Chapter 3 of 11 · 19938 words · ~100 min read

Part II

. pp. ix.-xii. (E. C. B.)

ANTHONY OF PADUA, SAINT (1195-1231), the most celebrated of the followers of Saint Francis of Assisi, was born at Lisbon on the 15th of August 1195. In his fifteenth year he entered the Augustinian order, and subsequently joined the Franciscans in 1220. He wished to devote himself to missionary labours in North Africa, but the ship in which he sailed was cast by a storm on the coast of Sicily, whence he made his way to Italy. He taught theology at Bologna, Toulouse, Montpellier and Padua, and won a great reputation as a preacher throughout Italy. He was the leader of the rigorous party in the Franciscan order against the mitigations introduced by the general Elias. His death took place at the convent of Ara Coeli, near Padua, on the 13th of June 1231. He was canonized by Gregory IX. in the following year, and his festival is kept on the 13th of June. He is regarded as the patron saint of Padua and of Portugal, and is appealed to by devout clients for finding lost objects. The meagre accounts of his life which we possess have been supplemented by numerous popular legends, which represent him as a continuous worker of miracles, and describe his marvellous eloquence by pictures of fishes leaping out of the water to hear him. There are many confraternities established in his honour throughout Christendom, and the number of "pious" biographies devoted to him would fill many volumes.

The most trustworthy modern works are by A. Lepitre, _St Antoine de Padoue_ (Paris, 1902, in _Les Saints_ series: good bibliography; Eng. trans. by Edith Guest, London, 1902), and by Leopold de Cherance, _St Antoine de Padoue_ (Paris, 1895; Eng. trans., London, 1896). His works, consisting of sermons and a mystical commentary on the Bible, were published in an appendix to those of St Francis, in the _Annales Minorum_ of Luke Wadding (Antwerp, 1623), and are also reproduced by Horoy, _Medii aevi bibliotheca patristica_ (1880, vi. pp. 555 et sqq.); see art. "Antonius von Padua" in Herzog-Hauck, _Realencyklopadie_.

ANTHONY, SUSAN BROWNELL (1820-1906), American reformer, was born at Adams, Massachusetts, on the 15th of February 1820, the daughter of Quakers. Soon after her birth, her family moved to the state of New York, and after 1845 she lived in Rochester. She received her early education in a school maintained by her father for his own and neighbours' children, and from the time she was seventeen until she was thirty-two she taught in various schools. In the decade preceding the outbreak of the Civil War she took a prominent part in the anti-slavery and temperance movements in New York, organizing in 1852 the first woman's state temperance society in America, and in 1856 becoming the agent for New York state of the American Anti-slavery Society. After 1854 she devoted herself almost exclusively to the agitation for woman's rights, and became recognized as one of the ablest and most zealous advocates, both as a public speaker and as a writer, of the complete legal equality of the two sexes. From 1868 to 1870 she was the proprietor of a weekly paper, _The Revolution_, published in New York, edited by Mrs Elizabeth Cady Stanton, and having for its motto, "The true republic--men, their rights and nothing more; women, their rights and nothing less." She was vice-president-at-large of the National Woman's Suffrage Association from the date of its organization in 1869 until 1892, when she became president. For casting a vote in the presidential election of 1872, as, she asserted, the Fourteenth Amendment to the Federal Constitution entitled her to do, she was arrested and fined $100, but she never paid the fine. In collaboration with Mrs Elizabeth Cady Stanton, Mrs Matilda Joslyn Gage, and Mrs Ida Husted Harper, she published _The History of Woman Suffrage_ (4 vols., New York, 1884-1887). She died at Rochester, New York, on the 13th of March 1906.

See Mrs Ida Husted Harper's _Life and Work of Susan B. Anthony_ (3 vols., Indianapolis, 1898-1908).

ANTHOZOA (i.e. "flower-animals"), the zoological name for a class of marine polyps forming "coral" (q.v.). Although corals have been familiar objects since the days of antiquity, and the variety known as the precious red coral has been for a long time an article of commerce in the Mediterranean, it was only in the 18th century that their true nature and structure came to be understood. By the ancients and the earlier naturalists of the Christian era they were regarded either as petrifactions or as plants, and many supposed that they occupied a position midway between minerals and plants. The discovery of the animal nature of red coral is due to J.A. de Peyssonel, a native of Marseilles, who obtained living specimens from the coral fishers on the coast of Barbary and kept them alive in aquaria. He was thus able to see that the so-called "flowers of coral" were in fact nothing else than minute polyps resembling sea-anemones. His discovery, made in 1727, was rejected by the Academy of Sciences of France, but eventually found acceptance at the hands of the Royal Society of London, and was published by that body in 1751. The structure and classification of polyps, however, were at that time very imperfectly understood, and it was fully a century before the true anatomical characters and systematic position of corals were placed on a secure basis.

The hard calcareous substance to which the name coral is applied is the supporting skeleton of certain members of the _Anthozoa_, one of the classes of the phylum Coelentera. The most familiar Anthozoan is the common sea-anemone, _Actinia equina_, L., and it will serve, although it does not form a skeleton or _corallum_, as a good example of the structure of a typical Anthozoan polyp or zooid. The individual animal or zooid of _Actinia equina_ has the form of a column fixed by one extremity, called the _base_, to a rock or other object, and bearing at the opposite extremity a crown of _tentacles_. The tentacles surround an area known as the _peristome_, in the middle of which there is an elongated mouth-opening surrounded by tumid lips. The mouth does not open directly into the general cavity of the body, as is the case in a hydrozoan polyp, but into a short tube called the _stomodaeum_, which in its turn opens below into the general body-cavity or _coelenteron_. In

## Actinia and its allies, and most generally, though not invariably, in

Anthozoa, the stomodaeum is not circular, but is compressed from side to side so as to be oval or slit-like in transverse section. At each end of the oval there is a groove lined by specially long vibratile cilia. These grooves are known as the _sulcus_ and _sulculus_, and will be more

## particularly described hereafter. The elongation of the mouth and

stomodaeum confer a bilateral symmetry on the body of the zooid, which is extended to other organs of the body. In Actinia, as in all Anthozoan zooids, the coelenteron is not a simple cavity, as in a Hydroid, but is divided by a number of radial folds or curtains of soft tissue into a corresponding number of radial chambers. These radial folds are known as _mesenteries_, and their position and relations may be understood by reference to figs. 1 and 2. Each mesentery is attached by its upper margin to the peristome, by its outer margin to the body-wall, and by its lower margin to the basal disk. A certain number of mesenteries, known as complete mesenteries, are attached by the upper parts of their internal margins to the stomodaeum, but below this level their edges hang in the coelenteron. Other mesenteries, called incomplete, are not attached to the stomodaeum, and their internal margins are free from the peristome to the basal disk. The lower part of the free edge of every mesentery, whether complete or incomplete, is thrown into numerous puckers or folds, and is furnished with a glandular thickening known as a _mesenterial filament_. The reproductive organs or gonads are borne on the mesenteries, the germinal cells being derived from the inner layer or endoderm.

[Illustration: FIG. 1. Diagrammatic longitudinal section of an Anthozoan zooid,

m, Mesentery. s, Stoma. t, Tentacles. lm, Longitudinal muscle. st, Stomodaeum. d, Diagonal Muscle. sc, Sulcus. go, Gonads. r, Rotteken's muscle.]

In common with all Coelenterate animals, the walls of the columnar body and also the tentacles and peristome of Actinia are composed of three layers of tissue. The external layer, or ectoderm, is made up of cells, and contains also muscular and nervous elements. The preponderating elements of the ectodermic layer are elongated columnar cells, each containing a nucleus, and bearing cilia at their free extremities. Packed in among these are _gland cells, sense cells_, and _cnidoblasts_. The last-named are specially numerous on the tentacles and on some other regions of the body, and produce the well-known "thread cells," or _nematocysts_, so characteristic of the Coelentera. The inner layer or endoderm is also a cellular layer, and is chiefly made up of columnar cells, each bearing a cilium at its free extremity and terminating internally in a long muscular fibre. Such cells, made up of epithelial and muscular components, are known as epithelio-muscular or myo-epithelial cells. In Actinians the epithelio-muscular cells of the endoderm are crowded with yellow spherical bodies, which are unicellular plants or Algae, living symbiotically in the tissues of the zooid. The endoderm contains in addition gland cells and nervous elements. The middle layer or mesogloea is not originally a cellular layer, but a gelatinoid structureless substance, secreted by the two cellular layers. In the course of development, however, cells from the ectoderm and endoderm may migrate into it. In _Actinia equina_ the mesogloea consists of fine fibres imbedded in a homogeneous matrix, and between the fibres are minute branched or spindle-shaped cells. For further details of the structure of Actinians, the reader should consult the work of O. and R. Hertwig.

[Illustration: FIG. 2.--1, Portion of epithelium from the tentacle of an

## Actinian, showing three supporting cells and one sense cell (sc); 2, a

cnidoblast with enclosed nematocyst from the same specimen; 3 and 4 two forms of gland cell from the stomodaeum; 5a, 5b, epithelio-muscular cells from the tentacle in different states of contraction; 5c, an epithelio-muscular cell from the endoderm, containing a symbiotic zooxanthella; 6, a ganglion cell from the ectoderm of the peristome. (After O. and R. Hertwig.)]

The Anthozoa are divisible into two sub-classes, sharply marked off from one another by definite anatomical characters. These are the ALCYONARIA and the ZOANTHARIA. To the first-named belong the precious red coral and its allies, the sea-fans or Gorgoniae, to the second belong the white or Madreporarian corals.

[Illustration: FIG. 3.--An expanded Alcyonarian zooid, showing the mouth surrounded by eight pinnate tentacles. st, Stomodaeum in the the centre of the transparent body; m, mesenteries; asm, asulcar mesenteries; B, spicules, enlarged.]

Alcyonaria.--In this sub-class the zooid has very constant anatomical characters, differing in some important respects from the Actinian zooid, which has been taken as a type. There is only one ciliated groove, the sulcus, in the stomodaeum. There are always eight tentacles, which are hollow and fringed on their sides, with hollow projections or pinnae; and always eight mesenteries, all of which are complete, i.e. inserted on the stomodaeum. The mesenteries are provided with well-developed longitudinal retractor muscles, supported on longitudinal folds or plaits of the mesogloea, so that in cross-section they have a branched appearance. These _muscle-banners_, as they are called, have a highly characteristic arrangement; they are all situated on those faces of the mesenteries which look towards the sulcus. (fig. 4). Each mesentery has a filament; but two of them, namely, the pair farthest from the sulcus, are longer than the rest, and have a different form of filament. It has been shown that these asulcar filaments are derived from the ectoderm, the remainder from the endoderm. The only exceptions to this structure are found in the arrested or modified zooids, which occur in many of the colonial Alcyonaria. In these the tentacles are stunted or suppressed and the mesenteries are ill-developed, but the sulcus is unusually large and has long cilia. Such modified zooids are called siphonozooids, their function being to drive currents of fluid through the canal-systems of the colonies to which they belong. With very few exceptions a calcareous skeleton is present in all Alcyonaria; it usually consists of spicules of carbonate of lime, each spicule being formed within an ectodermic cell (fig. 3, B). Most commonly the spicule-forming cells pass out of the ectoderm and are imbedded in the mesogloea, where they may remain separate from one another or may be fused together to form a strong mass. In addition to the spicular skeleton an organic horny skeleton is frequently present, either in the form of a horny external investment (_Cornularia_), or an internal axis (_Gorgonia_), or it may form a matrix in which spicules are imbedded (_Keroeides, Meistodes_).

[Illustration: FIG. 4.--Transverse section of an Alcyonarian zooid mm, Mesenteries; mb, muscle banners; sc, sulcus; st, stomodaeum.]

