater horse-shoe bat in Europe; but it is, apparently, not found on the same bat in the British Islands. In Somaliland and in India it is found on other bats.With certain exceptions, Ungulates are remarkably free from fleas. This great order of mammals includes a variety of hoofed animals: oxen, sheep, goats, deer, pigs, camels, giraffes and antelopes.The only true fleas found on these are two species of the genusVermipsylla, which resemble the chigoes in so far that the pregnant females burrow into the host and expand there. One species has been found on camels and horses in Transcaucasia; another on roe-deer in Northern China. The female of the last is often found ensconced on the inside of the nostrils of the deer. Of course chigoes may attack domestic Ungulates of all kinds; but no other members of the familyPulicidæor typical fleas except those two above mentioned have been found on hoofed mammals.Insectivora such as moles, shrews and hedgehogs are the hosts of a great variety of species. The same thing may be said of the Rodents, which includeporcupines, squirrels, rats, mice and a vast number of other small mammals whose geographical distribution includes almost the whole of the habitable globe. Probably more different species of fleas have been collected from Insectivora and Rodents than from all the other orders of mammals grouped together.The Carnivora, excluding the Pinnepedia, or seals, sea-lions and walruses, harbour numerous species.Among the Edentata a very remarkable and highly specialised genus of fleas is parasitic on armadilloes in South America. This genus (Malacopsylla) consists of two species only, which are confined to South America and are found on the armadilloes and on carnivorous animals which probably have preyed on them. The thorax of these fleas is much reduced and very small in size. Their piercing organs are slender and weak, but they possess enormous spines on the legs with which they hold on to their hosts. These two South American fleas (M. grossiventrisandM. androcli) will be referred to again later as striking examples of fleas with strongly developed legs and weakly constructed mouth-parts. The contrary combination of powerful mouths and degenerate legs is also found in other groups of fleas, as will be seen in the chapter on the chigoes.The Marsupials of Australia and South America have special fleas which were probably associated with this strange order of pouched mammals beforethey became divided into the American and Australian groups. Fleas have been collected on the spiny ant-eater (Echidna) which belongs to the lowest order of Monotremes or egg-laying mammals.On almost every form of bird, including the most aquatic kinds, fleas of various species have been obtained.Only one instance has been recorded of a flea occurring on a reptile. A female of one of the species of burrowing chigoes (Echidnophaga ambulans) from Australia was collected by Dr Woodward from the Brown Snake (Diemenia superciliosa). This reptile, which is well known in Australia, belongs to a sub-family that contains some of the most deadly poisonous snakes and is allied to the cobras. The Brown Snake is a terrestrial snake, and one must regard the presence of the flea on such a host as a rare and chance occurrence. The snake was captured at Herdman’s Lake, near Perth in West Australia. The same species of flea has also been obtained from the phalangers (Trichosurus) which live in the tops of the Australian gum-trees; from the little terrestrial and nocturnal rat kangaroos (Bettongia); and from the banded ant-eater (Myrmecobius), another Australian Marsupial. It is possible that the flea moved from some small mammal which was being devoured by the snake and managed to fix itself between the scaly plates of the reptile.When fleas are hatched in a nest they have no choice but to attach themselves to the young mammals or birds. But even in that case they frequently leave their hosts and do not for very long remain stationary. Moreover, when a host dies and becomes cold the fleas invariably leave their quarters, which explains how it may happen that Carnivora get infested with the fleas of their prey. This change of hosts which is always occurring makes it impossible to draw conclusions from material collected in zoological gardens where many animals are herded together. In menageries, too, the normal conditions of breeding are absent. A German naturalist collected 2036 fleas from theatres, concert-halls, ball-rooms, schools and barracks in the grand-duchy of Baden and found that more than fifty per cent. were dog-fleas (Ctenocephalus canis). What the proportion may be in other parts of Europe we have no materials from which to form a judgment. In zoological gardens cat-fleas (Ct. felis) are generally numerous in most of the cages.It is, of course, well known to every zoologist that species are not fixed or constant and that various forms of mammal or of bird tend to show geographical variations. When a long series of skins are laid out on a table and carefully examined it is seldom that those from the west of any great region cannot be picked out and distinguished from those obtained inthe east. So we also get northern and southern forms of the same species varying slightly. These variations are perceptible in many forms of insects, and zoologists now describe these local races as subspecies and designate them with trinomials. No one, however, knows enough as yet about all the various forms which are assumed by fleas to attempt, except in a few instances, to do so in the case of these animals.The study of Siphonaptera is still quite in its infancy. We know little or nothing of the minute geographical variation of fleas. That there is such a thing can already be seen in a few species. In the meantime the study of variation must be postponed until collectors have amassed a more plentiful amount of material; and it is best to treat all forms which are to all appearance constantly different as being specifically distinct until more is known about variation.Any classification of fleas that may now be attempted can only be tentative. It will be enough for present purposes if the reader will remember that the Order Siphonaptera can be divided into three groups or families: (1) the chigoes and their allies, which are the most parasitic fleas (Sarcopsyllidæ); (2) the typical fleas to which the majority of species belong (Pulicidæ); and (3) the bat-fleas (Ceratopsyllidæ), which have certain peculiarities that will be described in a later chapter.Of the antiquity of fleas, and of the period in geological history when the order made its appearance, little can be said. When it was thought that fleas were confined, as parasites, to warm-blooded mammals and birds, evolutionists were inclined to say that the parasites could not have appeared before their hosts. The discovery of a flea on a reptile opened the vista of possibly enormous antiquity stretching back to Permian or Carboniferous ages. The fossil record is most meagre. If we reject as too doubtful the supposed remains of a flea from the lower Oligocene strata at Aix in Provence, only one undoubted fossil has been discovered. Nor does it seem certain that fleas are entirely restricted to preying on vertebrates. Dr Dampf introduced a number of common bird-fleas (Ceratophyllus gallinæ) of both sexes to some hairy caterpillars. He observed that several of the fleas buried their heads in the hairy covering of the larvæ and remained some time in the attitude of sucking blood. While this was going on the victims made violent demonstrations of annoyance and discomfort. He also observed that a naked caterpillar was not attacked.Mr Boden has also recorded how he found in a seed-warehouse some peas that were being eaten by two species of Lepidopterous larvæ. On bringing these home and keeping them in a jar, he found among them some small larvæ which ultimatelyturned into fleas, probablyPulex irritans. These fleas, being confined without other food, were observed to prey on the Lepidopterous larvæ and to feed freely on their juices. The larvæ which were attacked by fleas pined and died. The fluid from the stomach of the fleas when they were crushed was transparent and not red like vertebrate blood which often exudes when a mammalian flea is pinched and cracked open.A French entomologist has also reported that the numerous fleas which swarm in the dwelling-houses of Corsica, for want of other nourishment turn their attention to flies that may be incapable of flight.The only fossil remains of a flea that have, so far, been discovered are a single insect in a piece of Baltic amber of Oligocene age. Many organic remains have been preserved in this fashion, but this is the first mammalian parasite that has been found. The flea is admirably protected by its semi-transparent surroundings, and the most minute details of structure, the arrangement of bristles on the body, and the number of segments to the labial palpi can be discerned. This unique object is in the collection of Professor Klebs. The first point to note is that a flea of this antiquity hardly differs from the existing insects of the present day. It has been referred to an existing genus (Palæopsylla) of which there areat least four species living. Three of these are parasites of the mole, and the fourth is found on shrews. There is good reason to suppose that the host of the fossil was some insectivorous mammal. The early specialisation of fleas is strikingly illustrated. This insect is already adapted for life on some warm-blooded animal. It has a thoracic comb, and its mouth-parts are in all respects like those of a modern flea. It belongs to a genus which is still commonly distributed over Europe. When we consider how remote are the chances that a mammalian flea should first get embedded in amber and should, subsequently, be detected and described by a naturalist, we may well understand that the owner of the fossil asked, though without success, £1200 for it.The ordinary person regards fleas as a subject for humour of an obvious and familiar kind. The utilitarian despises a man who can cheerfully spend his time in collecting fleas. Yet it seems probable that a study of their forms and habits may be of immediate benefit to the human race. The discovery that fleas are connected with the spread of plague is an instance of apparently unprofitable scientific labour proving of direct advantage to mankind. An accurate knowledge of the structure and habits of fleas is now seen to be of importance to all who are engaged in fighting one of the most dreaded infectious diseases. When plague breaks out men of science now at onceturn their attention to the fleas. This is likely to prove more directly efficacious than the mediæval custom of marking the house with a red cross and inscribing the legend, “God have mercy on us.”

