Chapter 17

(H. F. Ba.)

1The word “function” (from Lat.fungi, to perform) has many uses, with the fundamental sense of an activity special or proper to an office, business or profession, or to an organ of an animal or plant, the definite work for which the organ is an apparatus. From the use of the word, as in the Italianfunzione, for a ceremony of the Roman Church, “function” is often employed for a public ceremony of any kind, and loosely of a social entertainment or gathering.

FUNDY, BAY OF,an inlet of the North Atlantic, separating New Brunswick from Nova Scotia. It is 145 m. long and 48 m. wide at the mouth, but gradually narrows towards the head, where it divides into Chignecto Bay to the north, which subdivides into Shepody Bay and Cumberland Basin (the French Beaubassin), and Minas Channel, leading into Minas Basin, to the east and south. Off its western shore opens Passamaquoddy Bay, a magnificent sheet of deep water with good anchorage, receiving the waters of the St Croix river and forming part of the boundary between New Brunswick and the state of Maine, The Bay of Fundy is remarkable for the great rise and fall of the tide, which at the head of the bay has been known to reach 62 ft. In Passamaquoddy Bay the rise and fall is about 25 ft., which gradually increases toward the narrow upper reaches. At spring tides the water in the Bay of Fundy is 19 ft. higher than it is in Bay Verte, in Northumberland Strait, only 15 m. distant. Though the bay is deep, navigation is rendered dangerous by the violence and rapidity of the tide, and in summer by frequent fogs. At low tide, at such points as Moncton or Amherst, only an expanse of red mud can be seen, and the tide rushes in a bore or crest from 3 to 6 ft. in height. Large areas of fertile marshes are situated at the head of the bay, and the remains of a submerged forest show that the land has subsided in the latest geological period at least 40 ft. The bay receives the waters of the St Croix and St John rivers, and has numerous harbours, of which the chief are St Andrews (on Passamaquoddy Bay) and St John in New Brunswick, and Digby and Annapolis (on an inlet known as Annapolis Basin) in Nova Scotia. It was first explored by the Sieur de Monts (d.c.1628) in 1604 and named by him La Baye Française.

FUNERAL RITES,the ceremonies associated with different methods of disposing of the dead. (See alsoBurial and Burial Acts;Cemetery; andCremation.) In general we have little record, except in their tombs, of races which, in a past measured not merely by hundreds but by thousands of years, occupied the earth; and exploration of these often furnishes our only clue to the religions, opinions, customs, institutions and arts of long vanished societies. In the case of the great culture folks of antiquity, the Babylonians, Egyptians, Hindus, Persians, Greeks and Romans, we have, besides their monuments, the evidence of their literatures, and so can know nearly as much of their rites as we do of our own. The rites of modern savages not only help us to interpret prehistoric monuments, but explain peculiarities in our own rituals and in those of the culture folks of the past of which the significance was lost or buried under etiological myths. We must not then confine ourselves to the rites of a few leading races, neglecting their less fortunate brethren who have never achieved civilization. It is better to try to classify the rites of all races alike according as they embody certain leading conceptions of death, certain fears, hopes, beliefs entertained about the dead, about their future, and their relations with the living.

The main ideas, then, underlying funeral rites may roughly be enumerated as follows:1. The pollution or taboo attaching to a corpse.2. Mourning.3. The continued life of the dead as evinced in the housing and equipment of the dead, in the furnishing of food for them, and in the orientation and posture assigned to the body.4. Communion with the dead in a funeral feast and otherwise.5. Sacrifice for the dead and expiation of their sins.6. Death witchery.7. Protection of the dead from ghouls.8. Fear of ghosts.

The main ideas, then, underlying funeral rites may roughly be enumerated as follows:

1. The pollution or taboo attaching to a corpse.

2. Mourning.

3. The continued life of the dead as evinced in the housing and equipment of the dead, in the furnishing of food for them, and in the orientation and posture assigned to the body.

4. Communion with the dead in a funeral feast and otherwise.

5. Sacrifice for the dead and expiation of their sins.

6. Death witchery.

7. Protection of the dead from ghouls.

8. Fear of ghosts.

1. A dead body is unclean, and the uncleanness extends to things and persons which touch it. Hence the Jewish law (Num. v. 2) enacted that “whoever is unclean by the dead shall be put outside the camp, that they defile not the camp in the midst whereof the Lord dwells.” Such persons were unclean until the even, and might not eat of the holy things unless they bathed their flesh in water. A high priest might on no account “go in to any dead body” (Lev. xxi. 11). Why a corpse is so widely tabooed is not certain; but it is natural to see one reason in the corruption which in warm climates soon sets in. The common experience that where one has died another is likely to do so may also have contributed, though, of course, there was no scientific idea of infection. The old Persian scriptures are full of this taboo. He who has touched a corpse is “powerless in mind, tongue and hand” (Zend AvestainSacred Books of the East, pt. i. p. 120), and the paralysis is inflicted by the innumerabledrugsor evil spirits which invest a corpse. Fire and earth, being alike creations of the good and pure godAhuramazda, a body must not be burned or buried; and so the ancient Persians and their descendants the Parsees build Dakmas or “towers of silence” on hill-tops far from human habitations. Inside these the corpses are laid on a flagged terrace which drains into a central pit. Twice a year the bones, picked clean by dogs and birds of prey, are collected in the pit, and when it is full another tower is built. In ancient times perhaps the bodies of the magi or priests alone were exposed at such expense; the common folk were covered with wax and laid in the earth, the wax saving the earth from pollution. In Rome and Greece the corpse was buried by night, lest it should pollute the sunlight; and a trough of water was set at the door of the house of death that men might purify themselves when they came out, before mixing in general society. Priests and magistrates in Rome might not meet or look on a corpse, for they were thereby rendered unclean and incapable of fulfilling their official duties without undergoing troublesome rites of purification. At a Roman funeral, when the remains had been laid in the tomb, all present were sprinkled with lustral water from a branch of olive or laurel calledaspergillum; and when they had gone home they were asperged afresh and stepped over a fire. The house was also swept out with a broom, probably lest the ghost of the dead should be lying about the floor. Many races, to avoid pollution, destroy the house and property of the deceased. Thus the Navahos pull down the hut in which he died, leaving its ruins on the ground; but if it be an expensive hut, a shanty is extemporized alongside, into which the dying man is transferred before death. No one will use the timbers of a hut so ruined. A burial custom of the Solomon Islands, noted by R.H. Codrington (The Melanesians, p. 255), may be dictated by the same scruple. There “the mourners having hung up a dead man’s arms on his house make great lamentations; all remains afterwards untouched, the house goes to ruin, mantled, as time goes on, with the vines of the growing yams, a picturesque and indeed, perhaps, a touching sight; for these things are not set up that they may in a ghostly manner accompany their former owner.” H. Oldenberg (Religion des Veda, p. 426) describes how Hindus shave themselves and cut off their nails after a death, at the same time that they wash, renew the hearth fire, and furnish themselves with new vessels. For the hair and nails may harbour pollution, just as the medieval Greeks believed that evil spirits could lurk in a man’s beard (Leo Allatius,De opinionibus quorundam Graecorum). The dead man’s body is shorn and the nails cut for a kindred reason; for it must be purified as much as can be before it is burned as an offering on the pyre and before he enters on a new sphere of existence.

2. We are accustomed to regard mourning costume as primarily an outward sign of our grief. Originally, however, the special garb seems to have been intended to warn the general public that persons so attired were unclean. In ancient Rome mourners stayed at home and avoided all feasts and amusements; laying aside gold, purple and jewels, they wore black dresses calledlugubriaor even skins. They cut neither hair nor beard, nor lighted fire. Under the emperors women began to wear white. On the west coast of Africa negroes wear white, on the Gold Coast red. The Chinese wear hemp, which is cheap, for mourning dress must as a rule be destroyed when the season of grief is past to get rid of the taboo. Among the Aruntas of Australia the wives of a dead man smear themselves with white pipe-clay until the last ceremonies are finished, sometimes adding ashes—this not to conceal themselves from the ghost (which may partly be the aim of some mourning costumes), but to show the ghost that they are duly sorrowing for their loss. These widows must not talk except on their hands for a whole year. “Among the Maoris,” says Frazer (Golden Bough, i. 323), “anyone who had handled a corpse, helped to convey it to the grave, or touched a dead man’s bones; was cut off from all intercourse and almost all communication with mankind. He could not enter any house, or come into contact with any person or thing, without utterly bedevilling them. He might not even touch food with his hands, which had become so frightfully tabooed or unclean as to be quite useless. Food would be set for him on the ground, and he would then sit or kneel down, and, with his hands carefully held behind his back, would gnaw at it as best he could.” Often a degraded outcast was kept in a village to feed mourners. Such a taboo is strictly similar to those which surround a sacred chief or his property, a menstruous woman or a homicide, rendering them dangerous to themselves and to all who approach them.

3. Primitive folk cannot conceive of a man’s soul surviving apart from his body, nor of another life as differing from this, and the dead must continue to enjoy what they had here. Accordingly the Patagonians kill horses at the grave that the dead may ride toAlhuemapu, or country of the dead. After a year they collect a chief’s bones, arrange them, tie them together and dress them in his best garments with beads and feathers. Then they lay him with his weapons in a square pit, round which dead horses are placed set upright on their feet by stakes. As late as 1781 in Poland F. Casimir’s horse was slain and buried with him. In the Caucasus a Christian lady’s jewels are buried with her. The Hindus used to burn a man’s widow on his pyre, because he could not do without her; and St Boniface commends the self-sacrifice of the Wend widows who in his day burned themselves alive on their husbands’ pyres.