Nearly all the Alcyonaria are colonial. Four solitary species have been described, viz. _Haimea funebris_ and _H. hyalina, Hartea elegans_, and _Monoxenia Darwinii_; but it is doubtful whether these are not the young forms of colonies. For the present the solitary forms may be placed in a grade, _Protal-cyonacea_, and the colonial forms may be grouped in another grade, _Synalcyonacea_. Every Alcyonarian colony is developed by budding from a single parent zooid. The buds are not direct outgrowths of the body-wall, but are formed on the courses of hollow out growths of the base or body-wall, called _solenia_. These form a more or less complicated canal system, lined by endoderm, and communicating with the cavities of the zooids. The most simple form of budding is found in the genus _Cornularia_, in which the mother zooid gives off from its base one or more simple radiciform outgrowths. Each outgrowth contains a single tube or solenium, and at a longer or shorter distance from the mother zooid a daughter zooid is formed as a bud. This gives off new outgrowths, and these, branching and anastomosing with one another, may form a network, adhering to stones, corals, or other objects, from which zooids arise at intervals. In _Clavularia_ and its allies each outgrowth contains several solenia, and the outgrowths may take the form of flat expansions, composed of a number of solenial tubes felted together to form a lamellar surface of attachment. Such outgrowths are called _stolons_, and a stolon may be simple, i.e. contain only one solenium, as in _Cornularia_, or may be complex and built up of many solenia, as in _Clavularia_. Further complications arise when the lower walls of the mother zooid become thickened and interpenetrated with solenia, from which buds are developed, so that lobose, tufted, or branched colonies are formed. The chief orders of the Synalcyonacea are founded upon the different architectural features of colonies produced by different modes of budding. We recognize six orders--the STOLONIFERA, ALCYONACEA, PSEUDAXONIA, AXIFERA, STELECHOTOKEA, and CORNOTHECALIA.

[Illustration: FIG. 5.

A. Skeleton of a young colony of _Tubipora purpurea_. st, Stolon; p, platform.

B. Diagrammatic longitudinal section of a corallite, showing two platforms, p and cup-shaped tabulae, t. (After S.J. Hickson.)]

In the order STOLONIRERA the zooids spring at intervals from branching or lamellar stolons, and are usually free from one another, except at their bases, but in some cases horizontal solenia arising at various heights from the body-wall may place the more distal portions of the zooids in communication with one another. In the genus _Tubipora_ these horizontal solenia unite to form a series of horizontal platforms (fig. 5). The order comprises the families _Cornulamdae, Syringopordae, Tubipondae_, and _Favositidae_. In the first-named, the zooids are united only by their bases and the skeleton consists of loose spicules. In the _Tubipondae_ the spicules of the proximal part of the body-wall are fused together to form a firm tube, the corallite, into which the distal part of the zooid can be retracted. The corallites are connected at intervals by horizontal platforms containing solenia, and at the level of each platform the cavity of the corallite is divided by a transverse calcareous partition, either flat or cup-shaped, called a _tabula_. Formerly all corals in which tabulae are present were classed together as Tabulata, but Tubipora is an undoubted Alcyonarian with a lamellar stolon, and the structure of the fossil genus Syringopora, which has vertical corallites united by horizontal solenia, clearly shows its affinity to Tubipora. The Favositidae, a fossil family from the Silurian and Devonian, have a massive corallum composed of numerous polygonal corallites closely packed together. The cavities of adjacent corallites communicate by means of numerous perforations, which appear to represent solenia, and numerous transverse tabulae are also present. In _Favosites hemisphaerica_ a number of radial spines, projecting into the cavity of the corallite, give it the appearance of a madreporarian coral.

[Illustration: FIG. 6.--Portion of a colony of _Coralinum rubrum_, showing expanded and contracted zooids. In the lower part of the figure the cortex has been cut away to show the _axis_, ax, and the longitudinal canals, lc, surrounding it.]

In the order ALCYONACEA the colony consists of bunches of elongate cylindrical zooids, whose proximal portions are united by solenia and compacted, by fusion of their own walls and those of the solenia, into a fleshy mass called the coenenchyma. Thus the coenenchyma forms a stem, sometimes branched, from the surface of which the free portions of the zooids project. The skeleton of the Alcyonacea consists of separate calcareous spicules, which are often, especially in the Nephthyidae, so abundant and so closely interlocked as to form a tolerably firm and hard armour. The order comprises the families _Xeniidae, Alcyonidae_ and _Nephthyidae_. _Alcyonium digitatum_, a pink digitate form popularly known as "dead men's fingers," is common in 10-20 fathoms of water off the English coasts.

[Illustration: FIG. 7.--The sea-fan (_Gorgonia cavolinii_).]

In the order PSEUDAXONIA the colonies are upright and branched, consisting of a number of short zooids whose proximal ends are imbedded in a coenenchyma containing numerous ramifying solenia and spicules. The coenenchyma is further differentiated into a medullary portion and a cortex. The latter contains the proximal moieties of the zooids and numerous but separate spicules. The medullary portion is densely crowded with spicules of different shape from those in the cortex, and in some forms the spicules are cemented together to form a hard supporting axis. There are four families of Pseudaxonia--the _Briareidae, Sclerogorgidae, Melitodidae_, and _Corallidae_. In the first-named the medulla is penetrated by solenia and forms an indistinct axis; in the remainder the medulla is devoid of solenia, and in the _Melitodidae_ and _Corallidae_ it forms a dense axis, which in the _Melitodidae_ consists of alternate calcareous and horny joints. The precious red coral of commerce, _Corallium rubrum_ (fig. 6), a member of the family _Corallidae_, is found at depths varying from 15 to 120 fathoms the Mediterranean Sea, chiefly on the African coast. It owes its commercial value to the beauty of its hard red calcareous axis which in life is covered by a cortex in which the proximal moieties of the zooids are imbedded. _Corallium rubrum_ has been the subject of a beautifully-illustrated memoir by de Lacaze-Duthiers, which should be consulted for details of anatomy.

The AXIFERA comprise those corals that have a horny or calcified axis, which in position corresponds to the axis of the Pscudaxonia, but, unlike it, is never formed of fused spicules; the most familiar example is the pink sea-fan, _Gorgonia cavolinii_, which is found in abundance in 10-25 fathoms of water off the English coasts (fig. 7). In this order the axis is formed as an ingrowth of the ectoderm of the base of the mother zooid of the colony, the cavity of the ingrowth being filled by a horny substance secreted by the ectoderm. In _Gorgonia_ the axis remains horny throughout life, but in many forms it is further strengthened by a deposit of calcareous matter In the family _Isidinae_ the axis consists of alternate segments of horny and calcareous substance, the latter being amorphous. The order contains six families--the _Dasygorgidae, Isidae, Primnoidae, Muriceidae, Plexauridae_, and _Gorgoniaae_.

[Illustration: FIG. 8.

A. Colony of _Pennatula phosphorea_ from the metarachidial aspect. p, The peduncle.

B. Section of the rachis bearing a single pinna, a, Axis; b, metarachidial; c, prorachidial; d, pararachidial stem canals.]

In the order STELECHOTOKEA the colony consists of a stem formed by a greatly-elongated mother zooid, and the daughter zooids are borne as lateral buds on the stem. In the section _Asiphonacea_ the colonies are upright and branched, springing from membranous or ramifying stolons. They resemble and are closely allied to certain families of the Cornulariidae, differing from them only in mode of budding and in the dispostion of the daughter zooids round a central, much-elongated mother zooid. The section contains two families, the _Telestidae_ and the _Coelogorgidae_. The second section comprises the _Pennatulacea_ or sea-pens, which are remarkable from the fact that the colony is not fixed by the base to a rock or other object, but is imbedded in sand or mud by the proximal portion of the stem known as the peduncle. In the typical genus, Pennatula (fig. 8), the colony looks like a feather having a stem divisible into an upper moiety or rachis, bearing lateral central leaflets (pinnae), and a lower peduncle, which is sterile and imbedded in sand or mud. The stem represents a greatly enlarged and elongated mother zooid. It is divided longitudinally by a

## partition separating a so-called "ventral" or prorachidial canal from

a so-called "dorsal" or metarachidial canal. A rod-like supporting axis of peculiar texture is developed in the longitudinal partition, and a longitudinal canal is hollowed out on either side of the axis in the substance of the longitudinal partition, so that there are four stem-canals in all. The prorachidial and metarachidial aspects of the rachis are sterile, but the sides or pararachides bear numerous daughter zooids of two kinds--(1) fully-formed autozooids, (2) small stunted siphonozooids. The pinnae are formed by the elongated autozooids, whose proximal portions are fused together to form a leaf-like expansion, from the upper edge of which the distal extremities of the zooids project. The siphonozooids are very numerous and lie between the bases at the pinnae on the pararachides; they extend also on the prorachidial and metarachidial surfaces. The calcareous skeleton of the Pennatulacea consists of scattered spicules, but in one species, _Protocaulon molle_, spicules are absent. Although of great interest the Pennatulacea do not form an enduring skeleton or "coral," and need not be considered in detail in this place.

[Illustration: FIG. 9.

A, Portion of the surface of a colony of _Heliopora coerulea_ magnified, showing two calices and the surrounding coenenchymal tubes.

B, Single zooid with the adjacent soft tissues as seen after removal of the skeleton by decalcification. Z1, the distal, and Z2, the proximal or intracalicular portion of the zooid; ec, ectoderm; ct, coenenchymal tubes; sp, superficial network of solenia.]

The order COENOTHECALIA is represented by a single living species, _Heliopora coerulea_, which differs from all recent Alcyonaria in the fact that its skeleton is not composed of spicules, but is formed as a secretion from a layer of cells called calicoblasts, which originate from the ectoderm. The corallum of Heliopora is of a blue colour, and has the form of broad, upright, lobed, or digitate masses flattened from side to side. The surfaces are pitted all over with perforations of two kinds, viz. larger star-shaped cavities, called _calices_, in which the zooids are lodged, and very numerous smaller round or polygonal apertures, which in life contain as many short unbranched tubes, known as the _coenenchymal tubes_ (fig. 9, A). The walls of the calices and coenenchymal tubes are formed of flat plates of calcite, which are so disposed that the walls of one tube enter into the composition of the walls of adjacent tubes, and the walls of the calices are formed by the walls of adjacent coenenchymal tubes. Thus the architecture of the Helioporid colony differs entirely from such forms as Tubipora or Favosites, in which each corallite has its own distinct and proper wall. The cavities both of the calices and coenenchymal tubes of Heliopora are closed below by horizontal

## partitions or _tabulae_, hence the genus was formerly included in the

group Tabulata, and was supposed to belong to the madreporarian corals, both because of its lamellar skeleton, which resembles that of a Madrepore, and because each calicle has from twelve to fifteen radial partitions or septa projecting into its cavity. The structure of the zooid of Heliopora, however, is that of a typical Alcyonarian, and the septa have only a resemblance to, but no real homology with, the similarly named structures in madreporarian corals. _Heliopora coerulea_ is found between tide-marks on the shore platforms of coral islands. The order was more abundantly represented in Palaeozoic times by the _Heliolitidae_ from the Upper and Lower Silurian and the Devonian, and by the _Thecidae_ from the Wenlock limestone. In _Heliolites porosus_ the colonies had the form of spheroidal masses; the calices were furnished with twelve pseudosepta, and the coenenchymal tubes were more or less regularly hexagonal.

[Illustration with caption: FIG. 10.

A, _Edwardsia claparedii_ (after A. Andres). Cap, capitulum; sc, scapus; ph, physa.

B, Transverse section of the same, showing the arrangement of the mesenteries, s, Sulcus; sl, sulculus.

C, Transverse section of _Halcampa_. d, d, Directive mesenteries; st, stomodaeum.]