c1

INTRODUCTORY

Fleasform a group of insects that have, until recently, been little studied by zoologists. We call them insects because they are jointed animals, or Arthropods, with three pairs of legs in the adult condition. The reader will best understand the position which fleas occupy in the general classification of animals by remembering that the arthropods, or jointed animals, are one of a dozen subkingdoms, or phyla, to which the various members of the great animal kingdom have been assigned. There is good ground for believing that all the animals included in each phylum trace their ancestry back to a common primitive form which lived in more or less remote ages. Besides (1)Insects, the arthropods, or jointed animals, include (2)Crustaceans, such as crabs, lobsters, shrimps, wood-lice, water-fleas and barnacles; (3)Myriapods, such as centipedes and millipedes; and (4)Arachnids, such as spiders, scorpions, mites and ticks. To all these varied forms of animal life fleas, and other insects, are therefore more or less nearly related.

The animals belonging to this large and important collection, which compose the arthropod phylum, have certain common characteristic features. We find a body made up of a series of more or less completely similar segments placed one behind the other. In this they resemble certain worms which are far less highly organised. The body is elongated, symmetrical on either side, and the mouth and anus are at opposite ends. There is, however, an important advance on the segmented worms. Each typical segment carries a pair of appendages which are very different from the foot-stumps that are found on certain worms. These appendages of arthropods are divisible into distinct limb-segments, separated from one another by moveable joints, and acted upon by special muscles.

The common ancestor of all the various arthropods which are found living on the earth to-day, was probably composed of a series of segments each very similar to the last and each bearing a pair of very similar appendages. In the course of ages, these appendages have been astoundingly modified in form and in function. So it happens that we find in the arthropods of the present day pairs of antennæ, of mandibles and other mouth-parts, of pincers, of legs, of swimming-feet and of tail pieces which on close examination can all be traced back to a common structure. The body-segments, also, have beenstrangely fused together and modified. All that has been so far said applies equally to fleas and to other insects.

It is of great interest, when one comes to make a minute study of the form and external structure of a flea, to try and trace the modifications that must have taken place in the course of descent from the ancestral arthropod; but the relationship of fleas to other insects living at the present day is of more immediate concern. Insects are highly specialized arthropods and fleas are highly specialized insects. This means that they have become vastly modified from the primitive ancestral type and fitted thereby for a life among certain defined and peculiar surroundings.

It will be unnecessary to remind the reader who knows anything of zoology or of botany that all classification is now based on descent. Since naturalists have abandoned a belief in the special creation of the various species of animals now living on the earth and have conclusively shown that they have arisen by descent and modification from other forms, the problem is to reconstruct a vast genealogical tree. What then were the ancestors of the fleas and to what other insects, in consequence, do they appear to be related?

It is probable that the ancestors of the fleas were winged insects, and that the organs of flight weregradually lost, as they became useless, when a partially parasitic life was adopted. At one time entomologists regarded fleas as wingless flies and placed them in the order Diptera. Certain supposed scaly plates on their bodies were regarded as the atrophied relics of wings. It is, however, more than doubtful whether this view is correct; and all modern entomologists who have given any special study to fleas are agreed that they are sufficiently unlike any other living insects to deserve a place in an order by themselves. To this order the nameSiphonapterahas been given: which means that the insects comprised in it are provided with sucking mouths and are destitute of wings. Another name for the order is Aphaniptera, but this is gradually falling into disuse. Linnæus (1758) only mentions two species of flea: the human flea which he appropriately namedPulex irritans, and the chigoe of hot countries which he calledPulex penetrans, from the habit which the female has of burrowing under the skin of her victims. At the time of writing, about 460 species of flea have been described and named; but some of the names are doubtless synonymous, and the actual number of separable species that have been discovered is somewhere about four hundred. The vast majority of these have been described within the last few years, which shows what can be done when attention is turned to any neglected group of animals. Therecan be no doubt that many undiscovered species still remain, and will now, in due course, be collected, described and named.

The position which should be assigned to the order Siphonaptera in the general scheme of insect classification is a question on which the most learned modern entomologists have disputed with considerable vigour. Some see the nearest relatives among the beetles, others among the flies. The majority, as we shall see later on, would place them near the Diptera: but since no convincing arguments have been produced on either side it may be wisest to regard the question as still at present unsolved.

Fleas belong to one of the groups of insects which go through a complete metamorphosis. Their life-history consequently falls into four divisions: egg, larva, pupa and imago. If the climate permits, the female flea lays her eggs all the year round, and from one to five are dropped at a time. Unlike those of many other parasites they are never attached to the hairs of the hosts, but appear to be deposited indiscriminately on the floors of houses or in the nests and sleeping places of their hosts. The eggs generally hatch in a few days, and a minute, white, wormlike larva emerges (Fig. 1). The larvæ, of some, and possibly of all, fleas are provided with a wonderful adaptation in the shape of an egg-breaker or hatching-spine. This is a thin plate, like the edge of a knife, where the point of the head comes in contact with the shell. The movements of the prisoner make a slight split in the egg-shell, which then bursts asunder. This organ has vanished in later larval life, and it is probably lost after the first moult. The larva is legless and has thirteen segments. It grows rapidly, and, as it grows, moults its skin several times. It is provided with mouth-parts adapted for biting, and eats any decaying organic refuse. The larvæ may be reared on the sweepings of an ordinary room or the dirty scurf which collects at the bottom of old birds’ nests. It is hardly necessary to add that the mother takes no interest whatever in the larvæ and that the belief that she feeds them on dried blood is not based on any sound foundations.

i6

Fig. 1. The larva of a flea. The body consists of thirteen segments and is legless. On the fore part of the head are the antennæ and on the upper part of the head is shown the knife-like edge of the egg-breaker. The mouth-parts are adapted for biting. On the last segment of the body are the two caudal stylets.

Fig. 1. The larva of a flea. The body consists of thirteen segments and is legless. On the fore part of the head are the antennæ and on the upper part of the head is shown the knife-like edge of the egg-breaker. The mouth-parts are adapted for biting. On the last segment of the body are the two caudal stylets.

Fig. 1. The larva of a flea. The body consists of thirteen segments and is legless. On the fore part of the head are the antennæ and on the upper part of the head is shown the knife-like edge of the egg-breaker. The mouth-parts are adapted for biting. On the last segment of the body are the two caudal stylets.

The larval stage lasts some days, and the animal spins a small cocoon before pupating. In the course of a few more days, the time probably depending on the weather, the perfect flea emerges. The larvæ generally live in places where the perfect insects will have an opportunity of finding a host as soon as they leave the pupal envelope. The nests of their hosts where the young are being reared are always favourite places. It seems possible that the comparative immunity from fleas which hoofed mammals or Ungulates enjoy may be due to the fact that the young beast follows its mother from the time of birth instead of passing its early life helpless in a nest.

Observations made on the development of the dog-flea (Ctenocephalus canis) in India show that eggs laid on October 17 hatched on October 19. The larva spun its cocoon on October 25 and the mature flea emerged on November 2. In Northern Europe the human flea takes about four weeks in summer and six weeks in winter to pass through its metamorphosis.

Unlike many parasitic insects, fleas do not constantly pass their time upon the bodies of their victims. The greater part of their life is probably spent on the ground, in the house, or nest, of the mammal or bird which serves them with blood. In this respect there is considerable difference in the habits of different species of flea. Some attach themselves to an animal and actually burrow into the skin. These are the most parasitic species. Some only come to feed and leave to lay their eggs. Many probably do not suck blood more than once in their lives.