The tumuli met with all over the north of Europe (in the Orkneys alone 2000 remain) are regular houses of the dead, models of those they occupied in life. The greater the dignity of the deceased, the loftier was his barrow. Silbury hill is 170 ft. high; the tomb of Alyattes, father of Croesus, was a fourth of a league round; the Pyramids are still the largest buildings in existence; at Oberea in Tahiti is a barrow 267 ft. long, 87 wide and 44 high. Some Eskimo just leave a dead man’s body in his house, and shut it up, often leaving by his side a dog’s head to guide him on his last journey, along with his tools and kayak. The Sea Dyaks set a chief adrift in his war canoe with his weapons. So in Norse story Hake “was laid wounded on a ship with the dead men and arms; the ship was taken out to sea and set on fire.” The Viking was regularly buried in his ship or boat under a great mound. He sailed after death to Valhalla. In the ship was laid a stone as anchor and the tools, clothes, weapons and treasures of the dead. The Egyptians, whose land was the gift of the river Nile, equally believed that the dead crossed over water, and fashioned the hearse in the form of a boat. Hence perhaps was derived the Greek myth of Charon and the Styx, and the custom, which still survives in parts of Europe, of placing a coin in the mouth of the dead with which to pay the ferryman. The Egyptians placed in the tomb books of a kind to guide the dead to the next world. The Copts in a later age did the same, and to this custom we owe the recovery in Egypt of much ancient literature. The Armenians till lately buried with a priest his missal or gospel.

In Egyptian entombments of the XIIth to the XIVth dynasties were added above the sepulchres what Professor Petrie terms soul-houses, viz. small models of houses furnished with couch and table, &c., for the use of thekaor double whenever it might wish to come above ground and partake of meats and drinks. They recall, in point of size, the hut-urns of the Etruscans, but the latter had another use, for they contain incinerated remains. Etruscan tombs, like those of Egypt and Asia Minor, were made to resemble the dwelling-houses of the living, and furnished with coffered ceilings, panelled walls, couches, stools, easy chairs with footstools attached, all hewn out of the living rock (Dennis,Cities and Cemeteries of Etruria, vol i. p. lxx.).

Of the old Peruvian mummies in the Kircherian Museum at Rome, several are of women with babies in their arms, whence it is evident that a mother had her suckling buried with her; it would console her in the next world and could hardly survive her in this. The practice of burying ornaments, tools and weapons with the dead characterizes the inhumations of the Quaternary epoch, as if in that dim and remote age death was already regarded as the portal of another life closely resembling this. The cups, tools, weapons, ornaments and other articles deposited with the dead are often carefully broken or turned upside down and inside out; for the soul ormanesof objects is liberated by such fracture or inversion and so passes into thedead man’s use and possession. For the same reason where the dead are burned, their properties are committed to the flames. The ghost of the warrior has a ghostly sword and buckler to fight with and a ghostly cup to drink from, and he is also nourished by the impalpable odour and reek of the animal victims sacrificed over his grave. Instead of valuable objects cheap images and models are often substituted; and why not, if the mere ghosts of the things are all that the wraith can enjoy? Thus Marco Polo (ii. 76) describes how in the land of Kinsay (Hang-chau) “the friends and relations make a great mourning for the deceased, and clothe themselves in hempen garments, and follow the corpse, playing on a variety of instruments and singing hymns to their idols. And when they come to the burning place they take representations of things cut out of parchment, such as caparisoned horses, male and female slaves, camels, armour, suits of cloth of gold (and money), in great quantities, and these things they put on the fire along with the corpse so that they are all burned with it. And they tell you that the dead man shall have all these slaves and animals of which the effigies are burned, alive in flesh and blood, and the money in gold, at his disposal in the next world; and that the instruments which they have caused to be played at his funeral, and the idol hymns that have been chaunted shall also be produced again to welcome him in the next world.” The manufacture of such papersimulacrafor consumption at funerals is still an important industry in Chinese cities. The ancient Egyptians, assured that a man’skaor double shall revivify his body, took pains to guard the flesh from corruption, steeping the corpse in natron and stuffing it with spices. A body so prepared is called a mummy (q.v.), and the custom was already of a hoary antiquity in 3200B.C., when the oldest dated mummy we have was made. The bowels, removed in the process, were placed in jars over the corpse in the tomb, together with writing tablets, books, musical instruments, &c., of the dead. Cemeteries also remain full of mummies of crocodiles, cats, fish, cows and other sacred animals. The Greeks settled in Egypt learned to mummify their dead, but the custom was abhorrent to the Jews, although the Christian belief in the resurrection of the flesh must have been formed to a large extent under Egyptian influence. Half the superiority of the Jewish to other ancient religions lay in this, that it prescribed no funeral rites other than the simplest inhumation.

The dead all over the world and from remote antiquity have been laid not anyhow in the earth, but with the feet and face towards the region in which their future will be spent; the Samoans and Fijians towards the far west whither their souls have preceded them; the Guarayos with head turned eastwards because their god Tamoi has in that quarter “his happy hunting grounds where the dead will meet again” (Tylor,Prim. Cult.ii. 422). The legend is that Christ was buried with His head to the west, and the church follows the custom, more ancient than itself, of laying the dead looking to the East, because that is the attitude of prayer, and because at the last trump they will hurry eastwards. So in Eusebius (Hist. Eccl.430.19) a martyr explains to his pagan judge that the heavenly Jerusalem, the fatherland of the pious, lay exactly in the east at the rising place of the sun. Where the body is laid out straight it is difficult to discern the presence of any other idea than that it is at rest. In Scandinavian barrows,e.g.in the one opened at Goldhavn in 1830, the skeletons have been found seated on a low stone bench round the wall of the grave chamber facing its opening, which always looks south or east, never north. Here the dead were continuing the drinking bouts they enjoyed on earth.

The Peruvians mummified their dead and placed them jointed and huddled up with knees to chin, looking toward the sunset, with the hands held before the face. In the oldest prehistoric tombs along the Nile the bodies are doubled up in the same position. It would seem as if in these and numerous other similar cases the dead were deliberately given in their graves the attitude of a foetus in the womb, and, as Dr Budge remarks (Egyptian Ideas of the Future Life, London, 1899, p. 162), “we may perhaps be justified in seeing in this custom the symbol of a hope that, as the child is born from this position into the world, so might the deceased be born into the life beyond the grave.” The late Quaternary skeletons of the Mentone cave were laid in a layer of ferrugineous earth specially laid down for them, and have contracted a red colour therefrom. Many other prehistoric skeletons found in Italy have a reddish colour, perhaps for the same reason, or because, as often to-day, the bones were stripped of flesh and painted. Ambrose relates that the skeletons of the martyrs Gervasius and Protasius, which he found and depositedA.D.386 under the altar of his new basilica in Milan, weremirae magnitudinis ut prisca aetas ferebat, and were also coloured red. He imagined the red to be the remains of the martyrs’ blood!Hic sanguis clamat coloris indicio.Salomon Reinach has rightly divined that what Ambrose really hit upon was a prehistoric tomb. Red earth was probably chosen as a medium in which to lay a corpse because demons flee from red. Sacred trees and stones are painted red, and for the most solemn of their rites savages bedaub themselves with red clay. It is a favourite taboo colour.

4. A feast is an essential feature of every primitive funeral, and in the Irish “wake” it still survives. A dead man’s soul or double has to be fed at the tomb itself, perhaps to keep it from prowling about the homes of the survivors in search of victuals; and such food must also be supplied to the dead at stated intervals for months or years. Many races leave a narrow passage or tube open down to the cavity in which the corpse lies, and through it pour down drinks for the dead. Traces of such tubes are visible in the prehistoric tombs of the British Isles. However, such provision of food is not properly a funeral feast unless the survivors participate. In the Eastern churches and in Russia the departed are thus fed on the ninth, twelfth and fortieth days from death. “Ye appease the shades of the dead with wine and meals,” was the charge levelled at the Catholics by the 4th-century Manichaeans, and it has hardly ceased to be true even now after the lapse of sixteen centuries. The funeral feast proper, however, is either a meal of communion with or in the dead, which accompanies interment, or a banquet off the flesh of victims slain in atonement of the dead man’s sins. Some anthropologists see in the common meal held at the grave “the pledge and witness of the unity of the kin, the chief means, if not of making, at least of repairing and renewing it.”1The flesh provided at these banquets is occasionally that of the dead man himself; Herodotus and Strabo in antiquity relate this of several half-civilized races in the East and West, and a similar story is told by Marco Polo of certain Tatars. Nor among modern savages are funeral feasts off the flesh of the dead unknown, and they seem to be intended to effect and renew a sacramental union or kinship of the living with the dead. The Uaupes in the Amazons incinerate a corpse a month after death, pound up the ashes, and mix them with their fermented drink. They believe that the virtues of the dead will thus be passed on to his survivors. The life of the tribe is kept inside the tribe and not lost. Such cannibal sacraments, however, are rare, and, except in a very few cases, the evidence for them weak. The slaying and eating of animal victims, however, at the tomb is universal and bears several meanings, separately or all at once. The animals may be slain in order that their ghosts may accompany the deceased in his new life. This significance we have already dwelt upon. Or it is believed that the shade feeds upon them, as the shades came up from Hades and lapped up out of a trench the blood of the animals slain by Ulysses. The survivors by eating the flesh of a victim, whose blood and soul the dead thus consume, sacramentally confirm the mystic tie of blood kinship with the dead. Or lastly, the victim may be offered for the sins of the dead. His sins are even supposed to be transferred into it and eaten by the priest. Such expiatory sacrifices of animals for the dead survive in the Christian churches of Armenia, Syria and of the East generally. Their vicarious character is emphasized in the prayers which accompany them, but the popular understanding of them probably combines all the meanings above enumerated. It has been suggested by Robertson Smith (Religion of the Semites, 336) that the world-wide customs oftearing the hair, rending the garments, and cutting and wounding the body were originally intended to establish a life-bond between the dead and the living. The survivors, he argues, in leaving portions of their hair and garments, and yet more by causing their own blood to stream over the corpse from self-inflicted wounds, by cutting off a finger and throwing it into the grave, leave what is eminently their own with the dead, so drawing closer their tie with him. Conversely, many savages daub themselves with the blood and other effluences of their dead kinsmen, and explain their custom by saying that in this way a portion of the dead is incorporated in themselves. Often the survivors, especially the widows, attach the bones or part of them to their persons and wear them, or at least keep them in their houses. The retention of the locks of the deceased and of parts of his dress is equally common. There is also another side to such customs. Having in their possession bits of the dead, and being so far in communion with him, the survivors are surer of his friendship. They have ensured themselves against ghosts who are apt to be by nature envious and mischievous. But whatever their original significance, the tearing of cheeks and hair and garments and cutting with knives are mostly expressions of real sorrow, and, as Robertson Smith remarks, of deprecation and supplication to an angry god or spirit. It must not be supposed that the savage or ancient man feels less than ourselves the poignancy of loss.