Zoantharia.--In this sub-class the arrangement of the mesenteries is subject to a great deal of variation, but all the types hitherto observed may be referred to a common plan, illustrated by the living genus _Edwardsia_ (fig. 10, A, B). This is a small solitary Zoantharian which lives embedded in sand. Its body is divisible into three portions, an upper _capitulum_ bearing the mouth and tentacles, a median _scapus_ covered by a friable cuticle, and a terminal physa which is rounded. Both capitulum and physa can be retracted within the scapus. There are from sixteen to thirty-two simple tentacles, but only eight mesenteries, all of which are complete. The stomodaeum is compressed laterally, and is furnished with two longitudinal grooves, a sulcus and a sulculus. The arrangement of the muscle-banners on the mesenteries is characteristic. On six of the mesenteries the muscle-banners have the same position as in the Alcyonaria, namely, on the sulcar faces; but in the two remaining mesenteries, namely, those which are attached on either side of the sulcus, the muscle-banners are on the opposite or sulcular faces. It is not known whether all the eight mesenteries of _Edwardsia_ are developed simultaneously or not, but in the youngest form which has been studied all the eight mesenteries were present, but only two of them, namely the sulco-laterals, bore mesenterial filaments, and so it is presumed that they are the first pair to be developed. In the common sea-anemone, _Actinia equina_ (which has already been quoted as a type of Anthozoan structure), the mesenteries are numerous and are arranged in cycles. The mesenteries of the first cycle are complete (i.e. are attached to the stomodaeum), are twelve in number, and arranged in couples, distinguishable by the position of the muscle-banners. In the four couples of mesenteries which are attached to the sides of the elongated stomodaeum the muscle-banners of each couple are turned towards one another, but in the sulcar and sulcular couples, known as the directive mesenteries, the muscle-banners are on the outer faces of the mesenteries, and so are turned away from one another (see fig. 10, C). The space enclosed between two mesenteries of the same couple is called an _entocoele_; the space enclosed between two mesenteries of adjacent couples is called an _exocoele_. The second cycle of mesenteries consists of six couples, each formed in an exocoele of the primary cycle, and in each couple the muscle-banners are _vis-a-vis_. The third cycle comprises twelve couples, each formed in an exocoele between the primary and secondary couples and so on, it being a general rule (subject, however, to exceptions) that new mesenterial couples are always formed in the exocoeles, and not in the entocoeles.

[Illustration: FIG. 11.--A, Diagram showing the sequence of mesenterial development in an Actinian. B, Diagrammatic transverse section of _Gonactinia prolifera_.]

While the mesenterial couples belonging to the second and each successive cycle are formed simultaneously, those of the first cycle are formed in successive pairs, each member of a pair being placed on opposite sides of the stomodaeum. Hence the arrangement in six couples is a secondary and not a primary feature. In most Actinians the mesenteries appear in the following order:--At the time when the stomodaeum is formed, a single pair of mesenteries, marked I, I in the diagram (fig. 11, A), makes its appearance, dividing the coelenteric cavity into a smaller sulcar and a large sulcular chamber. The muscle-banners of this pair are placed on the sulcar faces of the mesenteries. Next, a pair of mesenteries, marked II, II in the diagram, is developed in the sulcular chamber, its muscle-banners facing the same way as those of I, I. The third pair is formed in the sulcar chamber, in close connexion with the sulcus, and in this case the muscle-banners are on the _sulcular_ faces. The fourth pair, having its muscle-banners on the sulcar faces, is developed at the opposite extremity of the stomodaeum in close connexion with the sulculus. There are now eight mesenteries present, having exactly the same arrangement as in Edwardsia. A pause in the development follows, during which no new mesenteries are formed, and then the six-rayed symmetry characteristic of a normal Actinian zooid is completed by the formation of the mesenteries V, V in the lateral chambers, and VI, VI in the sulco-lateral chambers, their muscle-banners being so disposed that they form couples respectively with II, II and I, I. In _Actinia equina_ the Edwardsia stage is arrived at somewhat differently. The mesenteries second in order of formation form the sulcular directives, those fourth in order of formation form with the fifth the sulculo-lateral couples of the adult.

[Illustration: FIG. 12.

A, Zoanthid colony, showing the expanded zooids.

B, Diagram showing the arrangement of mesenteries in a young Zoanthid.

C, Diagram showing the arrangement of mesenteries in an adult Zoanthid. 1, 2, 3, 4, Edwardsian mesenteries.]

As far as the anatomy of the zooid is concerned, the majority of the stony or madreporarian corals agree exactly with the soft-bodied

## Actinians, such as _Actinia equina_, both in the number and

arrangement of the adult mesenteries and in the order of development of the first cycle. The few exceptions will be dealt with later, but it may be stated here that even in these the first cycle of six couples of mesenteries is always formed, and in all the cases which have been examined the course of development described above is followed. There are, however, several groups of Zoantharia in which the mesenterial arrangement of the adult differs widely from that just described. But it is possible to refer all these cases with more or less certainty to the Edwardsian type.

The order ZOANTHIDEA comprises a number of soft-bodied Zoantharians generally encrusted with sand. Externally they resemble ordinary sea-anemones, but there is only one ciliated groove, the sulcus, in the stomodaeum, and the mesenteries are arranged on a peculiar pattern. The first twelve mesenteries are disposed in couples, and do not differ from those of Actinia except in size. The mesenterial pairs I, II and III are attached to the stomodaeum, and are called macromesenteries (fig. 12, B), but IV, V and VI are much shorter, and are called micromesenteries. The subsequent development is peculiar to the group. New mesenteries are formed only in the sulco-lateral exocoeles. They are formed in couples, each couple consisting of a macromesentery and a micromesentery, disposed so that the former is nearest to the sulcar directives. The derivation of the Zoanthidea from an Edwardsia form is sufficiently obvious.

The order CERIANTHIDEA comprises a few soft-bodied Zoantharians with rounded aboral extremities pierced by pores. They have two circlets of tentacles, a labial and a marginal, and there is only one ciliated groove in the stomodaeum, which appears to be the sulculus. The mesenteries are numerous, and the longitudinal muscles, though distinguishable, are so feebly developed that there are no muscle-banners. The larval forms of the type genus _Cerianthus_ float freely in the sea, and were once considered to belong to a separate genus, _Arachnactis_. In this larva four pairs of mesenteries having the typical Edwardsian arrangement are developed, but the fifth and sixth pairs, instead of forming couples with the first and second, arise in the sulcar chamber, the fifth pair inside the fourth, and the sixth pair inside the fifth. New mesenteries are continually added in the sulcar chamber, the seventh pair within the sixth, the eighth pair within the seventh, and so on (fig. 13). In the Cerianthidea, as in the Zoanthidea, much as the adult arrangement of mesenteries differs from that of Actinia, the derivation from an Edwardsia stock is obvious.

[Illustration: FIG. 13.

A, _Cerianthus solitarius_ (after A. Andres).

B, Transverse section of the stomodaeum, showing the sulculus, sl, and the arrangement of the mesenteries.

C, Oral aspect of _Arachnactis brachiolata_, the larva of _Cerianthus_, with seven tentacles.

D, Transverse section of an older larva. The numerals indicate the order of development of the mesenteries.]

The order ANTIPATHIDEA is a well-defined group whose affinities are more obscure. The type form, _Antipathes dichotoma_ (fig. 14), forms arborescent colonies consisting of numerous zooids arranged in a single series along one surface of a branched horny axis. Each zooid has six tentacles; the stomodaeum is elongate, but the sulcus and sulculus are very feebly represented. There are ten mesenteries in which the musculature is so little developed as to be almost indistinguishable. The sulcar and sulcular pairs of mesenteries are short, the sulco-lateral and sulculo-lateral pairs are a little longer, but the two transverse are very large and are the only mesenteries which bear gonads. As the development of the Antipathidea is unknown, it is impossible to say what is the sequence of the mesenterial development, but in _Leiopathes glaberrima_, a genus with twelve mesenteries, there are distinct indications of an Edwardsia stage.

[Illustration: FIG. 14.

A, Portion of a colony of _Antipathes dichotoma_.

B, Single zooid and axis of the same magnified. m, Mouth; mf mesenterial filament; ax, axis.

C, Transverse section through the oral cone of _Antipathella minor_, st, Stomodaeum; ov, ovary.]

There are, in addition to these groups, several genera of Actinians whose mesenterial arrangement differs from the normal type. Of these perhaps the most interesting is _Gonactinia prolifera_ (fig. 11, B), with eight macromesenteries arranged on the Edwardsian plan. Two pairs of micromesenteries form couples with the first and second Edwardsian pairs, and in addition there is a couple of micromesenteries in each of the sulculo-lateral exocoeles. Only the first and second pairs of Edwardsian macromesenteries are fertile, i.e. bear gonads.

The remaining forms, the ACTINIIDEA, are divisible into the Malacactiniae, or soft-bodied sea-anemones, which have already been described sufficiently in the course of this article, and the Scleractiniae (= Madreporaria) or true corals.

[Illustration: FIG. 15.--Corallum of _Caryophyllia_; semi-diagrammatic. th, Theca; c, costae; sp, septa; p, palus; col, columella.]

All recent corals, as has already been said, conform so closely to the anatomy of normal Actinians that they cannot be classified apart from them, except that they are distinguished by the possession of a calcareous skeleton. This skeleton is largely composed of a number of radiating plates or _septa_, and it differs both in origin and structure from the calcareous skeleton of all Alcyonaria except Heliopora. It is formed, not from fused spicules, but as a secretion of a special layer of cells derived from the basal ectoderm, and known as _calicoblasts_. The skeleton or corallum of a typical solitary coral--the common Devonshire cup-coral _Caryophyllia smithii_ (fig. 15) is a good example--exhibits the followings parts:--(1) The _basal plate_, between the zooid and the surface of attachment. (2) The _septa_, radial plates of calcite reaching from the periphery nearly or quite to the centre of the coral-cup or calicle. (3) The _theca_ or wall, which in many corals is not an independent structure, but is formed by the conjoined thickened peripheral ends of the septa. (4) The _columella_, a structure which occupies the centre of the calicle, and may arise from the basal plate, when it is called essential, or may be formed by union of trabecular offsets of the septa, when it is called unessential. (5) The _costae_, longitudinal ribs or rows of spines on the outer surface of the theca. True costae always correspond to the septa, and are in fact the peripheral edges of the latter. (6) _Epitheca_, an offset of the basal plate which surrounds the base of the theca in a ring-like manner, and in some corals may take the place of a true theca. (7) _Pali_, spinous or blade-like upgrowths from the bottom of the calicle, which project between the inner edges of certain septa and the columella. In addition to these parts the following structures may exist in corals:--_Dissepiments_ are oblique calcareous partitions, stretching from septum to septum, and closing the interseptal chambers below. The whole system of dissepiments in any given calicle is often called _endotheca_. _Synapticulae_ are calcareous bars uniting adjacent septa. _Tabulae_ are stout horizontal partitions traversing the centre of the calicle and dividing it into as many superimposed chambers. The septa in recent corals always bear a definite relation to the mesenteries, being found either in every entocoele or in every entocoele and exocoele. Hence in corals in which there is only a single cycle of mesenteries the septa are correspondingly few in number; where several cycles of mesenteries are present the septa are correspondingly numerous. In some cases--e.g. in some species of _Madrepora_--only two septa are fully developed, the remainder being very feebly represented.

[Illustration: FIG. 16.--Tangential section of a larva of _Astroides calicularis_ which has fixed itself on a piece of cork. ec, Ectoderm; en, endoderm; mg, mesogloea; m, m, mesenteries; s, septum; b, basal plate formed of ellipsoids of carbonate of lime secreted by the basal ectoderm; ep, epitheca. (After von Koch.)]