An animal which harbours fleas and which nourishes the adult insect with blood is called ahost. No fleas are more than what is called temporary parasites; which means that they pass but a portion of their lives on their hosts and frequently take occasion to hop on and off. All fleas, apparently, go from host to host. The labours of diligent collectors have proved that the greatmajority of mammals and birds have fleas. As a general rule, it is true to say that certain species of flea are associated with certain species of host. Thus man is the true host ofPulex irritans; the cat family are the true hosts of the cat-flea (Ctenocephalus felis); and the dog family are the true hosts of the dog-flea (Ctenocephalus canis). But the human flea is sometimes found on cats and dogs, and cat and dog-fleas occasionally bite human beings; and cat-fleas are found on dogs and dog-fleas are found on cats. All fleas, so far as we know, may occasionally pass from one species of host to another; but they do not, for the most part, seem to flourish in unaccustomed quarters. Some fleas are more catholic in their tastes than others. Some seem to be very strictly confined to one host, and even when starving only suck strange blood under protest. There is a species of flea that has only (except by accident) been found on the long-tailed field-mouse and another that has only been found on the hedgehog. Other fleas are commonly found on two absolutely distinct animals; a good instance of this is the human flea which, at all events in certain parts of England, is a regular parasite of the badger.

As distinguished from true or natural hosts one must separate what may be termed casual or accidental hosts. All animals which come in contact with one another, or which live in close proximity,may exchange fleas. So even bird-fleas may be collected from mammals and typically mammalian fleas from birds. In this fashion puzzles may arise which tax the ingenuity of the collector to solve. Bird-fleas are sometimes found on bats, and this may be obviously attributed to the bats having inhabited a hole which was tenanted by starlings or an old loft infested with the fleas of pigeons. All beasts of prey are sometimes found to harbour the fleas of animals they have devoured. Rabbits’ fleas are found on wild-cats; hedgehogs’ fleas on foxes; mice fleas on weasels; and fleas characteristic of small birds on stoats. So also in the case of mice, rats and voles with holes and runs in the same hedgerow, the parasites usually peculiar to one are not uncommonly found on the others. It is sometimes difficult to determine the true host of a flea.

Much more puzzling to explain are the reasons which confine a flea to a certain host and which cause closely allied hosts to have different fleas. The fleas from the house-martin and the sand-martin are quite different; those from the domestic fowl and the domestic pigeon are distinct species. The causes which have affected the evolution of the various forms of flea are too obscure to enable anyone at the present day to offer any satisfactory explanation.

Speaking generally, the fleas found on birds have points in common, and they probably form a naturalgroup to themselves. What may be called true bird-fleas have been collected from almost all European birds. An unwieldy genus (Ceratophyllus) comprises many species of different flea. Some species are very abundant and infest the nests of many different birds. Others are extremely rare. One of these rarities (C. vagabundus) is found in the nests of puffins and other sea-birds. Another has been collected on antarctic petrels. Penguins have a special genus of flea to themselves. A specimen, unique at one time (Ceratophyllus borealis), in Mr N. C. Rothschild’s collection was obtained from the gannet. It has now been found on rock-pipits in the Shetland Islands.

Two very rare fleas (C. farreniand C.rothschildi) are found in the nests of house-martins; yet the nests of these birds are infested with common species besides. A plague flea (Xenopsylla) has been found on an African swift.

Forty-six different species of flea have been found in the British Islands, but many of these are extremely scarce.

We know too little about the geographical distribution of fleas to lay down many accurate generalities. When a great deal more material has been collected and studied, it may be possible to show that certain groups are associated with certain regions of the earth or certain orders of animals. To some extent this is already seen to be the case. The fleasindigenous to the New World are distantly related to those of the Old World. Broadly speaking the geographical distribution of the parasite must follow that of the host. But sometimes the parasite is impatient of cold and cannot follow the host out of the tropics. The chigoes and their allies are fleas of hot countries. Different kinds of bats are found from the tropics to the Arctic circle, but the same bat-fleas are not found everywhere.

When a flea has a cosmopolitan range it is probable that it has travelled over the world in company with its host.

Monkeys have no fleas. This is an assertion that is commonly received with surprise and incredulity. Occasionally a gorilla or a chimpanzee may get a chigoe in its toe. And monkeys in zoological gardens or menageries are possibly exposed to the danger of catching an occasional human flea from the people who crowd round their cages. These are remote contingencies which may happen to anyone. Healthy wild monkeys are much too clean and active to harbour fleas. When they search one another’s fur in a fashion that must be familiar to most persons, they are clearing their coats of particles of scurf or of similar scraps of dirt and not of fleas. So, speaking generally, it may be said that no fleas have been found truly parasitic on monkeys.

Bats have fleas, but not in great abundance. Allbat-fleas are rare on their hosts and extremely difficult to find and collect. The same species are not found on fruit-bats and the ordinary smaller insect-eating bats. The geographical distribution of some bat-fleas is puzzling. For instance, one species (Ischnopsylla unipectinata) is found on the greater horse-shoe bat in Europe; but it is, apparently, not found on the same bat in the British Islands. In Somaliland and in India it is found on other bats.

With certain exceptions, Ungulates are remarkably free from fleas. This great order of mammals includes a variety of hoofed animals: oxen, sheep, goats, deer, pigs, camels, giraffes and antelopes.

The only true fleas found on these are two species of the genusVermipsylla, which resemble the chigoes in so far that the pregnant females burrow into the host and expand there. One species has been found on camels and horses in Transcaucasia; another on roe-deer in Northern China. The female of the last is often found ensconced on the inside of the nostrils of the deer. Of course chigoes may attack domestic Ungulates of all kinds; but no other members of the familyPulicidæor typical fleas except those two above mentioned have been found on hoofed mammals.

Insectivora such as moles, shrews and hedgehogs are the hosts of a great variety of species. The same thing may be said of the Rodents, which includeporcupines, squirrels, rats, mice and a vast number of other small mammals whose geographical distribution includes almost the whole of the habitable globe. Probably more different species of fleas have been collected from Insectivora and Rodents than from all the other orders of mammals grouped together.

The Carnivora, excluding the Pinnepedia, or seals, sea-lions and walruses, harbour numerous species.

Among the Edentata a very remarkable and highly specialised genus of fleas is parasitic on armadilloes in South America. This genus (Malacopsylla) consists of two species only, which are confined to South America and are found on the armadilloes and on carnivorous animals which probably have preyed on them. The thorax of these fleas is much reduced and very small in size. Their piercing organs are slender and weak, but they possess enormous spines on the legs with which they hold on to their hosts. These two South American fleas (M. grossiventrisandM. androcli) will be referred to again later as striking examples of fleas with strongly developed legs and weakly constructed mouth-parts. The contrary combination of powerful mouths and degenerate legs is also found in other groups of fleas, as will be seen in the chapter on the chigoes.

The Marsupials of Australia and South America have special fleas which were probably associated with this strange order of pouched mammals beforethey became divided into the American and Australian groups. Fleas have been collected on the spiny ant-eater (Echidna) which belongs to the lowest order of Monotremes or egg-laying mammals.

On almost every form of bird, including the most aquatic kinds, fleas of various species have been obtained.

Only one instance has been recorded of a flea occurring on a reptile. A female of one of the species of burrowing chigoes (Echidnophaga ambulans) from Australia was collected by Dr Woodward from the Brown Snake (Diemenia superciliosa). This reptile, which is well known in Australia, belongs to a sub-family that contains some of the most deadly poisonous snakes and is allied to the cobras. The Brown Snake is a terrestrial snake, and one must regard the presence of the flea on such a host as a rare and chance occurrence. The snake was captured at Herdman’s Lake, near Perth in West Australia. The same species of flea has also been obtained from the phalangers (Trichosurus) which live in the tops of the Australian gum-trees; from the little terrestrial and nocturnal rat kangaroos (Bettongia); and from the banded ant-eater (Myrmecobius), another Australian Marsupial. It is possible that the flea moved from some small mammal which was being devoured by the snake and managed to fix itself between the scaly plates of the reptile.