6. Death-witchery has close parallels in the witch and heretic hunts of the Christians, but, happily for us, only flourishes to-day among savages. Sixty % of the deaths which occur in West Africa are, according to Miss Mary Kingsley—a credible witness—believed to be due to witchcraft and sorcery. The blacks regard old age or effusion of blood as the sole legitimate causes of death. All ordinary diseases are in their opinion due to private magic on the part of neighbours, just as a widespread epidemic marks the active hatred “of some great outraged nature spirit, not of a mere human dabbler in devils.”2Similarly in Christian countries an epidemic is set down to the wrath of a God offended by the presence of Jews, Arians and other heretics. The duty of an African witch-doctor is to find out who bewitched the deceased, just as it was of an inquisitor to discover the heretic. Every African post-mortem accordingly involves the murder of the person or persons who bewitched the dead man and caused him to die. The death-rate by these means is nearly doubled; but, since the use of poison against an obnoxious neighbour is common, the right person is occasionally executed. It is also well for neighbours not to quarrel, for, if they do and one of them dies of smallpox, the other is likely to be slain as a witch, and his lungs, liver and spleen impaled on a pole at the entrance of the village. It is the same case with the Australian blacks: “no such thing as natural death is realized by the native; a man who dies has of necessity been killed by some other man, or perhaps even by a woman, and sooner or later that man or woman will be attacked. In the normal condition of the tribe every death meant the killing of another individual.”3

7. Lastly, a primitive interment guards against the double risk of the ghost haunting the living and of ghouls or vampires taking possession of the corpse. The latter end is likely to be achieved if the body is cremated, for then there is no nidus to harbour the demon; but whether, in the remote antiquity to which belong many barrows containing incinerated remains, this motive worked, cannot be ascertained. The Indo-European race seems to have cremated at an early epoch, perhaps before the several races of East and West separated. In Christian funeral rites many prayers are for the protection of the body from violation by vampires, and it would seem as if such a motive dictated the architectural solidity of some ancient tombs. Christian graves were for protection regularly sealed with the cross; and the following is a characteristic prayer from the old Armenian rite for the burial of a layman:

“Preserve, Almighty Lord, this man’s spirit with all saints and with all lovers of Thy holy name. And do Thou seal and guard the sepulchre of Thy servant, Thou who shuttest up the depths and sealest them with Thy almighty right hand ... so let the seal of Thy Lordship abide unmoved upon this man’s dwelling-place and upon the shrine which guards Thy servant. Andlet not any filthy and unclean devil dare to approach him, such as assail the body and souls of the heathen, who possess not the birth of the holy font, and have not the dread seal laid upon their graves.”

A terrible and revolting picture of the superstitious belief in ghouls which violate Christian tombs is given by Leo Allatius (who held it) in his tractDe opinionibus quorundam Graecorum(Paris, 1646). It was probably the fear of such demonic assaults on the dead that inspired the insanitary custom of burying the dead under the floors of churches, and as near as possible to the altar. In the Greek Church this practice was happily forbidden by the code of Justinian as well as by the older law in the case of churches consecrated withEncaeniaand deposition of relics. In the Armenian Church the same rule holds, and Ephrem Syrus in his testament particularly forbade his body to be laid within a church. Such prohibitions, however, are a witness to the tendency in question.

The custom of lighting candles round a dead body and watching at its side all night was originally due to the belief that a corpse, like a person asleep, is specially liable to the assaults of demons. The practice of tolling a bell at death must have had a similar origin, for it was a common medieval belief that the sound of a consecrated bell drives off the demons which when a man dies gather near in the air to waylay his fleeting soul. For a like reason the consecrated bread of the Eucharist was often buried with believers, and St Basil is said to have specially consecrated a Host to be placed in his coffin.

8. Some of the rites described under the previous heads may be really inspired by the fear of the dead haunting the living, but it must be kept in mind that the taboo attaching to a dead body is one thing and fear of a ghost another. A corpse is buried or burned, or scaffolded on a tree, a tower or a house-top, in order to get it out of the way and shield society from the dangerous infection of its taboo; but ghostsquâghosts need not be feared and a kinsman’s ghost usually is not. On the contrary, it is fed and consoled with everything it needs, is asked not to go away but to stay, is in a thousand ways assured of the sorrow and sympathy of the survivors. Even if the body be eaten, it is merely to keep the soul of the deceased inside the circle of kinsmen, and Strabo asserts that the ancient Irish and Massagetae regarded it as a high honour to be so consumed by relatives. In Santa Cruz in Melanesia they keep the bones for arrow heads and store a skull in a box and set food before it “saying that this is the man himself” (R.H. Codrington,The Melanesians, p. 264), or the skull and jaw bone are kept and “are calledmangite, which aresaka, hot with spiritual power, and by means of which the help of thelio’a, the powerful ghost of the man whose relics these are, can be obtained” (ibid.p. 262). Here we have the savage analogue to Christian relics. So the Australian natives make pointing sticks out of the small bones of the arm, with which to bewitch enemies.

We may conclude then that in the most primitive societies, where blood-kinship is the only social tie and root of social custom it is the shades, not of kinsmen, but of strangers, who as such are enemies, that are dangerous and uncanny. In more developed societies, however, all ghosts alike are held to be so; and if a ghost walks it is because its body has not been properly interred or because its owner was a malefactor. Still, even allowing for this, it remains true that for a friendly ghost the proper place is the grave and not the homes of the living, and accordingly the Aruntas with cries ofWah! Wah!with wearing of fantastic head-dresses, wild dancing and beating of the air with hands and weapons “drive the spirit away from the old camp which it is supposed to haunt,” and which has been set fire to, and hunt it at a run into the grave prepared, and there stamp it down into the earth. “The loud shouting of the men and women shows him that they do not wish to be frightened by him in his present state, and that they will be angry with him if he does not rest.”(Spencer and Gillen,Native Tribes of Central Australia, p. 508). In Mesopotamia cemeteries have been discovered where the sepulchral jars were set upside down, clearly by way of hindering the ghosts from escaping into the upper world. In the Dublin museum we see specimens of ancient Celtic tombs showing the same peculiarity. For a like reason perhaps the name of the dead must among the Aruntas not be uttered, nor the grave approached, by certain classes of kinsmen. The same repugnance to naming the dead exists all over the world, and leads survivors who share the dead man’s name to adopt another, at least for a time. If the dead man’s name was that of a plant, tree, animal or stream, that too is changed. Here is a potent cause of linguistic change, that also renders any historical tradition impossible. The survivors seem to fear that the ghost will come when he hears his name called; but it also hangs together with the taboo which hedges round the dead as it does kings, chieftains and priests.

Authorities.—B. Spencer and F.J. Gillen,The Native Tribes of Central Australia(London, 1899); F.B. Jevons,Introduction to History of Religion(London, 1896); E.S. Hartland,The Legend of Perseus, vol. ii.; J.G. Frazer,The Golden Bough(London, 1900); L.W. Faraday, “Custom and Belief in the Icelandic Sagas,” inFolk-lore, vol. xvii. No. 4; E.B. Tylor,Primitive Culture(London, 1903); E.A. W. Budge,The Mummy(Cambridge, 1893); C. Royer, “Les Rites funéraires aux époques préhistoriques,”Revue d’anthropologie(1876); Forrer,Über die Totenbestattung bei den Pfahlbauern(Ausland, 1885); J. Lubbock,Origin of Civilization(London, 1875) andPrehistoric Times(London, 1865); L.A. Muratori, “De antiquis Christianorum sepulchris,”Anecd. Graeca(Padua, 1709); Onaphr. Panvinius,De ritu sepeliendi mortuos apua veteres Christianos, reprinted in Volbeding’sThesaurus(Leipzig, 1841).

(F. C. C.)

1E.S. Hartland,Legend of Perseus(1895), ii. 278.2Mary Kingsley,West African Studies(1901), p. 178.3B. Spencer and F.J. Gillen,The Native Tribes of Central Australia(1899), p. 48.

1E.S. Hartland,Legend of Perseus(1895), ii. 278.

2Mary Kingsley,West African Studies(1901), p. 178.

3B. Spencer and F.J. Gillen,The Native Tribes of Central Australia(1899), p. 48.

FUNGI(pl. of Lat.fungus, a mushroom), the botanical name covering in the broad sense all the lower cellular Cryptogams devoid of chlorophyll, which arise from spores, and the thallus of which is either unicellular or composed of branched or unbranched tubes or cell-filaments (hyphae) with apical growth, or of more or less complex wefted sheets or tissue-like masses of such (mycelium). The latter may in certain cases attain large dimensions, and even undergo cell-divisions in their interior, resulting in the development of true tissues. The spores, which may be uni- or multicellular, are either abstricted free from the ends of hyphae (acrogenous), or formed from segments in their course (chlamydospores) or from protoplasm in their interior (endogenous). The want of chlorophyll restricts their mode of life—which is rarely aquatic—since they are therefore unable to decompose the carbon dioxide of the atmosphere, and renders them dependent on other plants or (rarely) animals for their carbonaceous food-materials. These they obtain usually in the form of carbohydrates from the dead remains of other organisms, or in this or other forms from the living cells of their hosts; in the former case they are termed saprophytes, in the latter parasites. While some moulds (Penicillium,Aspergillus) can utilize almost any organic food-materials, other fungi are more restricted in their choice—e.g.insect-parasites, horn- and feather-destroying fungi and parasites generally. It was formerly the custom to include with the Fungi the Schizomycetes or Bacteria, and the Myxomycetes or Mycetozoa; but the peculiar mode of growth and division, the cilia, spores and other peculiarities of the former, and the emission of naked amoeboid masses of protoplasm, which creep and fuse to streaming plasmodia, with special modes of nutrition and spore-formation of the latter, have led to their separation as groups of organisms independent of the true Fungi. On the other hand, lichens, previously regarded as autonomous plants, are now known to be dual organisms—fungi symbiotic with algae.