Though the corallum appears to live within the zooid, it is morphologically external to it, as is best shown by its developmental history. The larvae of corals are free swimming ciliated forms known as planulae, and they do not acquire a corallum until they fix themselves. A ring-shaped plate of calcite, secreted by the ectoderm, is then formed, lying between the embryo and the surface of attachment. As the mesenteries are formed, the endoderm of the basal disk lying above the basal plate is raised up in the form of radiating folds. There may be six of these folds, one in each entocoele of the primary cycle of mesenteries, or there may be twelve, one in each exocoele and entocoele. The ectoderm beneath each fold becomes detached from the surface of the basal plate, and both it and the mesogloea are folded conformably with the endoderm. The cells forming the limbs of the ectodermic folds secrete nodules of calcite, and these, fusing together, give rise to six (or twelve) vertical radial plates or septa. As growth proceeds new septa are formed simultaneously with the new couples of secondary mesenteries. In some corals, in which all the septa are entocoelic, each new system is embraced by a mesenteric couple; in others, in which the septa are both entocoelic and exocoelic, three septa are formed in every chamber between two primary mesenterial couples, one in the entocoele of the newly formed mesenterial couple of the secondary cycle, and one in each exocoele between a primary and a secondary couple. These latter are in turn embraced by the couples of the tertiary cycle of mesenteries, and new septa are formed in the exocoeles on either side of them, and so forth.

[Illustration: FIG. 17.--Transverse section through a zooid of _Cladocora_. The corallum shaded with dots, the mesogloea represented by a thick line. Thirty-two septa are present, six in the entocoeles of the primary cycle of mesenteries, I; six in the entocoeles of the secondary cycle of mesenteries, II; four in the entocoeles of the tertiary cycle of mesenteries, III, only four pairs of the latter being developed; and sixteen in the entocoeles between the mesenterial pairs. D, D, Directive mesenteries; st, stomodaeum. (After Duerden.)]

It is evident from an inspection of figs. 16 and 17 that every septum is covered by a fold of endoderm, mesogloea, and ectoderm, and is in fact pushed into the cavity of the zooid from without. The zooid then is, as it were, moulded upon the corallum. When fully extended, the upper part of the zooid projects for some distance out of the calicle, and its wall is reflected for some distance over the lip of the latter, forming a fold of soft tissue extending to a greater or less distance over the theca, and containing in most cases a cavity continuous over the lip of the calicle with the coelenteron. This fold of tissue is known as the _edge-zone_ In some corals the septa are solid imperforate plates of calcite, and their peripheral ends are either firmly welded together, or are united by interstitial pieces so as to form imperforate theca. In others the peripheral ends of the septa are united only by bars or trabeculae, so that the theca is perforate, and in many such perforate corals the septa themselves are pierced by numerous perforations. In the former, which have been called aporose corals, the only communication between the cavity of the edge-zone and the general cavity of the zooid is by way of the lip of the calicle; in the latter, or perforate corals, the theca is permeated by numerous branching and anastomosing canals lined by endoderm, which place the cavity of the edge-zone in communication with the general cavity of the zooid.

[Illustration: FIG. 18.

A, Schematic longitudinal section through a zooid and bud of _Stylophora digitata_. In A, B, and C the thick black lines represent the soft tissues; the corallum is dotted. s, Stomodaeum; c, c, coenosarc; col, columella, T tabulae.

B, Similar section through a single zooid and bud of _Astroides calicularis_.

C, Similar section through three corallites of _Lophohelia prolifera_. ez, Edge-zone.

D, Diagram illustrating the process of budding by unequal division.

E, Section through a dividing calicle of _Mussa_, showing the union of two septa in the plane of division and the origin of new septa at right angles to them.

(C original; the rest after von Koch.)]

A large number of corals, both aporose and perforate, are colonial. The colonies are produced by either budding or division. In the former case the young daughter zooid, with its corallum, arises wholly outside the cavity of the parent zooid, and the component parts of the young corallum, septa, theca, columella, &c., are formed anew in every individual produced. In division a vertical constriction divides a zooid into two equal or unequal parts, and the several parts of the two corals thus produced are severally derived from the corresponding parts of the dividing corallum. In colonial corals a bud is always formed from the edge-zone, and this bud develops into a new zooid with its corallum. The cavity of the bud in an aporose coral (fig. 18, A, C) does not communicate directly with that of the parent form, but through the medium of the edge-zone. As growth proceeds, and parent and bud become separated farther from one another, the edge-zone forms a sheet of soft tissue, bridging over the space between the two, and resting upon projecting spines of the corallum. This sheet of tissue is called the _coenosarc_. Its lower surface is clothed with a layer of calicoblasts which continue to secrete carbonate of lime, giving rise to a secondary deposit which more or less fills up the spaces between the individual coralla, and is distinguished as _coenenchyme_. This coenenchyme may be scanty, or may be so abundant that the individual corallites produced by budding seem to be immersed in it. Budding takes place in an analogous manner in perforate corals (fig. 18, B), but the presence of the canal system in the perforate theca leads to a modification of the process. Buds arise from the edge-zone which already communicate with the cavity of the zooid by the canals. As the buds develop the canal system becomes much extended, and calcareous tissue is deposited between the network of canals, the confluent edge-zones of mother zooid and bud forming a coenosarc. As the process continues a number of calicles are formed, imbedded in a spongy tissue in which the canals ramify, and it is impossible to say where the theca of one corallite ends and that of another begins. In the formation of colonies by division a constriction at right angles to the long axis of the mouth involves first the mouth, then the peristome, and finally the calyx itself, so that the previously single corallite becomes divided into two (fig. 18, E). After division the corallites continue to grow upwards, and their zooids may remain united by a bridge of soft tissue or coenosarc. But in some cases, as they grow farther apart, this continuity is broken, each corallite has its own edge-zone, and internal continuity is also broken by the formation of dissepiments within each calicle, all organic connexion between the two zooids being eventually lost. Massive meandrine corals are produced by continual repetition of a process of incomplete division, involving the mouth and to some extent the peristome: the calyx, however, does not divide, but elongates to form a characteristic meandrine channel containing several zooid mouths.

Corals have been divided into _Aporosa_ and _Perforata_, according as the theca and septa are compact and solid, or are perforated by pores containing canals lined by endoderm. The division is in many respects convenient for descriptive purposes, but recent researches show that it does not accurately represent the relationships of the different families. Various attempts have been made to classify corals according to the arrangement of the septa, the characters of the theca, the microscopic structure of the corallum, and the anatomy of the soft parts. The last-named method has proved little more than that there is a remarkable similarity between the zooids of all recent corals, the differences which have been brought to light being for the most part secondary and valueless for classificatory purposes. On the other hand, the study of the anatomy and development of the zooids has thrown much light upon the manner in which the corallum is formed, and it is now possible to infer the structure of the soft parts from a microscopical examination of the septa, theca, &c., with the result that unexpected relationships have been shown to exist between corals previously supposed to stand far apart. This has been particularly the case with the group of Palaeozoic corals formerly classed together as _Rugosa_. In many of these so-called rugose forms the septa have a characteristic arrangement, differing from that of recent corals chiefly in the fact that they show a tetrameral instead of a hexameral symmetry. Thus in the family _Stauridae_ there are four chief septa whose inner ends unite in the middle of the calicle to form a false columella, and in the _Zaphrentidae_ there are many instances of an arrangement, such as that depicted in fig. 19, which represents the septal arrangement of _Streptelasma corniculum_ from the lower Silurian. In this coral the calicle is divided into quadrants by four principal septa, the _main septum, counter septum_, and two _alar septa_. The remaining septa are so disposed that in the quadrants abutting on the chief septum they converge towards that septum, whilst in the other quadrants they converge towards the alar septa. The secondary septa show a regular gradation in size, and, assuming that the smallest were the most recently formed, it will be noticed that in the chief quadrants the youngest septa lie nearest to the main septum; in the other quadrants the youngest septa lie nearest to the alar septa. This arrangement, however, is by no means characteristic even of the Zaphrentidae, and in the family _Cyathophyllidae_ most of the genera exhibit a radial symmetry in which no trace of the bilateral arrangement described above is recognizable, and indeed in the genus _Cyathophyllum_ itself a radial arrangement is the rule. The connexion between the Cyathophyllidae and modern Astraeidae is shown by _Moseleya latistellata_, a living reef-building coral from Torres Strait. The general structure of this coral leaves no doubt that it is closely allied to the Astraeidae, but in the young calicles a tetrameral symmetry is indicated by the presence of four large septa placed at right angles to one another. Again, in the family _Amphiastraeidae_ there is commonly a single septum much larger than the rest, and it has been shown that in the young calicles, e.g. of _Thecidiosmilia_, two septa, corresponding to the main- and counter-septa of Streptelasma, are first formed, then two alar septa, and afterwards the remaining septa, the latter taking on a generally radial arrangement, though the original bilaterality is marked by the preponderance of the main septum. As the microscopic character of the corallum of these extinct forms agrees with that of recent corals, it may be assumed that the anatomy of the soft parts also was similar, and the tetrameral arrangement, when present, may obviously be referred to a stage when only the first two pairs of Edwardsian mesenteries were present and septa were formed in the intervals between them.

[Illustration: FIG. 19.--Diagram of the arrangement of the septa in a Zaphrentid coral. m, Main septum; c, counter septum; t, t, alar septa.]

Space forbids a discussion of the proposals to classify corals after the minute structure of their coralla, but it will suffice to say that it has been shown that the septa of all corals are built up of a number of curved bars called trabeculae, each of which is composed of a number of nodes. In many secondary corals (_Cyclolites, Thamnastraea_) the trabeculae are so far separate that the individual bars are easily recognizable, and each looks something like a bamboo owing to the thickening of the two ends of each node. The trabeculae are united together by these thickened internodes, and the result is a fenestrated septum, which in older septa may become solid and aporose by continual deposit of calcite in the fenestrae. Each node of a trabecula may be simple, i.e. have only one centre of calcification, or may be compound. The septa of modern perforate corals are shown to have a structure nearly identical with that of the secondary forms, but the trabeculae and their nodes are only apparent on microscopical examination. The aporose corals, too, have a practically identical structure, their compactness being due to the union of the trabeculae throughout their entire lengths instead of at intervals, as in the Perforata. Further, the trabeculae may be evenly spaced throughout the septum, or may be grouped together, and this feature is probably of value in estimating the affinities of corals. (For an account of coral formations see CORAL-REEFS.)

In the present state of our knowledge the Zoantharia in which a primary cycle of six couples of mesenteries is (or may be inferred to be) completed by the addition of two pairs to the eight Edwardsian mesenteries, and succeeding cycles are formed in the exocoeles of the pre-existing mesenterial cycles, may be classed in an order ACTINIIDEA, and this may be divided into the suborders _Malacactiniae_, comprising the soft-bodied Actinians, such as _Actinia, Sagartia, Bunodes_, &c., and the _Scleractiniae_, comprising the corals. The Scleractiniae may best be divided into groups of families which appear to be most closely related to one another, but it should not be forgotten that there is great reason to believe that many if not most of the extinct corals must have differed from modern Actiniidea in mesenterial characters, and may have only possessed Edwardsian mesenteries, or even have possessed only four mesenteries, in this respect showing close affinities to the Stauromedusae. Moreover, there are some modern corals in which the secondary cycle of mesenteries departs from the Actinian plan. For example, J.E. Duerden has shown that in _Porites_ the ordinary zooids possess only six couples of mesenteries arranged on the Actinian plan. But some zooids grow to a larger size and develop a number of additional mesenteries, which arise either in the sulcar or the sulcular entocoele, much in the same manner as in Cerianthus. Bearing this in mind, the following arrangement may be taken to represent the most recent knowledge of coral structure:--

GROUP A.

Family I. ZAPHRENTIDAE.--Solitary Palaeozoic corals with an epithecal wall. Septa numerous, arranged pinnately with regard to four principal septa. Tabulae present. One or more pits or fossulae present in the calicle. Typical genera--_Zaphrentis_, Raf. _Amplexus_, M. Edw. and H. _Streptelasma_, Hall. _Omphyma_, Raf.

Family 2. TURBINOLIDAE.--Solitary, rarely colonial corals, with radially arranged septa and without tabulae. Typical genera--_Flabellum_, Lesson. _Turbinolia_, M. Edw. and H. _Caryophyllia_, Lamarck. _Sphenotrochus_, Moseley, &c.