When fleas are hatched in a nest they have no choice but to attach themselves to the young mammals or birds. But even in that case they frequently leave their hosts and do not for very long remain stationary. Moreover, when a host dies and becomes cold the fleas invariably leave their quarters, which explains how it may happen that Carnivora get infested with the fleas of their prey. This change of hosts which is always occurring makes it impossible to draw conclusions from material collected in zoological gardens where many animals are herded together. In menageries, too, the normal conditions of breeding are absent. A German naturalist collected 2036 fleas from theatres, concert-halls, ball-rooms, schools and barracks in the grand-duchy of Baden and found that more than fifty per cent. were dog-fleas (Ctenocephalus canis). What the proportion may be in other parts of Europe we have no materials from which to form a judgment. In zoological gardens cat-fleas (Ct. felis) are generally numerous in most of the cages.

It is, of course, well known to every zoologist that species are not fixed or constant and that various forms of mammal or of bird tend to show geographical variations. When a long series of skins are laid out on a table and carefully examined it is seldom that those from the west of any great region cannot be picked out and distinguished from those obtained inthe east. So we also get northern and southern forms of the same species varying slightly. These variations are perceptible in many forms of insects, and zoologists now describe these local races as subspecies and designate them with trinomials. No one, however, knows enough as yet about all the various forms which are assumed by fleas to attempt, except in a few instances, to do so in the case of these animals.

The study of Siphonaptera is still quite in its infancy. We know little or nothing of the minute geographical variation of fleas. That there is such a thing can already be seen in a few species. In the meantime the study of variation must be postponed until collectors have amassed a more plentiful amount of material; and it is best to treat all forms which are to all appearance constantly different as being specifically distinct until more is known about variation.

Any classification of fleas that may now be attempted can only be tentative. It will be enough for present purposes if the reader will remember that the Order Siphonaptera can be divided into three groups or families: (1) the chigoes and their allies, which are the most parasitic fleas (Sarcopsyllidæ); (2) the typical fleas to which the majority of species belong (Pulicidæ); and (3) the bat-fleas (Ceratopsyllidæ), which have certain peculiarities that will be described in a later chapter.

Of the antiquity of fleas, and of the period in geological history when the order made its appearance, little can be said. When it was thought that fleas were confined, as parasites, to warm-blooded mammals and birds, evolutionists were inclined to say that the parasites could not have appeared before their hosts. The discovery of a flea on a reptile opened the vista of possibly enormous antiquity stretching back to Permian or Carboniferous ages. The fossil record is most meagre. If we reject as too doubtful the supposed remains of a flea from the lower Oligocene strata at Aix in Provence, only one undoubted fossil has been discovered. Nor does it seem certain that fleas are entirely restricted to preying on vertebrates. Dr Dampf introduced a number of common bird-fleas (Ceratophyllus gallinæ) of both sexes to some hairy caterpillars. He observed that several of the fleas buried their heads in the hairy covering of the larvæ and remained some time in the attitude of sucking blood. While this was going on the victims made violent demonstrations of annoyance and discomfort. He also observed that a naked caterpillar was not attacked.

Mr Boden has also recorded how he found in a seed-warehouse some peas that were being eaten by two species of Lepidopterous larvæ. On bringing these home and keeping them in a jar, he found among them some small larvæ which ultimatelyturned into fleas, probablyPulex irritans. These fleas, being confined without other food, were observed to prey on the Lepidopterous larvæ and to feed freely on their juices. The larvæ which were attacked by fleas pined and died. The fluid from the stomach of the fleas when they were crushed was transparent and not red like vertebrate blood which often exudes when a mammalian flea is pinched and cracked open.

A French entomologist has also reported that the numerous fleas which swarm in the dwelling-houses of Corsica, for want of other nourishment turn their attention to flies that may be incapable of flight.

The only fossil remains of a flea that have, so far, been discovered are a single insect in a piece of Baltic amber of Oligocene age. Many organic remains have been preserved in this fashion, but this is the first mammalian parasite that has been found. The flea is admirably protected by its semi-transparent surroundings, and the most minute details of structure, the arrangement of bristles on the body, and the number of segments to the labial palpi can be discerned. This unique object is in the collection of Professor Klebs. The first point to note is that a flea of this antiquity hardly differs from the existing insects of the present day. It has been referred to an existing genus (Palæopsylla) of which there areat least four species living. Three of these are parasites of the mole, and the fourth is found on shrews. There is good reason to suppose that the host of the fossil was some insectivorous mammal. The early specialisation of fleas is strikingly illustrated. This insect is already adapted for life on some warm-blooded animal. It has a thoracic comb, and its mouth-parts are in all respects like those of a modern flea. It belongs to a genus which is still commonly distributed over Europe. When we consider how remote are the chances that a mammalian flea should first get embedded in amber and should, subsequently, be detected and described by a naturalist, we may well understand that the owner of the fossil asked, though without success, £1200 for it.

The ordinary person regards fleas as a subject for humour of an obvious and familiar kind. The utilitarian despises a man who can cheerfully spend his time in collecting fleas. Yet it seems probable that a study of their forms and habits may be of immediate benefit to the human race. The discovery that fleas are connected with the spread of plague is an instance of apparently unprofitable scientific labour proving of direct advantage to mankind. An accurate knowledge of the structure and habits of fleas is now seen to be of importance to all who are engaged in fighting one of the most dreaded infectious diseases. When plague breaks out men of science now at onceturn their attention to the fleas. This is likely to prove more directly efficacious than the mediæval custom of marking the house with a red cross and inscribing the legend, “God have mercy on us.”