The number of species in 1889 was estimated by Saccardo at about 32,000, but of these 8500 were so-calledFungi imperfecti—i.e.forms of which we only know certain stages, such as conidia, pycnidia, &c., and which there are reasons for regarding as merely the corresponding stages of higher forms. Saccardo also included about 400 species of Myxomycetes and 650 of Schizomycetes. Allowing for these and for the cases, undoubtedly not few, where one and the same fungus has been described under different names, we obtain Schroeter’s estimate (in 1892) of 20,000 species. In illustration of the very different estimates that have been made, however, may be mentioned that of De Bary in 1872 of 150,000 species, and that of Cooke in 1895 of 40,000, and Massee in 1899 of over 50,000 species, the fact being that no sufficient data are as yet to hand for any accurate census. As regards their geographical distribution, fungi, like flowering plants, have no doubt their centres of origin and of dispersal; but we must not forget that every exchange of wood, wheat, fruits, plants, animals, or other commodities involves transmission of fungi from one country to another; while the migrations of birds and other animals, currents of air and water, and so forth, are particularly efficacious in transmitting these minute organisms. Against this, of course, it may be argued that parasitic forms can only go where their hosts grow, as is proved to be the case by records concerning the introduction ofPuccinia malvacearum,Peronospora viticola,Hemileia vastatrix, &c. Some fungi—e.g.moulds and yeasts—appear to be distributed all over the earth. That the north temperate regions appear richest in fungi may be due only to the fact that North America and Europe have been much more thoroughly investigated than other countries; it is certain that the tropics are the home of very numerous species. Again, the accuracy of the statement that the fleshy Agaricini, Polyporei,Pezizae, &c., are relatively rarer in the tropics may depend on the fact that they are more difficult to collect and remit for identification than the abundantly recorded woody and coriaceous forms of these regions. When we remember that many parts of the world are practically unexplored as regards fungi, and that new species are constantly being discovered in the United States, Australia and northern Europe—the best explored of all—it is clear that no very accurate census of fungi can as yet be made, and no generalizations of value as to their geographical distribution are possible.

The existence of fossil fungi is undoubted, though very few of the identifications can be relied on as regards species or genera. They extend back beyond the Carboniferous, where they occur as hyphae, &c., preserved in the fossil woods, but the best specimens are probably those in amber and in siliceous petrifactions of more recent origin.