Family 3. AMPHIASTRAEIDAE.--Mainly colonial, rarely solitary corals, with radial septa, but bilateral arrangement indicated by persistence of a main septum. Typical genera--_Amphiastraea_, Etallon. _Thecidiosmilia_.

Family 4. STYLINIDAE.--Colonial corals allied to the Amphiastraeidae, but with radially symmetrical septa arranged in cycles. Typical genera--_Stylina_, Lamarck (Jurassic). _Convexastraea_, D'Orb. (Jurassic). _Isastraea_, M. Edw. and H.(Jurassic). Ogilvie refers the modern genus _Galaxea_ to this family.

GROUP B.

Family 5. OCULINIDAE.--Branching or massive aporose corals, the calices projecting above the level of a compact coenenchyme formed from the coenosarc which covers the exterior of the corallum. Typical genera--_Lophohelia_, M. Edw. and H. _Oculina_, M. Edw. and H.

Family 6. POCILLOPORIDAE.--Colonial branching aporose corals, with small calices sunk in the coenenchyme. Tabulae present, and two larger septa, an axial and abaxial, are always present, with traces of ten smaller septa. Typical genera--_Pocillopora_, Lamarck. _Seriatopora_, Lamarck.

Family 7. MADREPORIDAE.--Colonial branching or palmate perforate corals, with abundant trabecular coenenchyme. Theca porous; septa compact and reduced in number. Typical genera--_Madrepora_, Linn. _Turbinaria_, Oken. _Montipora_, Quoy and G.

Family 8. PORITIDAE.--Incrusting or massive colonial perforate corals; calices usually in contact by their edges, sometimes disjunct and immersed in coenenchyme. Theca and septa perforate. Typical genera--_Porites_, M. Edw. and H. _Goniopora_, Quoy and G. _Rhodaraea_, M. Edw. and H.

GROUP C.

Family 9. CYATHOPHYLLIDAE.--Solitary and colonial aporose corals. Tabulae and vesicular endotheca present. Septa numerous, generally radial, seldom pinnate. Typical genera--_Cyathophyllum_, Goldfuss (Devonian and Carboniferous). _Moseleya_, Quelch (recent).

Family 10. ASTRAEIDAE.--Aporpse, mainly colonial corals, massive, branching, or maeandroid. Septa radial; dissepiments present; an epitheca surrounds the base of massive or maeandroid forms, but only surrounds individual corallites in simple or branching forms. Typical genera--_Goniastraea_, M. Edw. and H. _Heliastraea_, M. Edw. and H. _Maeandrina_, Lam. _Coeloria_, M. Edw. and H. _Favia_, Oken.

Family 11. FUNGIDAE.--Solitary and colonial corals, with numerous radial septa united by synapticulae. Typical genera--_Lophoseris_, M. Edw. and H. _Thamnastraea_, Le Sauvage. _Leptophyllia_, Reuss (Jurassic and Cretaceous). _Fungia_, Dana. _Siderastraea_, Blainv.

GROUP D.

Family 12. EUPSAMMIDAE.--Solitary or colonial perforate corals, branching, massive, or encrusting. Septa radial; the primary septa usually compact, the remainder perforate. Theca perforate. Synapticula present in some genera. Typical genera--_Stephanophyllia_, Michelin. _Eupsammia_, M. Edw. and H. _Astroides_, Blainv. _Rhodopsammia_, M. Edw. and H. _Dendrophyllia_, M. Edw. and H.

GROUP E.

Family 13. CYSTIPHYLLIDAE.--Solitary corals with rudimentary septa, and the calicle filled with vesicular endotheca. Genera--_Cystiphyllum_, Lonsdale (Silurian and Devonian). _Goniophyllum_, M. Edw. and H. (In this Silurian genus the calyx is provided with a movable operculum, consisting of four paired triangular pieces, the bases of each being attached to the sides of the calyx, and their apices meeting in the middle when the operculum is closed). _Calcecla_, Lam. (In this Devonian genus there is a single semicircular operculum furnished with a stout median septum and numerous feebly developed secondary septa. The calyx is triangular in section, pointed below, and the operculum is attached to it by hinge-like teeth.)

AUTHORITIES.--The following list contains only the names of the more important and more general works on the structure and classification of corals and on coral reefs. For a fuller bibliography the works marked with an asterisk should be consulted: * A. Andres, _Fauna und Flora des Golfes von Neapel_, ix. (1884); H.M. Bernard, "Catalogue of Madreporarian Corals" in Brit. Museum, ii. (1896), iii. (1897); * G.C. Bourne, "Anthozoa," in E. Ray Lankester's _Treatise on Zoology_, vol. ii. (London, 1900); G. Brook, "_Challenger_ Reports," _Zoology_, xxxii. (1899) (_Antipatharia_); "Cat. Madrep. Corals," Brit. Museum, i. (1893); D.C. Danielssen, "Report Norwegian North Atlantic Exploring Expedition," _Zoology_, xix. (1890); J.E. Duerden, "Some Results on the Morphology and Development of Recent and Fossil Corals," _Rep. Brit. Association_, 1903, pp. 684-685; "The Morphology of the Madreporaria," _Biol. Bullet_, vii. pp. 79-104; P.M. Duncan, _Journ. Linnean Soc._ xviii. (1885); P.H. Gosse, _Actinologia britannica_ (London, 1860); O. and R. Hertwig, _Die Actinien_ (Jena, 1879); R. Hertwig, "_Challenger_ Reports," _Zoology_, vi. (1882) and xxvi. (1888); * C.B. Klunzinger, _Die Korallthiere des Rothen Meeres_ (Berlin, 1877); * G. von Koch, _Fauna und Flora des Golfes van Neapel_, xv. (1887); _Mitth. Zool. Stat. Neapel_, ii. (1882) and xii. (1897); _Palaeontographica_, xxix. (1883); (also many papers in the _Morphol. Jahrbuch_ from 1878 to 1898); F. Koby, "Polypiers jurassiques de la Suisse," _Mem. Soc. Palaeont. Suisse_, vii.-xvi. (1880-1889); A. von Kolliker, "Die Pennatuliden," _Abh. d. Senck. Naturf. Gesell_. vii.; * "_Challenger_ Reports," _Zoology_, i. _Pennatulidae_ (1880); Koren and Danielssen, _Norske Nordhaus Exped., Alcyonida_ (1887); H. de Lacaze-Duthiers, _Hist. nat. du corail_ (Paris, 1864); H. Milne-Edwards and J. Haime, _Hist. nat. des coralliaires_ (Paris, 1857); H.N. Moseley, "_Challenger_ Reports," _Zoology_, ii. (1881); H.A. Nicholson, _Palaeozoic Tabulate Corals_ (Edinburgh, 1879); M.M. Ogilvie, _Phil. Transactions_, clxxxvii. (1896); E. Pratz, _Palaeontographica_, xxix. (1882); J.J. Quelch, "_Challenger_ Reports," _Zoology_, xvi. (1886); * P.S. Wright and Th. Studer, "_Challenger_ Reports," _Zoology_, xxxi. (1889). (G. C. B.)

ANTHRACENE (from the Greek [Greek: anthrax], coal), C14H10, a hydrocarbon obtained from the fraction of the coal-tar distillate boiling between 270 deg. and 400 deg. C. This high boiling fraction is allowed to stand for some days, when it partially solidifies. It is then separated in a centrifugal machine, the low melting-point impurities are removed by means of hot water, and the residue is finally hot-pressed. The crude anthracene cake is purified by treatment with the higher pyridine bases, the operation being carried out in large steam-jacketed boilers. The whole mass dissolves on heating, and the anthracene crystallizes out on cooling. The crystallized anthracene is then removed by a centrifugal separator and the process of solution in the pyridine bases is repeated. Finally the anthracene is purified by sublimation.

Many synthetical processes for the preparation of anthracene and its derivatives are known. It is formed by the condensation of acetylene tetrabromide with benzene in the presence of aluminium chloride:--

Br.CH.Br /CH\ C6H6 + | + C6H6 = 4HBr + C6H4< | >C6H4, Br.CH.Br \CH/

and similarly from methylene dibromide and benzene, and also when benzyl chloride is heated with aluminium chloride to 200 deg. C. By condensing ortho-brombenzyl bromide with sodium, C.L. Jackson and J.F. White (_Ber_., 1879, 12, p. 1965) obtained dihydro-anthracene

/CH2Br Br\ /CH2\ C6H4< + 4Na + >C6H4 = 4NaBr + C6H4< >C6H4. \Br BrCH2/ \CH2/

Anthracene has also been obtained by heating ortho-tolylphenyl ketone with zinc dust

/CH8 /CH \ C6H4< = H2O + C6H4< | >C6H4. \COC6H5 \CH /

Anthracene crystallizes in colourless monoclinic tables which show a fine blue fluorescence. It melts at 213 deg. C. and boils at 351 deg. C. It is insoluble in water, sparingly soluble in alcohol and ether, but readily soluble in hot benzene. It unites with picric acid to form a picrate, C14H10.C6H2(NO2)3.OH, which crystallizes in needles, melting at 138 deg. C. On exposure to sunlight a solution of anthracene in benzene or xylene deposits para-anthracene (C14H10)2, which melts at 244 deg. C. and passes back into the ordinary form. Chlorine and bromine form both addition and substitution products with anthracene; the addition product, anthracene dichloride, C14H10Cl2, being formed when chlorine is passed into a cold solution of anthracene in carbon bisulphide. On treatment with potash, it forms the substitution product, monochlor-anthracene, C14H9Cl. Nitro-anthracenes are not as yet known. The mono-oxyanthracenes (anthrols), C14H9OH or

/CH\ C6H4< | >C6H3OH \CH/

([alpha]) and ([beta]) resemble the phenols, whilst

/C(OH)\ C6H4< | >C6H4 \CH /

([gamma]) (anthranol) is a reduction product of anthraquinone. [beta]-anthrol and anthranol give the corresponding amino compounds (anthramines) when heated with ammonia.

Numerous sulphonic acids of anthracene are known, a monosulphonic acid being obtained with dilute sulphuric acid, whilst concentrated sulphuric acid produces mixtures of the anthracene disulphonic acids. By the

## action of sodium amalgam on an alcoholic solution of anthracene, an

anthracene dihydride, C14H12, is obtained, whilst by the use of stronger reducing agents, such as hydriodic acid and amorphous phosphorus, hydrides of composition C14H16 and C14H24 are produced.

Methyl and phenyl anthracenes are known; phenyl anthranol (phthalidin) being somewhat closely related to the phenolphthaleins (q.v.). Oxidizing agents convert anthracene into anthraquinone (q.v.); the production of this substance by oxidizing anthracene in glacial acetic acid solution, with chromic acid, is the usual method employed for the estimation of anthracene.

ANTHRACITE (Gr. [Greek: anthrax], coal), a term applied to those varieties of coal which do not give off tarry or other hydrocarbon vapours when heated below their point of ignition; or, in other words, which burn with a smokeless and nearly non-luminous flame. Other terms having the same meaning are, "stone coal" (not to be confounded with the German _Steinkohle_) or "blind coal" in Scotland, and "Kilkenny coal" in Ireland. The imperfect anthracite of north Devon, which however is only used as a pigment, is known as _culm_, the same term being used in geological classification to distinguish the strata in which it is found, and similar strata in the Rhenish hill countries which are known as the Culm Measures. In America, culm is used as an equivalent for waste or slack in anthracite mining.

Physically, anthracite differs from ordinary bituminous coal by its greater hardness, higher density, 1.3-1.4, and lustre, the latter being often semi-metallic with a somewhat brownish reflection. It is also free from included soft or fibrous notches and does not soil the fingers when rubbed. Structurally it shows some alteration by the development of secondary divisional planes and fissures so that the original stratification lines are not always easily seen. The thermal conductivity is also higher, a lump of anthracite feeling perceptibly colder when held in the warm hand than a similar lump of bituminous coal at the same temperature. The chemical composition of some typical anthracites is given in the article COAL.