c2CHAPTER IITHE EXTERNAL STRUCTURE OF A FLEAIncomparing the structure of a flea with that of a man, or any other of the higher animals, it is of the utmost importance to understand that the one has an internal and the other an external skeleton. In either case the skeleton serves as an attachment for the muscles by which the animal moves itself. Everyone is familiar with the external skeleton of a lobster and can see for himself how the muscles are attached. The structure of a flea, though so much smaller, is somewhat similar, except that the skeleton is composed of a horny substance known aschitininstead of being calcified. The chitinous cuticle entirely covers the flea, but it varies in hardness and thickness on different parts of the body. The epidermis, or true skin, lies immediately beneath. On those parts of the body which are to the rear of the head the chitin forms a series of plates or shields which overlap one another somewhat like thetiles of a house. The segmented structure of a flea is there most clearly seen; this we may suppose is an inheritance from the segmented worms.The chitin which forms the external skeleton of a flea is secreted by an outer layer of cells on the insect’s body. The deposit being thin at the joints, and thick on the plates, which serve for protection, the flea is encased in a suit of flexible armour. It is made of a fairly solid and dense substance, but, owing to the absence of carbonate and phosphate of lime, is much lighter than the familiar external skeleton of the lobster. Chitin is a very peculiar and durable substance which resists boiling in acids or alkalies. It is a structureless substance, in the sense that it does not consist of cells. Though horny in appearance it is, of course, in no sense true horn like that of the nails, hoofs, claws, and horns of vertebrates.The different species of flea vary considerably in size. Some are smaller than the familiar human flea. Others are much larger. A very large flea (Hystrichopsylla talpæ) is that found on the mole. The largest known flea (Dolichopsyllus stylosus) is found on small rodents in the United States where, as we know, all things are on a bigger scale than in the Old World. It is seven millimetres long.The colour of the horny integument varies from a pale or light yellow to a ruddy or dark brown.It is plentifully sprinkled over with spines, bristles, or hairs, directed backwards so as not to impede progress. The presence or absence, the arrangement on the body, and the size of these serve, along with other features, to distinguish different species. They seem materially to help a flea in those wriggles to escape with which we are all familiar. The bristles are not always the same in size and arrangement in the two sexes of the same species of flea. As a rule the males are more bristly than the females.These appendages of the flea’s integument are called by various writers either spines, bristles, or hairs. There is, however, no real distinction in the structure or nature of the appendages, and it is a question of degree which name is most appropriate.In distinguishing species, very little reliance can be placed on the colour of the flea. An insect newly emerged from the pupa is always lighter in colour; and the difference between the appearance of an empty stomach and a stomach gorged with red blood is surprising.The general external appearance of a typical flea belonging to the main familyPulicidæis fairly familiar to most persons. This is well seen in the figure of the oriental rat-flea. The body is compressed or flattened from side to side, and this is a feature which is extremely rare among insects. It doubtlessenables the animal to glide with greater facility through the hairs of its host. Like other insects, a flea is readily divided into a head, a thorax, and an abdomen. The head is rounded on the top and front and shows no obvious trace of segmentation; but what is known of the development of other insects leads one to think that it must properly be regarded as a number of segments closely fused together. On the under side of the front part of the head is a beak or proboscis for piercing and sucking, composed of the mouth-parts, whose structure is worthy of minute study. It will be best to examine them in detail in a subsequent chapter.Some fleas have eyes, others have none. The common mouse-flea (Leptopsylla musculi) is blind. The bat-fleas are also destitute of eyes. The nocturnal habits of their hosts would render eyes of little or no use. If eyes are present they are large and placed on either side of the head. Each is a simple eye or ocellus; the compound eyes, divided into a great number of hexagonal facets, which are characteristic of many insects, are never found in fleas.Nothing is known about the flea’s powers of vision, but there is no reason to suppose that they are at all acute. The eyes are marked with pigment. Ocelli appear to be primitive types of insect eye which are, perhaps, an inheritance from a wormlikeancestor. Presumably all the fleas of long ago had eyes and those that are now blind have lost their organs of sight from disuse. In their simplest condition, the eyes of the lower invertebrates only enable the creature, so far as one can judge, to distinguish light from darkness. Entomologists believe that the power of vision of ocelli is probably confined to very near objects and that this simple form of eye is more useful in dark places than the compound eyes. There is no reason for believing that fleas can distinguish colours or can discern any object which is more than a few inches away. It is enough for their purpose to perceive from which point light comes upon them and to make all despatch to escape in the opposite direction.In blind fleas there is often a spine where the eye should be. In one species the spine is rudimentary and there is some black pigment beneath it. It is not impossible that this is the vestige of a once functional eye. In one genus, however, the eye and the spine are both present. Of the fleas belonging to this genus one species is South American and the other European. The latter (Typhloceras poppei) is confined to the long-tailed field-mouse.The organs by which fleas keep in touch with the outward world, and with other fleas, are their antennæ. All fleas have antennæ; but unlike those of a moth, a beetle, or a grasshopper, each fits neatlyinto a groove at the side of the head and can be protruded when desired. This is another adaptation to enable the insect to creep swiftly through a forest of hairs.i26Fig. 2. Showing a type of (a)genaland (b)thoraciccombs of a flea, on the under part of the head and on the thorax respectively. Analagous combs are found in several other parasitic insects and on the abdominal segments of certain fleas.The combs which are found on the heads of many fleas are organs of exceptional interest (Fig. 2). They are toothed and horny appendages, which are connected with parasitic habits, for somewhat similar combs are found on several unrelated groups of parasitic insects, as, for instance, on parasitic beetles (Platypsyllus) found on the beaver, on insects allied to bed-bugs (Polyctenes) found on bats, and on wingless flies (Nycteribia) which infest Egyptian and South American bats. The majority of thePulicidæhave one or more combs with comparatively long teeth. These combs reach their maximumdevelopment in the bat-fleas which have no less than eight. Some Australian and South American fleas (Stephanocircus) have a helmet-like comb extending all round their heads. These combs are by some supposed to be of service in holding on to the hairs of the host; and, if one may judge from experiments made on live fleas in cotton wool, they are also used in moving forwards through the fur.All the chigoes (Sarcopsyllidæ) have a large triangular post-oral process which is more or less curved and probably prevents the flea slipping back as it pushes forward. Bat-fleas (Ceratopsyllidæ) have lobes or flaps placed two on each side of the head, which may possibly serve an identical purpose, but whether they do so is not known.These combs may be divided into three groups according to the part of the flea’s body on which they are found. Those found on the head are called genal combs and take the form shown inFig. 2. There are also combs found on the thorax. The fossil flea (Palæopsylla klebsi) described in the previous chapter has one of these thoracic combs. A certain number of fleas also have combs on the abdominal segments.There are really two types of toothed organs on fleas to both of which the name of comb may be given. One is composed of a sheet of chitin with a number of slits and teeth and resembles a true comb.The other consists merely of a number of highly chitinised bristles arranged in a row. They probably both serve the same function. Apart from their use as organs to assist movement onwards, they may also serve ashair-tightjoints and protect the flea from the inconvenience of getting the tips of the host’s fur into the joints of its horny armour.The size of the head compared to the thorax and abdomen varies considerably in different species. Some fleas have what may be called by comparison large heads and others very small ones.A small head is never found in a flea with powerful mouth-parts. The head being the bearer of piercing and sucking organs, which require strong muscles, there must be room not only for the organs but for their extensors and retractors.There are normally three rows of bristles on a flea’s head which divide the head into four sections. It is possible that these correspond to the four segments of the ancestral insect which are now fused together.The head of a flea is closely applied by the whole of its back surface to the body and that slender and conspicuous neck which is characteristic of the Diptera, or flies, is not to be found in any fleas. For this reason a flea cannot turn its head in any direction without at the same time following it round with its body.The thorax of a flea consists of three segments called respectively the prothorax, mesothorax and metathorax. The chitinous external skeleton which covers each of these three segments is primarily a hoop but each hoop is further subdivided into a number of complicated plates. Attached to the thorax are the three pairs of legs which are characteristic of all adult insects. The hind pair are very much the strongest (Fig. 3). They are the organs of hopping. It has often been pointed out that if men had the leaping powers of some fleas they would bound with ease backwards and forwards over the cross on the top of St. Paul’s Cathedral. Each leg consists of four segments beautifully articulated and plentifully supplied with bristles. At the end comes the foot with five very short segments. The last segment is provided with a pair of more or less formidable claws. Fleas use their legs for leaping, for running, and for clinging to their hosts. They also use their mouth-parts for the last purpose and it is worthy of note, as we shall see later on, that in those fleas in which the mouth-parts are shortest and weakest the legs are most liberally supplied with bristles and possess the stoutest claws. The legs of a flea are unique in the insect world owing to the enormous development of the segment nearest the body called the coxa. Most leaping insects rely for their activity on the muscles of the lower joints. In a grasshopper it is the third joint from the body (femur) which is so immensely enlarged. The three pairs of legs are each attached to a different thoracic segment.i30Fig. 3. The hind leg of a flea. The segment or joint nearest the body is thecoxawhich is unusually developed. Next come the smalltrochanterand the largerfemur. Thetibiawhich is long and slender follows. Then come the fivetarsiwith the sixth and ultimate segment provided with claws.When fleas walk, they are so to speak plantigrades walking on the sole of the foot, and all the tarsal or foot joints are applied to the surface of the ground. The claws serve as grips so as to make the most of any unevenness; and thus the insect drags itself along with surprising rapidity when it moves through the hairy coat of a mammal. But on an open surface fleas are not really rapid movers compared with many other insects.The two claws on the end of each ultimate foot segment are freely moveable and are in fact highly modified bristles or setæ.In all fleas one of the plates of the metathorax (or hindmost thoracic segment) called the epimeron, is large and prolonged towards the rear. It invariably bears a stigma. The epimeron is placed laterally to the first abdominal tergite. The older naturalists jumped to the conclusion that this was the remains of a wing. The best judges have, however, formed a decided opinion that no trace of the relic of a flying organ can be detected on the thorax of a flea. Heymons, a German entomologist, has also failed to detect any sign in dissections which he has made of the larvæ and the pupæ.The epimeron is in fact neither a scale nor awing but a portion of the thorax present in all insects. It is of no special service to the flea except as a portion of the thoracic armature which covers the body.The larva of a flea has no legs; the adult insect has six. A study of other embryo insects shows that the ancestors of insects had many legs. It is an interesting problem why insects lost the legs on their abdomens, why legs should now invariably be restricted to the thorax, and why there should never be more than three pairs. In the earliest known insects which lived on the earth, before winged forms were evolved, the number of legs was already six. But our knowledge of fleas is too small to attempt, at present, to trace their exact line of ancestral descent.The abdomen of a flea consists of ten segments. The horny plates which cover the dorsal side are called tergites; those on the ventral side sternites. In fleas, as in all holometabolous insects, that is those which pass through a complete metamorphosis, the sternite of the first abdominal segment is suppressed and has completely disappeared. The tergite which covers the dorsal part of the first abdominal segment nearest to the thorax is, however, always present.The ultimate segments of the male and female flea are modified for reproductive purposes and of these segments more must be said later.Having now given a rough outline of the external skeleton of a flea, it only remains to say something about the muscular system. Attached to the inside of the chitinous armature are an enormous number of muscles, whitish and almost transparent. They act as extensors, retractors, flexors, elevators and depressors. The joints and hinges of the skeleton allow of considerable, but not perfect, freedom. The muscles of locomotion are partly in the thorax and partly in the several joints of the legs. Our knowledge of the muscular system of fleas is very imperfect. But, as in other insects, the general arrangement of the muscles is based on the segmented structure of the body.For the reader who can accurately picture to himself the external structure of a flea and of the typical insects belonging to other orders, a few words may be said on the probable ancestry of fleas and their relationship to other living insects. This vexed and much debated question is still, as the older naturalists would have said,tremendum mysterium. Very little light has yet been thrown upon it, and the most divergent views have been expressed by learned and competent entomologists. A historic survey of the various opinions that have been held since the days of Linnæus would fill many pages; but a short summary of the different orders to which fleas have been referred by different zoologists will suffice.The older authors, Linnæus, Geoffroy, Cuvier and Duméril, and Gervais placed them among theApterabecause they were wingless. Kircher regarded them asOrthoptera, an order which includes grasshoppers and crickets; but he has had few followers. By Fabricius and by Illiger they were treated asHemipteraor bugs. Lameere, a Belgian, has recently expressed a decided view that fleas are really a family ofColeopteraor beetles. Those who have held the once orthodox opinion that they belonged to theDipteraor flies are Roesel, Oken, Straus-Dürkheim, Burmeister, Newman, Walker, von Siebold and Wagner.The structure of an adult flea, however, differs from that of an adult fly in the following noteworthy respects: the mouth-parts are differently constructed, the head of the flea is closely joined to its thorax, the three divisions of the thorax are not joined and fused, the flea is wingless, the eyes of fleas are simple ocelli, and there are differences of lesser importance in the stigmata, which give access to the tracheal system by which all insects breathe.The number of those who have regarded fleas as belonging to a distinct order of insects is considerable: they are Lamarck, De Geer, Latreille, Kirby and Spence, MacLeay, Leach, Dugès, Bouché, van der Hoeven, Westwood, Landois, Brauer, Kraepelin,and Taschenberg. Modern opinion is all but unanimous on this point.There remains, however, a second question. Even if it be agreed that there must be a distinct order forSuctoria, Aphaniptera, Siphonaptera, or fleas; where ought that order to be placed? In which other order of insects must we look for the nearest relations of fleas? For a time after the acceptance of the fact that insect forms have been evolved, and not separately created, the ancestors of fleas were searched for among some species of fly.Then Kraepelin rejected the view that flies were as closely related to fleas as most entomologists thought and his followers could only find points of difference and no points of resemblance. Dahl (1899), a German, then took up the cudgels for the fly theory. Dahl pointed out the resemblance between fleas and a group of flies calledPhoridæalso parasitic on warm-blooded animals. During the ensuing years the debate was resumed afresh with much liveliness and sometimes with a little acrimony.The fleas were placed by MacLeay and by Balbiani between theDipteraandHemiptera; by Leach between theHemipteraandLepidoptera; by Dugès between theHymenopteraandDiptera; by Brauer between theDipteraandColeoptera. Handlirsch thinks that fleas have no connection atall with beetles and Gross can find no signs of relationship with eitherColeopteraorDiptera.Embryology and the study of larval forms have thrown so much light on the ancestry of many animals, that it was hoped that a microscopic examination of the larvæ of fleas, in various stages of development, would produce some facts of importance. In this hope entomologists have, to a great extent, been disappointed. There seems to be much similarity between the embryos of beetles, moths, flies, wasps and fleas. Those who have dwelt on the likeness of the larval flea to the maggot of a fly seem to forget that the resemblance to an embryo beetle is nearly as strong.The young larva of the flea is very transparent and the digestive canal, heart and nervous system are easily recognised. The egg-shell breaker is an interesting example of the development of a temporary larval structure and it is the only known instance of such a structure in an insect. There are no traces of eyes. The antennæ are three-jointed. They are rather long and slender, being about one-third as long as the head. The head is well-developed and the larva has no feet.The biting mandibles are broad and triangular. Compared with those of other larvæ they are said to be more like the mandibles of coleopterous than of dipterous larvæ. The maxillæ, or second pair ofjaws, are somewhat reduced and rudimentary. The absence of eyes and of legs are points of similarity between the larvæ of fleas and flies. The maggot of a fly has also two pairs of jaws, and a pair of antennæ.At the tail end of the larval flea’s abdomen are two small projections called caudal stylets (Fig. 1). They are strong, recurved, chitinous, structures which prop up the body of the larva when it creeps and wriggles. There are similar props in the larvæ of certain beetles and no exactly similar organs are known in dipterous larvæ. But caudal stylets are of small taxonomic importance.In one respect the mature flea is certainly nearer to a beetle than to a fly: the three joints of the thorax are free as in a beetle and not fused as in a fly; but when one studies the mouth-parts, the true view seems to be that the mouth-parts of a flea are equally unlike those of a fly and those of a beetle. Such being the present state of our knowledge, one must wait for fresh light to be thrown on the matter by further researches. It seems unlikely that the immediate future will produce a solution of the problem.