Organs.—Individual hyphae or their branches often exhibit specializations of form. In many Basidiomycetes minute branches arise below the septa; their tips curve over the outside of the latter, and fuse with the cell above just beyond it, forming aclamp-connexion. Many parasitic hyphae put out minute lateral branches, which pierce the cell-wall of the host and form a peg-like (Trichosphaeria), sessile (Cystopus), or stalked (Hemileia), knot-like, or a more or less branched (Peronospora) or coiled (Protomyces) haustorium. InRhizopuscertain hyphae creep horizontally on the surface of the substratum, and then anchor their tips to it by means of a tuft of short branches (appressorium), the walls of which soften and gumthemselves to it, then another branch shoots out from the tuft and repeats the process, like a strawberry-runner. Appressoria are also formed by some parasitic fungi, as a minute flattening of the tip of a very short branch (Erysiphe), or the swollen end of any hypha which comes in contact with the surface of the host (Piptocephalis,Syncephalis), haustoria piercing in each case the cell-wall below. InBotrytisthe appressoria assume the form of dense tassels of short branches. InArthrobotrysside-branches of the mycelium sling themselves around the host (Tylenchus) much as tendrils round a support.Many fungi (Phallus,Agaricus,Fumago, &c.) when strongly growing put out ribbon-like or cylindrical cords, or sheet-like mycelial plates of numerous parallel hyphae, all growing together equally, and fusing by anastomoses, and in this way extend long distances in the soil, or over the surfaces of leaves, branches, &c. These mycelial strands may be white and tender, or the outer hyphae may be hard and black, and very often the resemblance of the subterranean forms to a root is so marked that they are termed rhizomorphs. The outermost hyphae may even put forth thinner hyphae, radiating into the soil like root-hairs, and the convergent tips may be closely appressed and so divided by septa as to resemble the root-apex of a higher plant (Armillaria mellea).Sclerotia.—Fungi, like other plants, are often found to store up large quantities of reserve materials (oil, glycogen, carbohydrates, &c.) in special parts of their vegetative tissues, where they lie accumulated between a period of active assimilation and one of renewed activity, forming reserves to be consumed particularly during the formation of large fructifications. These reserve stores may be packed away in single hyphae or in swollen cells, but the hyphae containing them are often gathered into thick cords or mycelial strands (Phallus, mushroom, &c.), or flattened and anastomosing ribbons and plates, often containing several kinds of hyphae (Merulius lacrymans). In other cases the strands undergo differentiation into an outer layer with blackened, hardened cell-walls and a core of ordinary hyphae, and are then termed rhizomorphs (Armillaria mellea), capable not only of extending the fungus in the soil, like roots, but also of lying dormant, protected by the outer casing. Such aggregations of hyphae frequently become knotted up into dense masses of interwoven and closely packed hyphae, varying in size from that of a pin’s head or a pea (Peziza,Coprinus) to that of a man’s fist or head, and weighing 10 to 25 ℔ or more (Polyporus Mylittae,P. tumulosus,Lentinus Woermanni,P. Sapurema, &c.). The interwoven hyphae fuse and branch copiously, filling up all interstices. They also undergo cutting up by numerous septa into short cells, and these often divide again in all planes, so that a pseudoparenchyma results, the walls of which may be thickened and swollen internally, or hardened and black on the exterior. In many cases the swollen cell-walls serve as reserves, and sometimes the substance is so thickly deposited in strata as to obliterate the lumen, and the hyphae become nodular (Polyporus sacer,P. rhinoceros,Lentinus Woermanni). The various sclerotia, if kept moist, give rise to the fructifications of the fungi concerned, much as a potato tuber does to a potato plant, and in the same way the reserve materials are consumed. They are principally Polyporei, Agaricini, Pezizae; none are known among the Phycomycetes, Uredineae or Ustilagineae. The functions of mycelial strands, rhizomorphs and sclerotia are not only to collect and store materials, but also to extend the fungus, and in many cases similar strands act as organs of attack. The same functions of storage in advance of fructification are also exercised by the stromata so common in Ascomycetes.Tissue Differentiations.—The simpler mycelia consist of hyphae all alike and thin-walled, or merely differing in the diameter of the branches of various orders, or in their relations to the environment, some plunging into the substratum like roots, others remaining on its surface, and others (aerial hyphae) rising into the air. Such hyphae may be multicellular, or they may consist of simple tubes with numerous nuclei and no septa (Phycomycetes), and are then non-cellular. In the more complex tissue-bodies of higher fungi, however, we find considerable differences in the various layers or strands of hyphae.An epidermis-like or cortical protective outer layer is very common, and is usually characterized by the close septation of the densely interwoven hyphae and the thickening and dark colour of their outer walls (sclerotia,Xylaria, &c.). Fibre-like hyphae with the lumen almost obliterated by the thick walls occur in mycelial cords (Merulius). Latex-tubes abound in the tissues ofLactarius,Stereum,Mycena,Fistulina, filled with white or coloured milky fluids, and Istvanffvi has shown that similar tubes with fluid or oily contents are widely spread in other Hymenomycetes. Sometimes fatty oil or watery sap is found in swollen hyphal ends, or such tubes contain coloured sap. Cystidia and paraphyses may be also classed here. InMerulius lacrymansHartig has observed thin-walled hyphae with large lumina, the septa of which are perforated like those of sieve-tubes.As regards its composition, the cell-wall of fungi exhibits variations of the same kind as those met with in higher plants. While the fundamental constituent is a cellulose in many Mucorini and other Phycomycetes, in others bodies like pectose, callose, &c., commonly occur, and Wisselingh’s researches show that chitin, a gluco-proteid common in animals, forms the main constituent in many cases, and is probably deposited directly as such, though, like the other substances, it may be mixed with cellulose. As in other cell-walls, so here the older membranes may be altered by deposits of various substances, such as resin, calcium oxalate, colouring matters; or more profoundly altered throughout, or in definite layers, by lignification, suberization (Trametes,Daedalea), or swelling to a gelatinous mucilage (Tremella,Gymnosporangium), while cutinization of the outer layers is common. One of the most striking alterations of cell-walls is that termedcarbonization, in which the substance gradually turns black, hard and brittle, as if charred—e.g.Xylaria,Ustulina, some sclerotia. At the other extreme the cell-walls of many lichen-fungi are soft and colourless, but turn blue in iodine, as does starch. The young cell-wall is always tenuous and flexible, and may remain so throughout, but in many cases thickenings and structural differentiations, as well as the changes referred to above, alter the primary wall considerably. Such thickening may be localized, andpits(e.g.Uredospores, septa of Basidiomycetes),spirals,reticulations,rings, &c. (capillitium fibres ofPodaxon,Calostoma,Battarrea), occur as in the vessels of higher plants, while sculptured networks, pittings and so forth are as common on fungus-spores as they are on pollen grains.Cell-Contents.—The cells of fungi, in addition to protoplasm, nuclei and sap-vacuoles, like other vegetable cells, contain formed and amorphous bodies of various kinds. Among those directly visible to the microscope are oil drops, often coloured (Uredineae) crystals of calcium oxalate (Phallus,Russula), proteid crystals (Mucor,Pilobolus, &c.) and resin (Polyporei). The oidia of Erysipheae contain fibrosin bodies and the hyphae of Saprolegnieae cellulin bodies, but starch apparently never occurs. Invisible to the microscope, but rendered visible by reagents, are glycogen,Mucor, Ascomycetes, yeast, &c. In addition to these cell-contents we have good indirect evidence of the existence of large series of other bodies, such as proteids, carbohydrates, organic acids, alkaloids, enzymes, &c. These must not be confounded with the numerous substances obtained by chemical analysis of masses of the fungus, as there is often no proof of the manner of occurrence of such bodies, though we may conclude with a good show of probability that some of them also exist preformed in the living cell. Such are sugars (glucose, mannite, &c.), acids (acetic, citric and a whole series of lichen-acids), ethereal oils and resinous bodies, often combined with the intense colours of fungi and lichens, and a number of powerful alkaloid poisons, such as muscarin (Amanita), ergotin (Claviceps), &c.Among the enzymes already extracted from fungi areinvertases(yeasts, moulds, &c.), which split cane-sugar and other complex sugars with hydrolysis into simpler sugars such as dextrose and levulose;diastases, which convert starches into sugars (Aspergillus, &c.);cytases, which dissolve cellulose similarly (Botrytis, &c.);peptases, using the term as a general one for all enzymes which convert proteids into peptones and other bodies (Penicillium, &c.); lipases, which break up fatty oils (Empusa,Phycomyces, &c.); oxydases, which bring about the oxidations and changes of colour observed inBoletus, andzymase, extracted by Buchner from yeast, which brings about the conversion of sugar into alcohol and carbon-dioxide. That such enzymes are formed in the protoplasm is evident from the behaviour of hyphae, which have been observed to pierce cell-membranes, the chitinous coats of insects, artificial collodion films and layers of wax, &c. That a fungus can secrete more than one enzyme, according to the materials its hyphae have to attack, has been shown by the extraction of diastase, inulase, trehalase, invertase, maltase, raffinase, malizitase, emulsin, trypsin and lipase fromAspergillusby Bourquelot, and similar events occur in other fungi. The same fact is indicated by the wide range of organic substances which can be utilized byPenicilliumand other moulds, and by the behaviour of parasitic fungi which destroy various cell-contents and tissues. Many of the coloured pigments of fungi are fixed in the cell-walls or excreted to the outside (Peziza aeruginosa). Matruchot has used them for staining the living protoplasm of other fungi by growing the two together. Striking instances of coloured mycella are afforded byCorticium sanguineum, blood-red;Elaphomyces Leveillei, yellow-green;Chlorosplenium aeruginosum, verdigris green; and theDematei, brown or black.Nuclei.—Although many fungi have been regarded as devoid of nuclei, and all have not as yet been proved to contain them, the numerous investigations of recent years have revealed them in the cells of all forms thoroughly examined, and we are justified in concluding that the nucleus is as essential to the cell of a fungus as to that of other organisms. The hyphae of many contain numerous, even hundreds of nuclei (Phycomycetes); those of others have several (Aspergillus) in each segment, or only two (Exoascus) or one (Erysiphe) in each cell. Even the isolated cells of the yeast plant have each one nucleus. As a rule the nuclei of the mycelium are very minute (1.5-2 μ inPhycomyces), but those of many asci and spores are large and easily rendered visible. As with other plants, so in fungi the essential process of fertilization consists in the fusion of two nuclei, but owing to the absence of well-marked sexual organs from many fungi, a peculiar interest attaches to certain nuclear fusions in the vegetative cells or in young spores of many forms. Thus in Ustilagineae the chlamydospores, and in Uredineaethe teleutospores, each contain two nuclei when young, which fuse as the spores mature. In young asci a similar fusion of two nuclei occurs, and also in basidia, in each case the nucleus of the ascus or of the basidium resulting from the fusion subsequently giving rise by division to the nuclei of the ascospores and basidiospores respectively. The significance of these fusions will be discussed under the various groups. Nuclear division is usually accompanied by all the essential features of karyokinesis.Spores.—No agreement has ever been arrived at regarding the consistent use of the term spore. This is apparently owing to the facts that too much has been attempted in the definition, and that differences arise according as we aim at a morphological or a physiological definition. Physiologically, any cell or group of cells separated off from a hypha or unicellular fungus, and capable of itself growing out—germinating—to reproduce the fungus, is a spore; but it is evident that so wide a definition does not exclude the ordinary vegetative cells of sprouting fungi, such as yeasts, or small sclerotium like cell-aggregates of forms likeConiothecium. Morphologically considered, spores are marked by peculiarities of form, size, colour, place of origin, definiteness in number, mode of preparation, and so forth, such that they can be distinguished more or less sharply from the hyphae which produce them. The only physiological peculiarity exhibited in common by all spores is that they germinate and initiate the production of a new fungus-plant. Whether a spore results from the sexual union of two similar gametes (zygospore) or from the fertilization of an egg-cell by the protoplasm of a male organ (oospore); or is developed asexually as a motile (zoospore) or a quiescent body cut off from a hypha (conidium) or developed along its course (oidium or chlamydospore), or in its protoplasm (endospore), are matters of importance which have their uses in the classification and terminology of spores, though in many respects they are largely of academic interest.Fig.2.—Peronospora parasitica(De Bary). Conidiophore with conidia.Klebs has attempted to divide spores into three categories as follows: (1) kinospores, arising by relatively simple cell-divisions and subserving rapid dissemination and propagation,e.g.zoospores, conidia, endogonidia, stylospores, &c.; (2) paulospores, due to simple rearrangement of cell-contents, and subserving the persistence of the fungus through periods of exigency,e.g.gemmae, chlamydospores, resting-cells, cysts, &c.; (3) carpospores, produced by a more or less complex formative process, often in special fructifications, and subserving either or both multiplication and persistence,e.g.zygospores, oospores, brand-spores, aecidiospores, ascospores, basidiospores, &c. Little or nothing is gained by these definitions, however, which are especially physiological. In practice these various kinds of spores of fungi receive further special names in the separate groups, and names, moreover, which will appear, to those unacquainted with the history, to have been given without any consistency or regard to general principles; nevertheless, for ordinary purposes these names are far more useful in most cases, owing to their descriptive character, than the proposed new names, which have been only partially accepted.Sporophores.—In some of the simpler fungi the spores are not borne on or in hyphae which can be distinguished from the vegetative parts or mycelium, but in the vast majority of cases the sporogenous hyphae either ascend free into the air or radiate into the surrounding water as distinct branches, or are grouped into special columns, cushions, layers or complex masses obviously different in colour, consistency, shape and other characters from the parts which gather up and assimilate the food-materials. The term “receptacle” sometimes applied to these spore-bearing hyphae is better replaced by sporophore. The sporophore is obsolete when the spore-bearing hyphae are not sharply distinct from the mycelium, simple when the constituent hyphae are isolated, and compound when the latter are conjoined. The chief distinctive characters of the sporogenous hyphae are their orientation, usually vertical; their limited apical growth; their peculiar branching, form, colour, contents, consistency; and their spore-production. According to the characters of the last, we might theoretically divide them into conidiophores, sporangiophores, gametophores, oidiophores, &c.; but since the two latter rarely occur, and more than one kind of spore or spore-case may occur on a sporophore, it is impossible to carry such a scheme fully into practice.A simple sporophore may be merely a single short hypha, the end of which stops growing and becomes cut off as a conidium by the formation of a septum, which then splits and allows the conidium to fall. More generally the hypha below the septum grows forwards again, and repeats this process several times before the terminal conidium falls, and so a chain of conidia results, the oldest of which terminates the series (Erysiphe); when the primary branch has thus formed a basipetal series, branches may arise from below and again repeat this process, thus forming a tuft (Penicillium). Or the primary hypha may first swell at its apex, and put forth a series of short peg-like branches (sterigmata) from the increased surface thus provided, each of which develops a similar basipetal chain of conidia (Aspergillus), and various combinations of these processes result in the development of numerous varieties of exquisitely branched sporophores of this type (Botrytis,Botryosporium,Verticillium, &c.).Fig. 3.—Cystopus candidus.A.a, Conidia.b, Conidiophores.c, Conidium emitting zoospores.d, Free zoospore.B.og, Oogonium.os, Oosphere.an, Antheridium.C. Formation of zoospores by oospores.z, Free zoospores.(After De Bary.)A second type is developed as follows: the primary hypha forms a septum below its apex as before, and the terminal conidium, thus abstricted, puts out a branch at its apex, which starts as a mere point and rapidly swells to a second conidium; this repeats the process, and so on, so that we now have a chain of conidia developed in acropetal succession, the oldest being below, and, as inPenicillium, &c., branches put forth lower down may repeat the process (Hormodendron). In all these cases we may speak of simple conidiophores. The simple sporophore does not necessarily terminate in conidia, however. InMucor, for example, the end of the primary hypha swells into a spheroidal head (sporangium), the protoplasm of which undergoes segmentation into more or less numerous globular masses, each of which secretes an enveloping cell-wall and becomes a spore (endospore), and branched systems of sporangia may arise as before (Thamnidium). Such may be termed sporangiophores. InSporodiniathe branches give rise also to short branches, which meet and fuse their contents to form zygospores. In Peronospora, Saprolegnia, &c., the ends of the branches swell up into sporangia, which develop zoospores in their interior (zoosporangia), or their contents become oospheres, which may be fertilized by the contents of other branches (antheridia) and so form egg-cases (oogonia). Since in such cases the sporophore bears sexual cells, they may be conveniently termed gametophores.Compound sporophores arise when any of the branched or unbranched types of spore-bearing hyphae described above ascend into the air in consort, and are more or less crowded into definite layers, cushions, columns or other complex masses. The same laws apply to the individual hyphae and their branches as to simple sporophores, and as long as the conidia, sporangia, gametes, &c., are borne on their external surfaces, it is quite consistent to speak of these as compound sporophores, &c., in the sense described, however complex they may become. Among the simplest cases are the sheet-like aggregates of sporogenous hyphae inPuccinia,Uromyces, &c., or of basidia inExobasidium,Corticium, &c., or of asci inExoascus,Ascocorticium, &c. In the former, where the layer is small, it is often termed a sorus, but where, as in the latter, the sporogenous layer is extensive, and spread out more or less sheet-like on the supporting tissues, it is more frequently termed a hymenium. Another simple case is that of the columnar aggregates of sporogenous hyphae in forms likeStilbum,Coremium, &c. These leadus to cases where the main mass of the sporophore forms a supporting tissue of closely crowded or interwoven hyphae, the sporogenous terminal parts of the hyphae being found at the periphery or apical regions only. Here we have the cushion-like type (stroma) ofNectriaand many Pyrenomycetes, the clavate “receptacle” ofClavaria, &c., passing into the complex forms met with inSparassis,Xylaria,Polyporei, andAgaricini, &c. In these cases the compound sporophore is often termed the hymenophore, and its various parts demand special names (pileus, stipes, gills, pores, &c.) to denote peculiarities of distribution of the hymenium over the surface.Other series of modifications arise in which the tissues corresponding to the stroma invest the sporogenous hyphal ends, and thus enclose the spores, asci, basidia, &c., in a cavity. In the simplest case the stroma, after bearing its crop of conidia or oidia, develops ascogenous branches in the loosened meshes of its interior (e.g.Onygena). Another simple case is where the plane or slightly convex surface of the stroma rises at its margins and overgrows the sporogenous hyphal ends, so that the spores, asci, &c., come to lie in the depression of a cavity—e.g.Solenia,Cyphella—and even simpler cases are met with inMortierella, where the zygospore is invested by the overgrowth of a dense mat of closely branching hyphae, and inGymnoascus, where a loose mat of similarly barren hyphae covers in the tufts of asci as they develop.In such examples as the above we may regard the hymenium (Solenia,Cyphella), zygospores, or asci as truly invested by later growth, but in the vast majority of cases the processes which result in the enclosure of the spores, asci, &c., in a “fructification” are much more involved, inasmuch as the latter is developed in the interior of hyphal tissues, which are by no means obviously homologous with a stroma. Thus inPenicillium,Eurotium,Erysiphe, &c., hyphal ends which are the initials of ascogenous branches, are invested by closely packed branches at an early stage of development, and the asci develop inside what has by that time become a complete investment. Whether a true sexual process precedes these processes or not does not affect the present question, the point being that the resulting spheroidal “fructification” (cleistocarp, perithecium) has a definite wall of its own not directly comparable with a stroma. In other cases (Hypomyces,Nectria) the perithecia arise on an already mature stroma, while yet more numerous examples can be given (Poronia,Hypoxylon,Claviceps, &c.) where the perithecia originate below the surface of a stroma formed long before. Similarly with the various types of conidial or oidial “fructifications,” termed pycnidia, spermogonia, aecidia, &c. In the simplest of these cases—e.g.Fumago—a single mycelial cell divides by septa in all three planes until a more or less solid clump results. Then a hollow appears in the centre owing to the more rapid extension of the outer parts, and into this hollow the cells lining it put forth short sporogenous branches, from the tips of which the spores (stylospores, conidia, spermatia) are abstricted. In a similar way are developed the pycnidia ofCicinnobolus,Pleospora,Cucurbitaria,Leptosphaeriaand others. In other cases (Diplodia,Aecidium, &c.) conidial or oidial “fructifications” arise by a number of hyphae interweaving themselves into a knot, as if they were forming a Sclerotium. The outer parts of the mass then differentiate as a wall or investment, and the interior becomes a hollow, into which hyphal ends grow and abstrict the spores. Much more complicated are the processes in a large series of “fructifications,” where the mycelium first develops a densely packed mass of hyphae, all alike, in which labyrinths of cavities subsequently form by separation of hyphae in the previously homogeneous mass, and the hymenium covers the walls of these cavities and passages as with a lining layer. Meanwhile differences in consistency appear in various strata, and a dense outer protective layer (peridium), soft gelatinous layers, and so on are formed, the whole eventually attaining great complexity—e.g.puff-balls, earth-stars and variousPhalloideae.Spore-Distribution.—Ordinary conidia and similarly abstricted dry spores are so minute, light and numerous that their dispersal is ensured by any current of air or water, and we also know that rats and other burrowing animals often carry them on their fur; similarly with birds, insects, slugs, worms, &c., on claws, feathers, proboscides, &c., or merely adherent to the slimy body. In addition to these accidental modes of dispersal, however, there is a series of interesting adaptations on the part of the fungus itself. Passing over the locomotor activity of zoospores (Pythium,Peronospora,Saprolegnia) we often find spores held under tension in sporangia (Pilobolus) or in asci (Peziza) until ripe, and then forcibly shot out by the sudden rupture of the sporangial wall under the pressure of liquid behind—mechanism comparable to that of a pop-gun, if we suppose air replaced by watery sap. Even a single conidium, held tense to the last moment by the elastic cell-wall, may be thus shot forward by a spurt of liquid under pressure in the hypha abstricting it (e.g.Empusa), and similarly withbasidiospores(Coprinus,Agaricus, &c.). A more complicated case is illustrated bySphaerobolus, where the entire mass of spores, enclosed in its own peridium, is suddenly shot up into the air like a bomb from a mortar by the elastic retroversion of a peculiar layer which, up to the last moment, surrounded the bomb, and then suddenly splits above, turns inside out, and drives the former as a projectile from a gun. Gelatinous or mucilaginous degenerations of cell-walls are frequently employed in the interests of spore dispersal. The mucilage surrounding endospores ofMucor, conidia ofEmpusa, &c., serves to gum the spore to animals. Such gums are formed abundantly in pycnidia, and, absorbing water, swell and carry out the spores in long tendrils, which emerge for days and dry as they reach the air, the glued spores gradually being set free by rain, wind, &c. In oidial chains (Sclerotinia) a minute double wedge of wall-substance arises in the middle lamella between each pair of contiguous oidia, and by its enlargement splits the separating lamella. These disjunctors serve as points of application for the elastic push of the swelling spore-ends, and as the connecting outer lamella of cell-wall suddenly gives way, the spores are jerked asunder. In many cases the slimy masses of spermatia (Uredineae), conidia (Claviceps), basidiospores (Phallus,Coprinus), &c., emit more or less powerful odours, which attract flies or other insects, and it has been shown that bees carry the fragrant oidia ofSclerotiniato the stigma ofVacciniumand infect it, and that flies carry away the foetid spores ofPhallus, just as pollen is dispersed by such insects. Whether the strong odour of trimethylamine evolved by the spores ofTilletiaattracts insects is not known.The recent observations and exceedingly ingenious experiments of Falck have shown that the sporophores of the Basidiomycetes—especially the large sporophores of such forms asBoletus,Polyporus—contain quantities of reserve combustible material which are burnt up by the active metabolism occurring when the fruit-body is ripe. By this means the temperature of the sporophore is raised and the difference between it and the surrounding air may be one of several degrees. As a result convection currents are produced in the air which are sufficient to catch the basidiospores in their fall and carry them, away from the regions of comparative atmospheric stillness near the ground, to the upper air where more powerful air-currents can bring about their wide distribution.