Anthracite may be considered to be a transition stage between ordinary bituminous coal and graphite, produced by the more or less complete elimination of the volatile constituents of the former; and it is found most abundantly in areas that have been subjected to considerable earth-movements, such as the flanks of great mountain ranges. The largest and most important anthracite region, that of the north-eastern portion of the Pennsylvania coal-field, is a good example of this; the highly contorted strata of the Appalachian region produce anthracite exclusively, while in the western portion of the same basin on the Ohio and its tributaries, where the strata are undisturbed, free-burning and coking coals, rich in volatile matter, prevail. In the same way the anthracite region of South Wales is confined to the contorted portion west of Swansea and Llanelly, the central and eastern portions producing steam, coking and house coals.

Anthracites of newer, tertiary or cretaceous age, are found in the Crow's Nest part of the Rocky Mountains in Canada, and at various points in the Andes in Peru.

The principal use of anthracite is as a smokeless fuel. In the eastern United States, it is largely employed as domestic fuel, usually in close stoves or furnaces, as well as for steam purposes, since, unlike that from South Wales, it does not decrepitate when heated, or at least not to the same extent. For proper use, however, it is necessary that the fuel should be supplied in pieces as nearly uniform in size as possible, a condition that has led to the development of the breaker which is so characteristic a feature in American anthracite mining (see COAL). The large coal as raised from the mine is passed through breakers with toothed rolls to reduce the lumps to smaller pieces, which are separated into different sizes by a system of graduated sieves, placed in descending order. Each size can be perfectly well burnt alone on an appropriate grate, if kept free from larger or smaller admixtures. The common American classification is as follows:--

Lump, steamboat, egg and stove coals, the latter in two or three sizes, all three being above 1-1/2 in. size on round-hole screens.

Chestnut below 1-1/2 inch above 7/8 inch. Pea " 7/8 " " 9/16 " Buckwheat " 9/16 " " 3/8 " Rice " 3/8 " " 3/16 " Barley " 3/16 " " 3/32 "

From the pea size downwards the principal use is for steam purposes. In South Wales a less elaborate classification is adopted; but great care is exercised in hand-picking and cleaning the coal from included

## particles of pyrites in the higher qualities known as best malting

coals, which are used for kiln-drying malt and hops.

Formerly, anthracite was largely used, both in America and South Wales, as blast-furnace fuel for iron smelting, but for this purpose it has been largely superseded by coke in the former country and entirely in the latter. An important application has, however, been developed in the extended use of internal combustion motors driven by the so-called "mixed," "poor," "semi-water" or "Dowson gas" produced by the gasification of anthracite with air and a small proportion of steam. This is probably the most economical method of obtaining power known; with an engine as small as 15 horse-power the expenditure of fuel is at the rate of only 1 lb per horse-power hour, and with larger engines it is proportionately less. Large quantities of anthracite for power purposes are now exported from South Wales to France, Switzerland and parts of Germany. (H. B.)

ANTHRACOTHERIUM ("coal-animal," so called from the fact of the remains first described having been obtained from the Tertiary lignite-beds of Europe), a genus of extinct artiodactyle ungulate mammals, characterized by having 44 teeth, with five semi-crescentic cusps on the crowns of the upper molars. In many respects, especially the form of the lower jaw, _Anthracotherium_, which is of Oligocene and Miocene age in Europe, and typifies the family _Anthracotheriidae_, is allied to the hippopotamus, of which it is probably an ancestral form. The European _A. magnum_ was as large as the last-mentioned animal, but there were several smaller species and the genus also occurs in Egypt, India and North America. (See ARTIODACTYLA.)

ANTHRAQUINONE, C14H8O2, an important derivative of anthracene, first prepared in 1834 by A. Laurent. It is prepared commercially from anthracene by stirring a sludge of anthracene and water in horizontal cylinders with a mixture of sodium bichromate and caustic soda. This suspension is then run through a conical mill in order to remove all grit, the cones of the mill fitting so tightly that water cannot pass through unless the mill is running; the speed of the mill when working is about 3000 revolutions per minute. After this treatment, the mixture is run into lead-lined vats and treated with sulphuric acid, steam is blown through the mixture in order to bring it to the boil, and the anthracene is rapidly oxidized to anthraquinone. When the oxidation is complete, the anthraquinone is separated in a filter press, washed and heated to 120 deg. C. with commercial oil of vitriol, using about 2-1/2 parts of vitriol to 1 of anthraquinone. It is then removed to lead-lined tanks and again washed with water and dried; the product obtained contains about 95% of anthraquinone. It may be purified by sublimation. Various synthetic processes have been used for the preparation of anthraquinone. A. Behr and W.A. v. Dorp (_Ber._, 1874, 7, p. 578) obtained orthobenzoyl benzoic acid by heating phthalic anhydride with benzene in the presence of aluminium chloride. This compound on heating with phosphoric anhydride loses water and yields anthraquinone,

/CO\ C6H6 /CO.C6H6 /CO\ C6H4< >O -> C6H4< -> C6H4< >C6H4. \CO/ \COOH \CO/

It may be prepared in a similar manner by heating phthalyl chloride with benzene in the presence of aluminium chloride. Dioxy- and tetraoxy-anthraquinones are obtained when meta-oxy- and dimeta-dioxy-benzoic acids are heated with concentrated sulphuric acid.

Anthraquinone crystallizes in yellow needles or prisms, which melt at 277 deg. C. It is soluble in hot benzene, sublimes easily, and is very stable towards oxidizing agents. On the other hand, it is readily attacked by reducing agents. With zinc dust in presence of caustic soda it yields the secondary alcohol oxan-thranol, C6H4 : CO.CHOH : C6H4, with tin and hydrochloric acid, the phenolic compound anthranol, C6H4 : CO.C(OH) : C6H4; and with hydriodic acid at 150 deg. C. or on distillation with zinc dust, the hydrocarbon anthracene, C14H10. When fused with caustic potash, it gives benzoic acid. It behaves more as a ketone than as a quinone, since with hydroxylamine it yields an oxime, and on reduction with zinc dust and caustic soda it yields a secondary alcohol, whilst it cannot be reduced by means of sulphurous acid. Various sulphonic acids of anthraquinone are known, as well as oxy-derivatives, for the preparation and properties of which see ALIZARIN.

ANTHRAX (the Greek for "coal," or "carbuncle," so called by the ancients because they regarded it as burning like coal; cf. the French equivalent _charbon_; also known as _fievre charbonneuse, Milzbrand_, splenic fever, and malignant pustule), an acute, specific, infectious, virulent disease, caused by the _Bacillus anthracis_, in animals, chiefly cattle, sheep and horses, and frequently occurring in workers in the wool or hair, as well as in those handling the hides or carcases, of beasts which have been affected.

_Animals._--As affecting wild as well as domesticated animals and man, anthrax has been widely diffused in one or more of its forms, over the surface of the globe. It at times decimates the reindeer herds in Lapland and the Polar regions, and is only too well known in the tropics and in temperate latitudes. It has been observed and described in Russia, Siberia, Central Asia, China, Cochin-China, Egypt, West Indies, Peru, Paraguay, Brazil, Mexico, and other parts of North and South America, in Australia, and on different parts of the African continent, while for other European countries the writings which have been published with regard to its nature, its peculiar characteristics, and the injury it inflicts are innumerable. Countries in which are extensive marshes, or the subsoil of which is tenacious or impermeable, are usually those most frequently and seriously visited. Thus there have been regions notorious for its prevalence, such as the marshes of Sologne, Dombes and Bresse in France; certain parts of Germany, Hungary and Poland; in Spain the half-submerged valleys and the maritime coasts of Catalonia, as well as the Romagna and other marshy districts of Italy; while it is epizootic, and even panzootic, in the swampy regions of Esthonia, Livonia, Courland, and especially of Siberia, where it is known as the _Sibirskaja jaswa_ (Siberian boil-plague). The records of anthrax go back to a very ancient date. It is supposed to be the murrain of Exodus. Classical writers allude to anthrax as if it were the only cattle disease worthy of mention (see Virgil, _Georg._ iii.). It figures largely in the history of the early and middle ages as a devastating pestilence attacking animals, and through them mankind; the oldest Anglo-Saxon manuscripts contain many fantastic recipes, leechdoms, charms and incantations for the prevention or cure of the "blacan blezene" (black blain) and the relief of the "elfshot" creatures. In the 18th and 19th centuries it sometimes spread like an epizootic over the whole of Europe, from Siberia to France. It was in this malady that disease-producing germs (_bacteria_) were first discovered, in 1840, by Pollender of Wipperfurth, and, independently, by veterinary surgeon Brauell of Dorpat, and their real character afterwards verified by C.J. Davaine (1812-1882) of Alfort in 1863; and it was in their experiments with this disease that Toussaint, Pasteur and J.B. Chauveau first showed how to make the morbific poison its own antidote. (See VIVISECTION.)

The symptoms vary with the species of animal, the mode of infection, and the seat of the primary lesion, internal or external. In all its forms anthrax is an inoculable disease, transmission being surely and promptly effected by this means, and it may be conveyed to nearly all animals by inoculation of a wound of the skin or through the digestive organs. Cattle, sheep and horses nearly always owe their infection to spores or bacilli ingested with their food or water, and pigs usually contract the disease by eating the flesh of animals dead of anthrax.

Internal anthrax, of cattle and sheep, exhibits no premonitory symptoms that can be relied on. Generally the first indication of an outbreak is the sudden death of one or more of the herd or flock. Animals which do not die at once may be noticed to stagger and tremble; the breathing becomes hurried and the pulse very rapid, while the heart beats violently; the internal temperature of the body is high, 104 deg. to 106 deg. F.; blood oozes from the nose, mouth and anus, the visible mucous membranes are dusky or almost black. The animal becomes weak and listless, the temperature falls and death supervenes in a few hours, being immediately preceded by delirium, convulsions or coma. While death is usually rapid or sudden when the malady is general, constituting what is designated splenic apoplexy, internal anthrax in cattle is not invariably fatal. In some cases the animal rallies from a first attack and gradually recovers.

In the external or localized form, marked by the formation of carbuncles before general infection takes place, death may not occur for several days. The carbuncles may appear in any part of the body, being preceded or accompanied by fever. They are developed in the subcutaneous connective tissue where this is loose and plentiful, in the interstices of the muscles, lymphatic glands, in the mucous membranes of the mouth and tongue (glossanthrax of cattle), pharynx and larynx (_anthrax angina_ of horses and pigs), and the rectum. They begin as small circumscribed swellings which are warm, slightly painful and oedematous. In from two to eight hours they attain a considerable size, are cold, painless and gangrenous, and when they are incised a quantity of a blood-stained gelatinous exudate escapes. When the swellings have attained certain proportions symptoms of general infection appear, and, running their course with great rapidity, cause death in a few hours. Anthrax of the horse usually begins as an affection of the throat or bowel. In the former there is rapid obstructive oedema of the mucous membrane of the pharynx and larynx with swelling of the throat and neck, fever, salivation, difficulty in swallowing, noisy breathing, frothy discharge from the nose and threatening suffocation. General invasion soon ensues, and the horse may die in from four to sixteen hours. The intestinal form is marked by high temperature, great prostration, small thready pulse, tumultuous action of the heart, laboured breathing and symptoms of abdominal pain with straining and diarrhoea. When moved the horse staggers and trembles. Profuse sweating, a falling temperature and cyanotic mucous membranes indicate the approach of a fatal termination.