c2

THE EXTERNAL STRUCTURE OF A FLEA

Incomparing the structure of a flea with that of a man, or any other of the higher animals, it is of the utmost importance to understand that the one has an internal and the other an external skeleton. In either case the skeleton serves as an attachment for the muscles by which the animal moves itself. Everyone is familiar with the external skeleton of a lobster and can see for himself how the muscles are attached. The structure of a flea, though so much smaller, is somewhat similar, except that the skeleton is composed of a horny substance known aschitininstead of being calcified. The chitinous cuticle entirely covers the flea, but it varies in hardness and thickness on different parts of the body. The epidermis, or true skin, lies immediately beneath. On those parts of the body which are to the rear of the head the chitin forms a series of plates or shields which overlap one another somewhat like thetiles of a house. The segmented structure of a flea is there most clearly seen; this we may suppose is an inheritance from the segmented worms.

The chitin which forms the external skeleton of a flea is secreted by an outer layer of cells on the insect’s body. The deposit being thin at the joints, and thick on the plates, which serve for protection, the flea is encased in a suit of flexible armour. It is made of a fairly solid and dense substance, but, owing to the absence of carbonate and phosphate of lime, is much lighter than the familiar external skeleton of the lobster. Chitin is a very peculiar and durable substance which resists boiling in acids or alkalies. It is a structureless substance, in the sense that it does not consist of cells. Though horny in appearance it is, of course, in no sense true horn like that of the nails, hoofs, claws, and horns of vertebrates.