Many fungi (Phallus,Agaricus,Fumago, &c.) when strongly growing put out ribbon-like or cylindrical cords, or sheet-like mycelial plates of numerous parallel hyphae, all growing together equally, and fusing by anastomoses, and in this way extend long distances in the soil, or over the surfaces of leaves, branches, &c. These mycelial strands may be white and tender, or the outer hyphae may be hard and black, and very often the resemblance of the subterranean forms to a root is so marked that they are termed rhizomorphs. The outermost hyphae may even put forth thinner hyphae, radiating into the soil like root-hairs, and the convergent tips may be closely appressed and so divided by septa as to resemble the root-apex of a higher plant (Armillaria mellea).

Sclerotia.—Fungi, like other plants, are often found to store up large quantities of reserve materials (oil, glycogen, carbohydrates, &c.) in special parts of their vegetative tissues, where they lie accumulated between a period of active assimilation and one of renewed activity, forming reserves to be consumed particularly during the formation of large fructifications. These reserve stores may be packed away in single hyphae or in swollen cells, but the hyphae containing them are often gathered into thick cords or mycelial strands (Phallus, mushroom, &c.), or flattened and anastomosing ribbons and plates, often containing several kinds of hyphae (Merulius lacrymans). In other cases the strands undergo differentiation into an outer layer with blackened, hardened cell-walls and a core of ordinary hyphae, and are then termed rhizomorphs (Armillaria mellea), capable not only of extending the fungus in the soil, like roots, but also of lying dormant, protected by the outer casing. Such aggregations of hyphae frequently become knotted up into dense masses of interwoven and closely packed hyphae, varying in size from that of a pin’s head or a pea (Peziza,Coprinus) to that of a man’s fist or head, and weighing 10 to 25 ℔ or more (Polyporus Mylittae,P. tumulosus,Lentinus Woermanni,P. Sapurema, &c.). The interwoven hyphae fuse and branch copiously, filling up all interstices. They also undergo cutting up by numerous septa into short cells, and these often divide again in all planes, so that a pseudoparenchyma results, the walls of which may be thickened and swollen internally, or hardened and black on the exterior. In many cases the swollen cell-walls serve as reserves, and sometimes the substance is so thickly deposited in strata as to obliterate the lumen, and the hyphae become nodular (Polyporus sacer,P. rhinoceros,Lentinus Woermanni). The various sclerotia, if kept moist, give rise to the fructifications of the fungi concerned, much as a potato tuber does to a potato plant, and in the same way the reserve materials are consumed. They are principally Polyporei, Agaricini, Pezizae; none are known among the Phycomycetes, Uredineae or Ustilagineae. The functions of mycelial strands, rhizomorphs and sclerotia are not only to collect and store materials, but also to extend the fungus, and in many cases similar strands act as organs of attack. The same functions of storage in advance of fructification are also exercised by the stromata so common in Ascomycetes.

Tissue Differentiations.—The simpler mycelia consist of hyphae all alike and thin-walled, or merely differing in the diameter of the branches of various orders, or in their relations to the environment, some plunging into the substratum like roots, others remaining on its surface, and others (aerial hyphae) rising into the air. Such hyphae may be multicellular, or they may consist of simple tubes with numerous nuclei and no septa (Phycomycetes), and are then non-cellular. In the more complex tissue-bodies of higher fungi, however, we find considerable differences in the various layers or strands of hyphae.

An epidermis-like or cortical protective outer layer is very common, and is usually characterized by the close septation of the densely interwoven hyphae and the thickening and dark colour of their outer walls (sclerotia,Xylaria, &c.). Fibre-like hyphae with the lumen almost obliterated by the thick walls occur in mycelial cords (Merulius). Latex-tubes abound in the tissues ofLactarius,Stereum,Mycena,Fistulina, filled with white or coloured milky fluids, and Istvanffvi has shown that similar tubes with fluid or oily contents are widely spread in other Hymenomycetes. Sometimes fatty oil or watery sap is found in swollen hyphal ends, or such tubes contain coloured sap. Cystidia and paraphyses may be also classed here. InMerulius lacrymansHartig has observed thin-walled hyphae with large lumina, the septa of which are perforated like those of sieve-tubes.