In splenic fever or splenic apoplexy, the most marked alterations observed after death are--the effects of rapid decomposition, evidenced by the foul odour, disengagement of gas beneath the skin and in the tissues and cavities of the body, yellow or yellowish-red gelatinous exudation into and between the muscles, effusion of citron or rust-coloured fluid in various cavities, extravasations of blood and local congestions throughout the body, the blood in the vessels generally being very dark and tar-like. The most notable feature, however, in the majority of cases is the enormous enlargement of the spleen, which is engorged with blood to such an extent that it often ruptures, while its tissue is changed into a violet or black fluid mass.

The bacillus of anthrax, under certain conditions, retains its vitality for a long time, and rapidly grows when it finds a suitable field in which to develop, its mode of multiplication being by scission and the formation of spores, and depending, to a great extent at least, on the presence of oxygen. The morbid action of the bacillus is indeed said to be due to its affinity for oxygen; by depriving the red corpuscles of the blood of that most essential gas, it renders the vital fluid unfit to sustain life. Albert Hoffa and others assert that the fatal lesions are produced by the poisonous action of the toxins formed by the bacilli and not by the blocking up of the minute blood-vessels, or the abstraction of oxygen from the blood by the bacilli.

It was by the cultivation of this micro-organism, or attenuation of the virus, that Pasteur was enabled to produce a prophylactic remedy for anthrax. His discovery was first made with regard to the cholera of fowls, a most destructive disorder which annually carries off great numbers of poultry. Pasteur produced his inoculation material by the cultivation of the bacilli at a temperature of 42 deg. C. in oxygen. Two vaccines are required. The first or weak vaccine is obtained by incubating a bouillon culture for twenty-four days at 42 deg. C., and the second or less attenuated vaccine by incubating a bouillon culture, at the same temperature, for twelve days. Pasteur's method of protective inoculation comprises two inoculations with an interval of twelve days between them. Immunity, established in about fifteen days after the injection of the second vaccine, lasts from nine months to a year.

Toussaint had, previous to Pasteur, attenuated the virus of anthrax by the action of heat; and Chauveau subsequently corroborated by numerous experiments the value of Toussaint's method, demonstrating that, according to the degree of heat to which the virus is subjected, so is its inocuousness when transferred to a healthy creature. In outbreaks of anthrax on farms where many animals are exposed to infection immediate temporary protection can be conferred by the injection of anthrax serum.

_Human Beings._--For many years cases of sudden death had been observed to occur from time to time among healthy men engaged in woollen manufactories, particularly in the work of sorting or combing wool. In some instances death appeared to be due to the direct inoculation of some poisonous material into the body, for a form of malignant pustule was observed upon the skin; but, on the other hand, in not a few cases without any external manifestation, symptoms of blood-poisoning, often proving rapidly fatal, suggested the probability of other channels for the introduction of the disease. In 1880 the occurrence of several such cases among woolsorters at Bradford, reported by Dr J.H. Bell of that town, led to an official inquiry in England by the Local Government Board, and an elaborate investigation into the pathology of what was then called "woolsorters' disease" was at the same time conducted at the Brown Institution, London, by Professor W.S. Greenfield. Among the results of this inquiry it was ascertained: (1) that the disease appeared to be identical with that occurring among sheep and cattle; (2) that in the blood and tissues of the body was found in abundance, as in the disease in animals, the _Bacillus anthracis_, and (3) that the skins, hair, wool, &c., of animals dying of anthrax retain this infecting organism, which, under certain conditions, finds ready access to the bodies of the workers.

Two well-marked forms of this disease in man are recognized, "external anthrax" and "internal anthrax." In external anthrax the infecting agent is accidentally inoculated into some portion of skin, the seat of a slight abrasion, often the hand, arm or face. A minute swelling soon appears at the part, and develops into a vesicle containing serum or bloody matter, and varying in size, but seldom larger than a shilling. This vesicle speedily bursts and leaves an ulcerated or sloughing surface, round about which are numerous smaller vesicles which undergo similar changes, and the whole affected part becomes hard and tender, while the surrounding surface participates in the inflammatory action, and the neighbouring lymphatic glands are also inflamed. This condition, termed "malignant pustule," is frequently accompanied with severe constitutional disturbance, in the form of fever, delirium, perspirations, together with great prostration and a tendency to death from septicaemia, although on the other hand recovery is not uncommon. It was repeatedly found that the matter taken from the vesicle during the progress of the disease, as well as the blood in the body after death, contained the _Bacillus anthracis_, and when inoculated into small animals produced rapid death, with all the symptoms and post-mortem appearances characteristic of che disease as known to affect them.

In internal anthrax there is no visible local manifestation of the disease, and the spores or bacilli appear to gain access to the system from the air charged with them, as in rooms where the contaminated wool or hair is unpacked, or again during the process of sorting. The symptoms usually observed are those of rapid physical prostration, with a small pulse, somewhat lowered temperature (rarely fever), and quickened breathing. Examination of the chest reveals inflammation of the lungs and pleura. In some cases death takes place by collapse in less than one day, while in others the fatal issue is postponed for three or four days, and is preceded by symptoms of blood-poisoning, including rigors, perspirations, extreme exhaustion, &c. In some cases of internal anthrax the symptoms are more intestinal than pulmonary, and consist in severe exhausting diarrhoea, with vomiting and rapid sinking. Recovery from the internal variety, although not unknown, is more rare than from the external, and its most striking phenomena are its sudden onset in the midst of apparent health, the rapid development of physical prostration, and its tendency to a fatal termination despite treatment. The post-mortem appearances in internal anthrax are such as are usually observed in septicaemia, but in addition evidence of extensive inflammation of the lungs, pleura and bronchial glands has in most cases been met with. The blood and other fluids and the diseased tissues are found loaded with the _Bacillus anthracis_.

Treatment in this disease appears to be of but little avail, except as regards the external form, where the malignant pustule may be excised or dealt with early by strong caustics to destroy the affected textures. For the relief of the general constitutional symptoms, quinine, stimulants and strong nourishment appear to be the only available means. An anti-anthrax serum has also been tried. As preventive measures in woollen manufactories, the disinfection of suspicious material, or the wetting of it before handling, is recommended as lessening the risk to the workers. (J. Mac.)

ANTHROPOID APES, or MANLIKE APES, the name given to the family of the Simiidae, because, of all the ape-world, they most closely resemble man. This family includes four kinds, the gibbons of S.E. Asia, the orangs of Borneo and Sumatra, the gorillas of W. Equatorial Africa, and the chimpanzees of W. and Central Equatorial Africa. Each of these apes resembles man most in some one physical characteristic: the gibbons in the formation of the teeth, the orangs in the brain-structure, the gorillas in size, and the chimpanzees in the sigmoid flexure of the spine. In general structure they all closely resemble human beings, as in the absence of tails; in their semi-erect position (resting on finger-tips or knuckles); in the shape of vertebral column, sternum and pelvis; in the adaptation of the arms for turning the palm uppermost at will; in the possession of a long vermiform appendix to the short caecum of the intestine; in the size of the cerebral hemispheres and the complexity of their convolutions. They differ in certain respects, as in the proportion of the limbs, in the bony development of the eyebrow ridges, and in the opposable great toe, which fits the foot to be a climbing and grasping organ.

Man differs from them in the absence of a hairy coat; in the development of a large lobule to the external ear; in his fully erect attitude; in his flattened foot with the non-opposable great toe; in the straight limb-bones; in the wider pelvis; in the marked sigmoid flexure of his spine; in the perfection of the muscular movements of the arm; in the delicacy of hand; in the smallness of the canine teeth and other dental peculiarities; in the development of a chin; and in the small size of his jaws compared to the relatively great size of the cranium. Together with man and the baboons, the anthropoid apes form the group known to science as Catarhini, those, that is, possessing a narrow nasal septum, and are thus easily distinguishable from the flat-nosed monkeys or Platyrhini. The anthropoid apes are arboreal and confined to the Old World. They are of special interest from the important place assigned to them in the arguments of Darwin and the Evolutionists. It is generally admitted now that no fundamental anatomical difference can be proved to exist between these higher apes and man, but it is equally agreed that none probably of the Simiidae is in the direct line of human ancestry. There is a great gap to be bridged between the highest anthropoid and the lowest man, and much importance has been attached to the discovery of an extinct primate, Pithecanthropus (q.v.), which has been regarded as the "missing link."

See Huxley's _Man's Place in Nature_ (1863); Robt. Hartmann's _Anthropoid Apes_ (1883; London, 1885); A.H. Keane's _Ethnology_ (1896); Darwin's _Descent of Man_ (1871; pop. ed., 1901); Haeckel's _Anthropogeny_ (Leipzig, 1874, 1903; Paris, 1877; Eng. ed., 1883); W.H. Flower and Rich. Lydekker, _Mammals Living and Extinct_ (London, 1891).

ANTHROPOLOGY (Gr. [Greek: anthropos] man, and [Greek: logos], theory or science), the science which, in its strictest sense, has as its object the study of man as a unit in the animal kingdom. It is distinguished from ethnology, which is devoted to the study of man as a _racial_ unit, and from ethnography, which deals with the _distribution_ of the races formed by the aggregation of such units. To anthropology, however, in its more general sense as the natural history of man, ethnology and ethnography may both be considered to belong, being related as parts to a whole.

Various other sciences, in conformity with the above definition, must be regarded as subsidiary to anthropology, which yet hold their own independent places in the field of knowledge. Thus anatomy and physiology display the structure and functions of the human body, while psychology investigates the operations of the human mind. Philology deals with the general principles of language, as well as with the relations between the languages of particular races and nations. Ethics or moral science treats of man's duty or rules of conduct toward his fellow-men. Sociology and the science of culture are concerned with the origin and development of arts and sciences, opinions, beliefs, customs, laws and institutions generally among mankind within historic time; while beyond the historical limit the study is continued by inferences from relics of early ages and remote districts, to interpret which is the task of prehistoric archaeology and geology.

I. _Man's Place in Nature._--In 1843 Dr J.C. Prichard, who perhaps of all others merits the title of founder of modern anthropology, wrote in his _Natural History of Man_:--

"The organized world presents no contrasts and resemblances more remarkable than those which we discover on comparing mankind with the inferior tribes. That creatures should exist so nearly approaching to each other in all the particulars of their physical structure, and yet differing so immeasurably in their endowments and capabilities, would be a fact hard to believe, if it were not manifest to our observation. The differences are everywhere striking: the resemblances are less obvious in the fulness of their extent, and they are never contemplated without wonder by those who, in the study of anatomy and physiology, are first made aware how near is man in his physical constitution to the brutes. In all the principles of his internal structure, in the composition and functions of his parts, man is but an animal. The lord of the earth, who contemplates the eternal order of the universe, and aspires to communion with its invisible Maker, is a being composed of the same materials, and framed on the same principles, as the creatures which he has tamed to be the servile instruments of his will, or slays for his daily food. The points of resemblance are innumerable; they extend to the most recondite arrangements of that mechanism which maintains instrumentally the physical life of the body, which brings forward its early development and admits, after a given period, its decay, and by means of which is prepared a succession of similar beings destined to perpetuate the race."