The different species of flea vary considerably in size. Some are smaller than the familiar human flea. Others are much larger. A very large flea (Hystrichopsylla talpæ) is that found on the mole. The largest known flea (Dolichopsyllus stylosus) is found on small rodents in the United States where, as we know, all things are on a bigger scale than in the Old World. It is seven millimetres long.

The colour of the horny integument varies from a pale or light yellow to a ruddy or dark brown.It is plentifully sprinkled over with spines, bristles, or hairs, directed backwards so as not to impede progress. The presence or absence, the arrangement on the body, and the size of these serve, along with other features, to distinguish different species. They seem materially to help a flea in those wriggles to escape with which we are all familiar. The bristles are not always the same in size and arrangement in the two sexes of the same species of flea. As a rule the males are more bristly than the females.

These appendages of the flea’s integument are called by various writers either spines, bristles, or hairs. There is, however, no real distinction in the structure or nature of the appendages, and it is a question of degree which name is most appropriate.

In distinguishing species, very little reliance can be placed on the colour of the flea. An insect newly emerged from the pupa is always lighter in colour; and the difference between the appearance of an empty stomach and a stomach gorged with red blood is surprising.

The general external appearance of a typical flea belonging to the main familyPulicidæis fairly familiar to most persons. This is well seen in the figure of the oriental rat-flea. The body is compressed or flattened from side to side, and this is a feature which is extremely rare among insects. It doubtlessenables the animal to glide with greater facility through the hairs of its host. Like other insects, a flea is readily divided into a head, a thorax, and an abdomen. The head is rounded on the top and front and shows no obvious trace of segmentation; but what is known of the development of other insects leads one to think that it must properly be regarded as a number of segments closely fused together. On the under side of the front part of the head is a beak or proboscis for piercing and sucking, composed of the mouth-parts, whose structure is worthy of minute study. It will be best to examine them in detail in a subsequent chapter.

Some fleas have eyes, others have none. The common mouse-flea (Leptopsylla musculi) is blind. The bat-fleas are also destitute of eyes. The nocturnal habits of their hosts would render eyes of little or no use. If eyes are present they are large and placed on either side of the head. Each is a simple eye or ocellus; the compound eyes, divided into a great number of hexagonal facets, which are characteristic of many insects, are never found in fleas.

Nothing is known about the flea’s powers of vision, but there is no reason to suppose that they are at all acute. The eyes are marked with pigment. Ocelli appear to be primitive types of insect eye which are, perhaps, an inheritance from a wormlikeancestor. Presumably all the fleas of long ago had eyes and those that are now blind have lost their organs of sight from disuse. In their simplest condition, the eyes of the lower invertebrates only enable the creature, so far as one can judge, to distinguish light from darkness. Entomologists believe that the power of vision of ocelli is probably confined to very near objects and that this simple form of eye is more useful in dark places than the compound eyes. There is no reason for believing that fleas can distinguish colours or can discern any object which is more than a few inches away. It is enough for their purpose to perceive from which point light comes upon them and to make all despatch to escape in the opposite direction.

In blind fleas there is often a spine where the eye should be. In one species the spine is rudimentary and there is some black pigment beneath it. It is not impossible that this is the vestige of a once functional eye. In one genus, however, the eye and the spine are both present. Of the fleas belonging to this genus one species is South American and the other European. The latter (Typhloceras poppei) is confined to the long-tailed field-mouse.

The organs by which fleas keep in touch with the outward world, and with other fleas, are their antennæ. All fleas have antennæ; but unlike those of a moth, a beetle, or a grasshopper, each fits neatlyinto a groove at the side of the head and can be protruded when desired. This is another adaptation to enable the insect to creep swiftly through a forest of hairs.

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Fig. 2. Showing a type of (a)genaland (b)thoraciccombs of a flea, on the under part of the head and on the thorax respectively. Analagous combs are found in several other parasitic insects and on the abdominal segments of certain fleas.

Fig. 2. Showing a type of (a)genaland (b)thoraciccombs of a flea, on the under part of the head and on the thorax respectively. Analagous combs are found in several other parasitic insects and on the abdominal segments of certain fleas.

Fig. 2. Showing a type of (a)genaland (b)thoraciccombs of a flea, on the under part of the head and on the thorax respectively. Analagous combs are found in several other parasitic insects and on the abdominal segments of certain fleas.

The combs which are found on the heads of many fleas are organs of exceptional interest (Fig. 2). They are toothed and horny appendages, which are connected with parasitic habits, for somewhat similar combs are found on several unrelated groups of parasitic insects, as, for instance, on parasitic beetles (Platypsyllus) found on the beaver, on insects allied to bed-bugs (Polyctenes) found on bats, and on wingless flies (Nycteribia) which infest Egyptian and South American bats. The majority of thePulicidæhave one or more combs with comparatively long teeth. These combs reach their maximumdevelopment in the bat-fleas which have no less than eight. Some Australian and South American fleas (Stephanocircus) have a helmet-like comb extending all round their heads. These combs are by some supposed to be of service in holding on to the hairs of the host; and, if one may judge from experiments made on live fleas in cotton wool, they are also used in moving forwards through the fur.

All the chigoes (Sarcopsyllidæ) have a large triangular post-oral process which is more or less curved and probably prevents the flea slipping back as it pushes forward. Bat-fleas (Ceratopsyllidæ) have lobes or flaps placed two on each side of the head, which may possibly serve an identical purpose, but whether they do so is not known.

These combs may be divided into three groups according to the part of the flea’s body on which they are found. Those found on the head are called genal combs and take the form shown inFig. 2. There are also combs found on the thorax. The fossil flea (Palæopsylla klebsi) described in the previous chapter has one of these thoracic combs. A certain number of fleas also have combs on the abdominal segments.

There are really two types of toothed organs on fleas to both of which the name of comb may be given. One is composed of a sheet of chitin with a number of slits and teeth and resembles a true comb.The other consists merely of a number of highly chitinised bristles arranged in a row. They probably both serve the same function. Apart from their use as organs to assist movement onwards, they may also serve ashair-tightjoints and protect the flea from the inconvenience of getting the tips of the host’s fur into the joints of its horny armour.

The size of the head compared to the thorax and abdomen varies considerably in different species. Some fleas have what may be called by comparison large heads and others very small ones.

A small head is never found in a flea with powerful mouth-parts. The head being the bearer of piercing and sucking organs, which require strong muscles, there must be room not only for the organs but for their extensors and retractors.

There are normally three rows of bristles on a flea’s head which divide the head into four sections. It is possible that these correspond to the four segments of the ancestral insect which are now fused together.

The head of a flea is closely applied by the whole of its back surface to the body and that slender and conspicuous neck which is characteristic of the Diptera, or flies, is not to be found in any fleas. For this reason a flea cannot turn its head in any direction without at the same time following it round with its body.

The thorax of a flea consists of three segments called respectively the prothorax, mesothorax and metathorax. The chitinous external skeleton which covers each of these three segments is primarily a hoop but each hoop is further subdivided into a number of complicated plates. Attached to the thorax are the three pairs of legs which are characteristic of all adult insects. The hind pair are very much the strongest (Fig. 3). They are the organs of hopping. It has often been pointed out that if men had the leaping powers of some fleas they would bound with ease backwards and forwards over the cross on the top of St. Paul’s Cathedral. Each leg consists of four segments beautifully articulated and plentifully supplied with bristles. At the end comes the foot with five very short segments. The last segment is provided with a pair of more or less formidable claws. Fleas use their legs for leaping, for running, and for clinging to their hosts. They also use their mouth-parts for the last purpose and it is worthy of note, as we shall see later on, that in those fleas in which the mouth-parts are shortest and weakest the legs are most liberally supplied with bristles and possess the stoutest claws. The legs of a flea are unique in the insect world owing to the enormous development of the segment nearest the body called the coxa. Most leaping insects rely for their activity on the muscles of the lower joints. In a grasshopper it is the third joint from the body (femur) which is so immensely enlarged. The three pairs of legs are each attached to a different thoracic segment.