As regards its composition, the cell-wall of fungi exhibits variations of the same kind as those met with in higher plants. While the fundamental constituent is a cellulose in many Mucorini and other Phycomycetes, in others bodies like pectose, callose, &c., commonly occur, and Wisselingh’s researches show that chitin, a gluco-proteid common in animals, forms the main constituent in many cases, and is probably deposited directly as such, though, like the other substances, it may be mixed with cellulose. As in other cell-walls, so here the older membranes may be altered by deposits of various substances, such as resin, calcium oxalate, colouring matters; or more profoundly altered throughout, or in definite layers, by lignification, suberization (Trametes,Daedalea), or swelling to a gelatinous mucilage (Tremella,Gymnosporangium), while cutinization of the outer layers is common. One of the most striking alterations of cell-walls is that termedcarbonization, in which the substance gradually turns black, hard and brittle, as if charred—e.g.Xylaria,Ustulina, some sclerotia. At the other extreme the cell-walls of many lichen-fungi are soft and colourless, but turn blue in iodine, as does starch. The young cell-wall is always tenuous and flexible, and may remain so throughout, but in many cases thickenings and structural differentiations, as well as the changes referred to above, alter the primary wall considerably. Such thickening may be localized, andpits(e.g.Uredospores, septa of Basidiomycetes),spirals,reticulations,rings, &c. (capillitium fibres ofPodaxon,Calostoma,Battarrea), occur as in the vessels of higher plants, while sculptured networks, pittings and so forth are as common on fungus-spores as they are on pollen grains.

Cell-Contents.—The cells of fungi, in addition to protoplasm, nuclei and sap-vacuoles, like other vegetable cells, contain formed and amorphous bodies of various kinds. Among those directly visible to the microscope are oil drops, often coloured (Uredineae) crystals of calcium oxalate (Phallus,Russula), proteid crystals (Mucor,Pilobolus, &c.) and resin (Polyporei). The oidia of Erysipheae contain fibrosin bodies and the hyphae of Saprolegnieae cellulin bodies, but starch apparently never occurs. Invisible to the microscope, but rendered visible by reagents, are glycogen,Mucor, Ascomycetes, yeast, &c. In addition to these cell-contents we have good indirect evidence of the existence of large series of other bodies, such as proteids, carbohydrates, organic acids, alkaloids, enzymes, &c. These must not be confounded with the numerous substances obtained by chemical analysis of masses of the fungus, as there is often no proof of the manner of occurrence of such bodies, though we may conclude with a good show of probability that some of them also exist preformed in the living cell. Such are sugars (glucose, mannite, &c.), acids (acetic, citric and a whole series of lichen-acids), ethereal oils and resinous bodies, often combined with the intense colours of fungi and lichens, and a number of powerful alkaloid poisons, such as muscarin (Amanita), ergotin (Claviceps), &c.

Among the enzymes already extracted from fungi areinvertases(yeasts, moulds, &c.), which split cane-sugar and other complex sugars with hydrolysis into simpler sugars such as dextrose and levulose;diastases, which convert starches into sugars (Aspergillus, &c.);cytases, which dissolve cellulose similarly (Botrytis, &c.);peptases, using the term as a general one for all enzymes which convert proteids into peptones and other bodies (Penicillium, &c.); lipases, which break up fatty oils (Empusa,Phycomyces, &c.); oxydases, which bring about the oxidations and changes of colour observed inBoletus, andzymase, extracted by Buchner from yeast, which brings about the conversion of sugar into alcohol and carbon-dioxide. That such enzymes are formed in the protoplasm is evident from the behaviour of hyphae, which have been observed to pierce cell-membranes, the chitinous coats of insects, artificial collodion films and layers of wax, &c. That a fungus can secrete more than one enzyme, according to the materials its hyphae have to attack, has been shown by the extraction of diastase, inulase, trehalase, invertase, maltase, raffinase, malizitase, emulsin, trypsin and lipase fromAspergillusby Bourquelot, and similar events occur in other fungi. The same fact is indicated by the wide range of organic substances which can be utilized byPenicilliumand other moulds, and by the behaviour of parasitic fungi which destroy various cell-contents and tissues. Many of the coloured pigments of fungi are fixed in the cell-walls or excreted to the outside (Peziza aeruginosa). Matruchot has used them for staining the living protoplasm of other fungi by growing the two together. Striking instances of coloured mycella are afforded byCorticium sanguineum, blood-red;Elaphomyces Leveillei, yellow-green;Chlorosplenium aeruginosum, verdigris green; and theDematei, brown or black.

Nuclei.—Although many fungi have been regarded as devoid of nuclei, and all have not as yet been proved to contain them, the numerous investigations of recent years have revealed them in the cells of all forms thoroughly examined, and we are justified in concluding that the nucleus is as essential to the cell of a fungus as to that of other organisms. The hyphae of many contain numerous, even hundreds of nuclei (Phycomycetes); those of others have several (Aspergillus) in each segment, or only two (Exoascus) or one (Erysiphe) in each cell. Even the isolated cells of the yeast plant have each one nucleus. As a rule the nuclei of the mycelium are very minute (1.5-2 μ inPhycomyces), but those of many asci and spores are large and easily rendered visible. As with other plants, so in fungi the essential process of fertilization consists in the fusion of two nuclei, but owing to the absence of well-marked sexual organs from many fungi, a peculiar interest attaches to certain nuclear fusions in the vegetative cells or in young spores of many forms. Thus in Ustilagineae the chlamydospores, and in Uredineaethe teleutospores, each contain two nuclei when young, which fuse as the spores mature. In young asci a similar fusion of two nuclei occurs, and also in basidia, in each case the nucleus of the ascus or of the basidium resulting from the fusion subsequently giving rise by division to the nuclei of the ascospores and basidiospores respectively. The significance of these fusions will be discussed under the various groups. Nuclear division is usually accompanied by all the essential features of karyokinesis.

Spores.—No agreement has ever been arrived at regarding the consistent use of the term spore. This is apparently owing to the facts that too much has been attempted in the definition, and that differences arise according as we aim at a morphological or a physiological definition. Physiologically, any cell or group of cells separated off from a hypha or unicellular fungus, and capable of itself growing out—germinating—to reproduce the fungus, is a spore; but it is evident that so wide a definition does not exclude the ordinary vegetative cells of sprouting fungi, such as yeasts, or small sclerotium like cell-aggregates of forms likeConiothecium. Morphologically considered, spores are marked by peculiarities of form, size, colour, place of origin, definiteness in number, mode of preparation, and so forth, such that they can be distinguished more or less sharply from the hyphae which produce them. The only physiological peculiarity exhibited in common by all spores is that they germinate and initiate the production of a new fungus-plant. Whether a spore results from the sexual union of two similar gametes (zygospore) or from the fertilization of an egg-cell by the protoplasm of a male organ (oospore); or is developed asexually as a motile (zoospore) or a quiescent body cut off from a hypha (conidium) or developed along its course (oidium or chlamydospore), or in its protoplasm (endospore), are matters of importance which have their uses in the classification and terminology of spores, though in many respects they are largely of academic interest.

Klebs has attempted to divide spores into three categories as follows: (1) kinospores, arising by relatively simple cell-divisions and subserving rapid dissemination and propagation,e.g.zoospores, conidia, endogonidia, stylospores, &c.; (2) paulospores, due to simple rearrangement of cell-contents, and subserving the persistence of the fungus through periods of exigency,e.g.gemmae, chlamydospores, resting-cells, cysts, &c.; (3) carpospores, produced by a more or less complex formative process, often in special fructifications, and subserving either or both multiplication and persistence,e.g.zygospores, oospores, brand-spores, aecidiospores, ascospores, basidiospores, &c. Little or nothing is gained by these definitions, however, which are especially physiological. In practice these various kinds of spores of fungi receive further special names in the separate groups, and names, moreover, which will appear, to those unacquainted with the history, to have been given without any consistency or regard to general principles; nevertheless, for ordinary purposes these names are far more useful in most cases, owing to their descriptive character, than the proposed new names, which have been only partially accepted.

Sporophores.—In some of the simpler fungi the spores are not borne on or in hyphae which can be distinguished from the vegetative parts or mycelium, but in the vast majority of cases the sporogenous hyphae either ascend free into the air or radiate into the surrounding water as distinct branches, or are grouped into special columns, cushions, layers or complex masses obviously different in colour, consistency, shape and other characters from the parts which gather up and assimilate the food-materials. The term “receptacle” sometimes applied to these spore-bearing hyphae is better replaced by sporophore. The sporophore is obsolete when the spore-bearing hyphae are not sharply distinct from the mycelium, simple when the constituent hyphae are isolated, and compound when the latter are conjoined. The chief distinctive characters of the sporogenous hyphae are their orientation, usually vertical; their limited apical growth; their peculiar branching, form, colour, contents, consistency; and their spore-production. According to the characters of the last, we might theoretically divide them into conidiophores, sporangiophores, gametophores, oidiophores, &c.; but since the two latter rarely occur, and more than one kind of spore or spore-case may occur on a sporophore, it is impossible to carry such a scheme fully into practice.

A simple sporophore may be merely a single short hypha, the end of which stops growing and becomes cut off as a conidium by the formation of a septum, which then splits and allows the conidium to fall. More generally the hypha below the septum grows forwards again, and repeats this process several times before the terminal conidium falls, and so a chain of conidia results, the oldest of which terminates the series (Erysiphe); when the primary branch has thus formed a basipetal series, branches may arise from below and again repeat this process, thus forming a tuft (Penicillium). Or the primary hypha may first swell at its apex, and put forth a series of short peg-like branches (sterigmata) from the increased surface thus provided, each of which develops a similar basipetal chain of conidia (Aspergillus), and various combinations of these processes result in the development of numerous varieties of exquisitely branched sporophores of this type (Botrytis,Botryosporium,Verticillium, &c.).

A.a, Conidia.

b, Conidiophores.

c, Conidium emitting zoospores.

d, Free zoospore.

B.og, Oogonium.

os, Oosphere.

an, Antheridium.

C. Formation of zoospores by oospores.

z, Free zoospores.

(After De Bary.)