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Fig. 3. The hind leg of a flea. The segment or joint nearest the body is thecoxawhich is unusually developed. Next come the smalltrochanterand the largerfemur. Thetibiawhich is long and slender follows. Then come the fivetarsiwith the sixth and ultimate segment provided with claws.

Fig. 3. The hind leg of a flea. The segment or joint nearest the body is thecoxawhich is unusually developed. Next come the smalltrochanterand the largerfemur. Thetibiawhich is long and slender follows. Then come the fivetarsiwith the sixth and ultimate segment provided with claws.

Fig. 3. The hind leg of a flea. The segment or joint nearest the body is thecoxawhich is unusually developed. Next come the smalltrochanterand the largerfemur. Thetibiawhich is long and slender follows. Then come the fivetarsiwith the sixth and ultimate segment provided with claws.

When fleas walk, they are so to speak plantigrades walking on the sole of the foot, and all the tarsal or foot joints are applied to the surface of the ground. The claws serve as grips so as to make the most of any unevenness; and thus the insect drags itself along with surprising rapidity when it moves through the hairy coat of a mammal. But on an open surface fleas are not really rapid movers compared with many other insects.

The two claws on the end of each ultimate foot segment are freely moveable and are in fact highly modified bristles or setæ.

In all fleas one of the plates of the metathorax (or hindmost thoracic segment) called the epimeron, is large and prolonged towards the rear. It invariably bears a stigma. The epimeron is placed laterally to the first abdominal tergite. The older naturalists jumped to the conclusion that this was the remains of a wing. The best judges have, however, formed a decided opinion that no trace of the relic of a flying organ can be detected on the thorax of a flea. Heymons, a German entomologist, has also failed to detect any sign in dissections which he has made of the larvæ and the pupæ.

The epimeron is in fact neither a scale nor awing but a portion of the thorax present in all insects. It is of no special service to the flea except as a portion of the thoracic armature which covers the body.

The larva of a flea has no legs; the adult insect has six. A study of other embryo insects shows that the ancestors of insects had many legs. It is an interesting problem why insects lost the legs on their abdomens, why legs should now invariably be restricted to the thorax, and why there should never be more than three pairs. In the earliest known insects which lived on the earth, before winged forms were evolved, the number of legs was already six. But our knowledge of fleas is too small to attempt, at present, to trace their exact line of ancestral descent.

The abdomen of a flea consists of ten segments. The horny plates which cover the dorsal side are called tergites; those on the ventral side sternites. In fleas, as in all holometabolous insects, that is those which pass through a complete metamorphosis, the sternite of the first abdominal segment is suppressed and has completely disappeared. The tergite which covers the dorsal part of the first abdominal segment nearest to the thorax is, however, always present.

The ultimate segments of the male and female flea are modified for reproductive purposes and of these segments more must be said later.

Having now given a rough outline of the external skeleton of a flea, it only remains to say something about the muscular system. Attached to the inside of the chitinous armature are an enormous number of muscles, whitish and almost transparent. They act as extensors, retractors, flexors, elevators and depressors. The joints and hinges of the skeleton allow of considerable, but not perfect, freedom. The muscles of locomotion are partly in the thorax and partly in the several joints of the legs. Our knowledge of the muscular system of fleas is very imperfect. But, as in other insects, the general arrangement of the muscles is based on the segmented structure of the body.

For the reader who can accurately picture to himself the external structure of a flea and of the typical insects belonging to other orders, a few words may be said on the probable ancestry of fleas and their relationship to other living insects. This vexed and much debated question is still, as the older naturalists would have said,tremendum mysterium. Very little light has yet been thrown upon it, and the most divergent views have been expressed by learned and competent entomologists. A historic survey of the various opinions that have been held since the days of Linnæus would fill many pages; but a short summary of the different orders to which fleas have been referred by different zoologists will suffice.

The older authors, Linnæus, Geoffroy, Cuvier and Duméril, and Gervais placed them among theApterabecause they were wingless. Kircher regarded them asOrthoptera, an order which includes grasshoppers and crickets; but he has had few followers. By Fabricius and by Illiger they were treated asHemipteraor bugs. Lameere, a Belgian, has recently expressed a decided view that fleas are really a family ofColeopteraor beetles. Those who have held the once orthodox opinion that they belonged to theDipteraor flies are Roesel, Oken, Straus-Dürkheim, Burmeister, Newman, Walker, von Siebold and Wagner.

The structure of an adult flea, however, differs from that of an adult fly in the following noteworthy respects: the mouth-parts are differently constructed, the head of the flea is closely joined to its thorax, the three divisions of the thorax are not joined and fused, the flea is wingless, the eyes of fleas are simple ocelli, and there are differences of lesser importance in the stigmata, which give access to the tracheal system by which all insects breathe.

The number of those who have regarded fleas as belonging to a distinct order of insects is considerable: they are Lamarck, De Geer, Latreille, Kirby and Spence, MacLeay, Leach, Dugès, Bouché, van der Hoeven, Westwood, Landois, Brauer, Kraepelin,and Taschenberg. Modern opinion is all but unanimous on this point.

There remains, however, a second question. Even if it be agreed that there must be a distinct order forSuctoria, Aphaniptera, Siphonaptera, or fleas; where ought that order to be placed? In which other order of insects must we look for the nearest relations of fleas? For a time after the acceptance of the fact that insect forms have been evolved, and not separately created, the ancestors of fleas were searched for among some species of fly.

Then Kraepelin rejected the view that flies were as closely related to fleas as most entomologists thought and his followers could only find points of difference and no points of resemblance. Dahl (1899), a German, then took up the cudgels for the fly theory. Dahl pointed out the resemblance between fleas and a group of flies calledPhoridæalso parasitic on warm-blooded animals. During the ensuing years the debate was resumed afresh with much liveliness and sometimes with a little acrimony.

The fleas were placed by MacLeay and by Balbiani between theDipteraandHemiptera; by Leach between theHemipteraandLepidoptera; by Dugès between theHymenopteraandDiptera; by Brauer between theDipteraandColeoptera. Handlirsch thinks that fleas have no connection atall with beetles and Gross can find no signs of relationship with eitherColeopteraorDiptera.

Embryology and the study of larval forms have thrown so much light on the ancestry of many animals, that it was hoped that a microscopic examination of the larvæ of fleas, in various stages of development, would produce some facts of importance. In this hope entomologists have, to a great extent, been disappointed. There seems to be much similarity between the embryos of beetles, moths, flies, wasps and fleas. Those who have dwelt on the likeness of the larval flea to the maggot of a fly seem to forget that the resemblance to an embryo beetle is nearly as strong.

The young larva of the flea is very transparent and the digestive canal, heart and nervous system are easily recognised. The egg-shell breaker is an interesting example of the development of a temporary larval structure and it is the only known instance of such a structure in an insect. There are no traces of eyes. The antennæ are three-jointed. They are rather long and slender, being about one-third as long as the head. The head is well-developed and the larva has no feet.

The biting mandibles are broad and triangular. Compared with those of other larvæ they are said to be more like the mandibles of coleopterous than of dipterous larvæ. The maxillæ, or second pair ofjaws, are somewhat reduced and rudimentary. The absence of eyes and of legs are points of similarity between the larvæ of fleas and flies. The maggot of a fly has also two pairs of jaws, and a pair of antennæ.

At the tail end of the larval flea’s abdomen are two small projections called caudal stylets (Fig. 1). They are strong, recurved, chitinous, structures which prop up the body of the larva when it creeps and wriggles. There are similar props in the larvæ of certain beetles and no exactly similar organs are known in dipterous larvæ. But caudal stylets are of small taxonomic importance.

In one respect the mature flea is certainly nearer to a beetle than to a fly: the three joints of the thorax are free as in a beetle and not fused as in a fly; but when one studies the mouth-parts, the true view seems to be that the mouth-parts of a flea are equally unlike those of a fly and those of a beetle. Such being the present state of our knowledge, one must wait for fresh light to be thrown on the matter by further researches. It seems unlikely that the immediate future will produce a solution of the problem.

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