A second type is developed as follows: the primary hypha forms a septum below its apex as before, and the terminal conidium, thus abstricted, puts out a branch at its apex, which starts as a mere point and rapidly swells to a second conidium; this repeats the process, and so on, so that we now have a chain of conidia developed in acropetal succession, the oldest being below, and, as inPenicillium, &c., branches put forth lower down may repeat the process (Hormodendron). In all these cases we may speak of simple conidiophores. The simple sporophore does not necessarily terminate in conidia, however. InMucor, for example, the end of the primary hypha swells into a spheroidal head (sporangium), the protoplasm of which undergoes segmentation into more or less numerous globular masses, each of which secretes an enveloping cell-wall and becomes a spore (endospore), and branched systems of sporangia may arise as before (Thamnidium). Such may be termed sporangiophores. InSporodiniathe branches give rise also to short branches, which meet and fuse their contents to form zygospores. In Peronospora, Saprolegnia, &c., the ends of the branches swell up into sporangia, which develop zoospores in their interior (zoosporangia), or their contents become oospheres, which may be fertilized by the contents of other branches (antheridia) and so form egg-cases (oogonia). Since in such cases the sporophore bears sexual cells, they may be conveniently termed gametophores.

Compound sporophores arise when any of the branched or unbranched types of spore-bearing hyphae described above ascend into the air in consort, and are more or less crowded into definite layers, cushions, columns or other complex masses. The same laws apply to the individual hyphae and their branches as to simple sporophores, and as long as the conidia, sporangia, gametes, &c., are borne on their external surfaces, it is quite consistent to speak of these as compound sporophores, &c., in the sense described, however complex they may become. Among the simplest cases are the sheet-like aggregates of sporogenous hyphae inPuccinia,Uromyces, &c., or of basidia inExobasidium,Corticium, &c., or of asci inExoascus,Ascocorticium, &c. In the former, where the layer is small, it is often termed a sorus, but where, as in the latter, the sporogenous layer is extensive, and spread out more or less sheet-like on the supporting tissues, it is more frequently termed a hymenium. Another simple case is that of the columnar aggregates of sporogenous hyphae in forms likeStilbum,Coremium, &c. These leadus to cases where the main mass of the sporophore forms a supporting tissue of closely crowded or interwoven hyphae, the sporogenous terminal parts of the hyphae being found at the periphery or apical regions only. Here we have the cushion-like type (stroma) ofNectriaand many Pyrenomycetes, the clavate “receptacle” ofClavaria, &c., passing into the complex forms met with inSparassis,Xylaria,Polyporei, andAgaricini, &c. In these cases the compound sporophore is often termed the hymenophore, and its various parts demand special names (pileus, stipes, gills, pores, &c.) to denote peculiarities of distribution of the hymenium over the surface.

Other series of modifications arise in which the tissues corresponding to the stroma invest the sporogenous hyphal ends, and thus enclose the spores, asci, basidia, &c., in a cavity. In the simplest case the stroma, after bearing its crop of conidia or oidia, develops ascogenous branches in the loosened meshes of its interior (e.g.Onygena). Another simple case is where the plane or slightly convex surface of the stroma rises at its margins and overgrows the sporogenous hyphal ends, so that the spores, asci, &c., come to lie in the depression of a cavity—e.g.Solenia,Cyphella—and even simpler cases are met with inMortierella, where the zygospore is invested by the overgrowth of a dense mat of closely branching hyphae, and inGymnoascus, where a loose mat of similarly barren hyphae covers in the tufts of asci as they develop.

In such examples as the above we may regard the hymenium (Solenia,Cyphella), zygospores, or asci as truly invested by later growth, but in the vast majority of cases the processes which result in the enclosure of the spores, asci, &c., in a “fructification” are much more involved, inasmuch as the latter is developed in the interior of hyphal tissues, which are by no means obviously homologous with a stroma. Thus inPenicillium,Eurotium,Erysiphe, &c., hyphal ends which are the initials of ascogenous branches, are invested by closely packed branches at an early stage of development, and the asci develop inside what has by that time become a complete investment. Whether a true sexual process precedes these processes or not does not affect the present question, the point being that the resulting spheroidal “fructification” (cleistocarp, perithecium) has a definite wall of its own not directly comparable with a stroma. In other cases (Hypomyces,Nectria) the perithecia arise on an already mature stroma, while yet more numerous examples can be given (Poronia,Hypoxylon,Claviceps, &c.) where the perithecia originate below the surface of a stroma formed long before. Similarly with the various types of conidial or oidial “fructifications,” termed pycnidia, spermogonia, aecidia, &c. In the simplest of these cases—e.g.Fumago—a single mycelial cell divides by septa in all three planes until a more or less solid clump results. Then a hollow appears in the centre owing to the more rapid extension of the outer parts, and into this hollow the cells lining it put forth short sporogenous branches, from the tips of which the spores (stylospores, conidia, spermatia) are abstricted. In a similar way are developed the pycnidia ofCicinnobolus,Pleospora,Cucurbitaria,Leptosphaeriaand others. In other cases (Diplodia,Aecidium, &c.) conidial or oidial “fructifications” arise by a number of hyphae interweaving themselves into a knot, as if they were forming a Sclerotium. The outer parts of the mass then differentiate as a wall or investment, and the interior becomes a hollow, into which hyphal ends grow and abstrict the spores. Much more complicated are the processes in a large series of “fructifications,” where the mycelium first develops a densely packed mass of hyphae, all alike, in which labyrinths of cavities subsequently form by separation of hyphae in the previously homogeneous mass, and the hymenium covers the walls of these cavities and passages as with a lining layer. Meanwhile differences in consistency appear in various strata, and a dense outer protective layer (peridium), soft gelatinous layers, and so on are formed, the whole eventually attaining great complexity—e.g.puff-balls, earth-stars and variousPhalloideae.

Spore-Distribution.—Ordinary conidia and similarly abstricted dry spores are so minute, light and numerous that their dispersal is ensured by any current of air or water, and we also know that rats and other burrowing animals often carry them on their fur; similarly with birds, insects, slugs, worms, &c., on claws, feathers, proboscides, &c., or merely adherent to the slimy body. In addition to these accidental modes of dispersal, however, there is a series of interesting adaptations on the part of the fungus itself. Passing over the locomotor activity of zoospores (Pythium,Peronospora,Saprolegnia) we often find spores held under tension in sporangia (Pilobolus) or in asci (Peziza) until ripe, and then forcibly shot out by the sudden rupture of the sporangial wall under the pressure of liquid behind—mechanism comparable to that of a pop-gun, if we suppose air replaced by watery sap. Even a single conidium, held tense to the last moment by the elastic cell-wall, may be thus shot forward by a spurt of liquid under pressure in the hypha abstricting it (e.g.Empusa), and similarly withbasidiospores(Coprinus,Agaricus, &c.). A more complicated case is illustrated bySphaerobolus, where the entire mass of spores, enclosed in its own peridium, is suddenly shot up into the air like a bomb from a mortar by the elastic retroversion of a peculiar layer which, up to the last moment, surrounded the bomb, and then suddenly splits above, turns inside out, and drives the former as a projectile from a gun. Gelatinous or mucilaginous degenerations of cell-walls are frequently employed in the interests of spore dispersal. The mucilage surrounding endospores ofMucor, conidia ofEmpusa, &c., serves to gum the spore to animals. Such gums are formed abundantly in pycnidia, and, absorbing water, swell and carry out the spores in long tendrils, which emerge for days and dry as they reach the air, the glued spores gradually being set free by rain, wind, &c. In oidial chains (Sclerotinia) a minute double wedge of wall-substance arises in the middle lamella between each pair of contiguous oidia, and by its enlargement splits the separating lamella. These disjunctors serve as points of application for the elastic push of the swelling spore-ends, and as the connecting outer lamella of cell-wall suddenly gives way, the spores are jerked asunder. In many cases the slimy masses of spermatia (Uredineae), conidia (Claviceps), basidiospores (Phallus,Coprinus), &c., emit more or less powerful odours, which attract flies or other insects, and it has been shown that bees carry the fragrant oidia ofSclerotiniato the stigma ofVacciniumand infect it, and that flies carry away the foetid spores ofPhallus, just as pollen is dispersed by such insects. Whether the strong odour of trimethylamine evolved by the spores ofTilletiaattracts insects is not known.

The recent observations and exceedingly ingenious experiments of Falck have shown that the sporophores of the Basidiomycetes—especially the large sporophores of such forms asBoletus,Polyporus—contain quantities of reserve combustible material which are burnt up by the active metabolism occurring when the fruit-body is ripe. By this means the temperature of the sporophore is raised and the difference between it and the surrounding air may be one of several degrees. As a result convection currents are produced in the air which are sufficient to catch the basidiospores in their fall and carry them, away from the regions of comparative atmospheric stillness near the ground, to the upper air where more powerful air-currents can bring about their wide distribution.

Classification.—It has been accepted for some time now that the majority of the fungi proper fall into three main groups, the Phycomycetes, Ascomycetes and Basidiomycetes, the Schizomycetes and Myxomycetes (Mycetozoa) being considered as independent groups not coming under the true fungi.

The chief schemes of classification put forward in detail have been those of P.A. Saccardo (1882-1892), of Oskar Brefeld and Von Tavel (1892), of P.E.L. Van Tieghem (1893) and of J. Schroeter (1892). The scheme of Brefeld, which was based on the view that the Ascomycetes and Basidiomycetes were completely asexual and that these two groups had been derived from one division (Zygomycetes) of the Phycomycetes, has been very widely accepted. The recent work of the last twelve years has shown, however, that the two higher groups of fungi exhibit distinct sexuality, of either a normal or reduced type, and has also rendered very doubtful the view of the origin of these two groups from the Phycomycetes. The real difficulty of classification of the fungi lies in the polyphyletic nature of the group. There is very little doubt that the primitive fungi have been derived by degradation from the lower algae. It appears, however, that such a degradation has occurred not only once in evolution but on several occasions, so that we have in the Phycomycetes not a series of naturally related forms, but groups which have arisen perfectly independently of one another from various groups of the algae. It is also possible in the absence of satisfactory intermediate forms that the Ascomycetes and Basidiomycetes have also been derived from the algae independently of the Phycomycetes, and perhaps of one another.

A natural classification on these lines would obviously be very complicated, so that in the present state of our knowledge it will be best to retain the three main groups mentioned above, bearing in mind that the Phycomycetes especially are far from being a natural group. The following gives a tabular survey of the scheme adopted in the present article:

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