The acknowledgment of man's structural similarity with the anthropomorphous species nearest approaching him, viz.: the higher or anthropoid apes, had long before Prichard's day been made by Linnaeus, who in his _Systema Naturae_ (1735) grouped them together as the highest order of Mammalia, to which he gave the name of Primates. The _Amoenitates Academicae_ (vol. vi., Leiden, 1764), published under the auspices of Linnaeus, contains a remarkable picture which illustrates a discourse by his disciple Hoppius, and is here reproduced (see Plate, fig. 1). In this picture, which shows the crudeness of the zoological notions current in the 18th century as to both men and apes, there are set in a row four figures: (a) a recognizable orang-utan, sitting and holding a staff; (b) a chimpanzee, absurdly humanized as to head, hands, and feet; (c) a hairy woman, with a tail a foot long; (d) another woman, more completely coated with hair. The great Swedish naturalist was possibly justified in treating the two latter creatures as quasi-human, for they seem to be grotesque exaggerations of such tailed and hairy human beings as really, though rarely, occur, and are apt to be exhibited as monstrosities (see Bastian and Hartmann, _Zeitschrift fur Ethnologie_, Index, "Geschwanzte Menschen"; Gould and Pile, _Anomalies and Curiosities of Medicine_, 1897). To Linnaeus, however, they represented normal anthropomorpha or man-like creatures, vouched for by visitors to remote parts of the world. This opinion of the Swedish naturalist seems to have been little noticed in Great Britain till it was taken up by the learned but credulous Scottish judge, Lord Monboddo (see his _Origin and Progress of Language_, 1774, &c.; _Antient Metaphysics_, 1778). He had not heard of the tailed men till he met with them in the work of Linnaeus, with whom he entered into correspondence, with the result that he enlarged his range of mankind with races of sub-human type. One was founded on the description by the Swedish sailor Niklas Koping of the ferocious men with long tails inhabiting the Nicobar Islands. Another comprised the orang-utans of Sumatra, who were said to take men captive and set them to work as slaves. One of these apes, it was related, served as a sailor on board a Jamaica ship, and used to wait on the captain. These are stories which seem to carry their own explanation. When the Nicobar Islands were taken over by the British government two centuries later, the native warriors were still wearing their peculiar loin-cloth hanging behind in a most tail-like manner (E.H. Man, _Journal Anthropological Institute_, vol. xv. p. 442). As for the story of the orang-utan cabin boy, this may even be verbally true, it being borne in mind that in the Malay languages the term _orang-utan_, "man of the forest," was originally used for inland forest natives and other rude men, rather than for the _miyas_ apes to which it has come to be generally applied by Europeans. The speculations as to primitive man connected with these stories diverted the British public, headed by Dr Johnson, who said that Monboddo was "as jealous of his tail as a squirrel." Linnaeus's primarily zoological classification of man did not, however, suit the philosophical opinion of the time, which responded more readily to the systems represented by Buffon, and later by Cuvier, in which the human mind and soul formed an impassable wall of

## partition between him and other mammalia, so that the definition of

man's position in the animal world was treated as not belonging to zoology, but to metaphysics and theology. It has to be borne in mind that Linnaeus, plainly as he recognized the likeness of the higher simian and the human types, does not seem to have entertained the thought of accounting for this similarity by common descent. It satisfied his mind to consider it as belonging to the system of nature, as indeed remained the case with a greater anatomist of the following century, Richard Owen. The present drawing, which under the authority of Linnaeus shows an anthropomorphic series from which the normal type of man, the _Homo sapiens_, is conspicuously absent, brings zoological similarity into view without suggesting kinship to account for it. There are few ideas more ingrained in ancient and low civilization than that of relationship by descent between the lower animals and man. Savage and barbaric religions recognize it, and the mythology of the world has hardly a more universal theme. But in educated Europe such ideas had long been superseded by the influence of theology and philosophy, with which they seemed too incompatible. In the 19th century, however, Lamarck's theory of the development of new species by habit and circumstance led through Wallace and Darwin to the doctrines of the hereditary transmission of acquired characters, the survival of the fittest, and natural selection. Thenceforward it was impossible to exclude a theory of descent of man from ancestral beings whom zoological similarity connects also, though by lines of descent not at all clearly defined, with ancestors of the anthropomorphic apes. In one form or another such a theory of human descent has in our time become part of an accepted framework of zoology, if not as a demonstrable truth, at any rate as a working hypothesis which has no effective rival.

The new development from Linnaeus's zoological scheme which has thus ensued appears in Huxley's diagram of simian and human skeletons (fig. 2, (a) gibbon; (b) orang; (c) chimpanzee; (d) gorilla; (e) man). Evidently suggested by the Linnean picture, this is brought up to the modern level of zoology, and continued on to man, forming an introduction to his zoological history hardly to be surpassed. Some of the main points it illustrates may be briefly stated here, the reader being referred for further information to Huxley's _Essays_. In tracing the osteological characters of apes and man through this series, the general system of the skeletons, and the close correspondence in number and arrangement of vertebrae and ribs, as well as in the teeth, go far towards justifying the opinion of hereditary connexion. At the same time, the comparison brings into view differences in human structure adapted to man's pre-eminent mode of life, though hardly to be accounted its chief causes. It may be seen how the arrangement of limbs suited for going on all-fours belongs rather to the apes than to man, and walking on the soles of the feet rather to man than the apes. The two modes of progression overlap in human life, but the child's tendency when learning is to rest on the soles of the feet and the palms of the hands, unlike the apes, which support themselves on the sides of the feet and the bent knuckles of the hands. With regard to climbing, the long stretch of arm and the grasp with both hands and feet contribute to the arboreal life of the apes, contrasting with what seem the mere remains of the climbing habit to be found even among forest savages. On the whole, man's locomotive limbs are not so much specialized to particular purposes, as generalized into adaptation to many ends. As to the mechanical conditions of the human body, the upright posture has always been recognized as the chief. To it contributes the balance of the skull on the cervical vertebrae, while the human form of the pelvis provides the necessary support to the intestines in the standing attitude. The marked curvature of the vertebral column, by breaking the shock to the neck and head in running and leaping, likewise favours the erect position. The lowest coccygeal vertebrae of man remain as a rudimentary tail. While it is evident that high importance must be attached to the adaptation of the human body to the life of diversified intelligence and occupation he has to lead, this must not be treated as though it were the principal element of the superiority of man, whose comparison with all lower genera of mammals must be mainly directed to the intellectual organ, the brain. Comparison of the brains of vertebrate animals (see BRAIN) brings into view the immense difference between the small, smooth brain of a fish or bird and the large and convoluted organ in man. In man, both size and complexity contribute to the increased area of the cortex or outer layer of the brain, which has been fully ascertained to be the seat of the mysterious processes by which sensation furnishes the groundwork of thought. Schafer (_Textbook of Physiology_, vol. ii. p. 697) thus defines it: "The cerebral cortex is the seat of the intellectual functions, of intelligent sensation or consciousness, of ideation, of volition, and of memory."

The relations between man and ape are most readily stated in comparison with the gorilla, as on the whole the most anthropomorphous ape. In the general proportions of the body and limbs there is a marked difference between the gorilla and man. The gorilla's brain-case is smaller, its trunk larger, its lower limbs shorter, its upper limbs longer in proportion than those of man. The differences between a gorilla's skull and a man's are truly immense. In the gorilla, the face, formed largely by the massive jaw-bones, predominates over the brain-case or cranium; in the man these proportions are reversed. In man the occipital foramen, through which passes the spinal cord, is placed just behind the centre of the base of the skull, which is thus evenly balanced in the erect posture, whereas the gorilla, which goes habitually on all fours, and whose skull is inclined forward, in accordance with this posture has the foramen farther back. In man the surface of the skull is comparatively smooth, and the brow-ridges project but little, while in the gorilla these ridges overhang the cavernous orbits like penthouse roofs. The absolute capacity of the cranium of the gorilla is far less than that of man; the smallest adult human cranium hardly measuring less than 63 cub. in., while the largest gorilla cranium measured had a content of only 34-1/2 cub. in. The largest proportional size of the facial bones, and the great projection of the jaws, confer on the gorilla's skull its small facial angle and brutal character, while its teeth differ from man's in relative size and number of fangs. Comparing the lengths of the extremities, it is seen that the gorilla's arm is of enormous length, in fact about one-sixth longer than the spine, whereas a man's arm is one-fifth shorter than the spine; both hand and foot are proportionally much longer in the gorilla than in man; the leg does not so much differ. The vertebral column of the gorilla differs from that of man in its curvature and other characters, as also does the conformation of its narrow pelvis. The hand of the gorilla corresponds essentially as to bones and muscles with that of man, but is clumsier and heavier; its thumb is "opposable" like a human thumb, that is, it can easily meet with its extremity the extremities of the other fingers, thus possessing a character which does much to make the human hand so admirable an instrument; but the gorilla's thumb is proportionately shorter than man's. The foot of the higher apes, though often spoken of as a hand, is anatomically not such, but a prehensile foot. It has been argued by Sir Richard Owen and others that the position of the great toe converts the foot of the higher apes into a hand, an extremely important distinction from man; but against this Professor T.H. Huxley maintained that it has the characteristic structure of a foot with a very movable great toe. The external unlikeness of the apes to man depends much on their hairiness, but this and some other characteristics have no great zoological value. No doubt the difference between man and the apes depends, of all things, on the relative size and organization of the brain. While similar as to their general arrangement to the human brain, those of the higher apes, such as the chimpanzee, are much less complex in their convolutions, as well as much less in both absolute and relative weight--the weight of a gorilla's brain hardly exceeding 20 oz., and a man's brain hardly weighing less thin 32 oz., although the gorilla is considerably the larger animal of the two.

These anatomical distinctions are undoubtedly of great moment, and it is an interesting question whether they suffice to place man in a zoological order by himself. It is plain that some eminent zoologists, regarding man as absolutely differing as to mind and spirit from any other animal, have had their discrimination of mere bodily differences unconsciously sharpened, and have been led to give differences, such as in the brain or even the foot of the apes and man, somewhat more importance than if they had merely distinguished two species of apes. Many naturalists hold the opinion that the anatomical differences which separate the gorilla or chimpanzee from man are in some respects less than those which separate these man-like apes from apes lower in the scale. Yet all authorities class both the higher and lower apes in the same order. This is Huxley's argument, some prominent points of which are the following: As regards the proportion of limbs, the hylobates or gibbon is as much longer in the arms than the gorilla as the gorilla is than the man, while on the other hand, it is as much longer in the legs than the man as the man is than the gorilla. As to the vertebral column and pelvis, the lower apes differ from the gorilla as much as, or more than, it differs from man. As to the capacity of the cranium, men differ from one another so extremely that the largest known human skull holds nearly twice the measure of the smallest, a larger proportion than that in which man surpasses the gorilla; while, with proper allowance for difference of size of the various species, it appears that some of the lower apes fall nearly as much below the higher apes. The projection of the muzzle, which gives the character of brutality to the gorilla as distinguished from the man, is yet further exaggerated in the lemurs, as is also the backward position of the occipital foramen. In characters of such importance as the structure of the hand and foot, the lower apes diverge extremely from the gorilla; thus the thumb ceases to be opposable in the American monkeys, and in the marmosets is directed forwards, and armed with a curved claw like the other digits, the great toe in these latter being insignificant in proportion. The same argument can be extended to other points of anatomical structure, and, what is of more consequence, it appears true of the brain. A series of the apes, arranged from lower to higher orders, shows gradations from a brain little higher that that of a rat, to a brain like a small and imperfect imitation of a man's; and the greatest structural break in the series lies not between man and the man-like apes, but between the apes and monkeys on one side, and the lemurs on the other. On these grounds Huxley, restoring in principle the Linnean classification, desired to include man in the order of _Primates_. This order he divided into seven families: first, the _Anthropini_, consisting of man only; second, the _Catarhini_ or Old World apes; third, the _Platyrhini_, all New World apes, except the marmosets; fourth, the _Arclopithecini_, or marmosets; fifth, the _Lemurini_, or lemurs; sixth and seventh, the _Cheiromyini_ and _Galeopithecini_.

It is in assigning to man his place in nature on psychological grounds that the greater difficulty arises. Huxley acknowledged an immeasurable and practically infinite divergence, ending in the present enormous psychological gulf between ape and man. It is difficult to account for this intellectual chasm as due to some minor structural difference. The opinion is deeply rooted in modern as in ancient thought, that only a distinctively human element of the highest import can account for the severance between man and the highest animal below him. Differences in the mechanical organs, such as the perfection of the human hand as an instrument, or the adaptability of the human voice to the expression of human thought, are indeed of great value. But they have not of themselves such value, that to endow an ape with the hand and vocal organs of a man would be likely to raise it through any large part of the interval that now separates it from humanity. Much more is to be said for the view that man's larger and more highly organized brain accounts for those mental powers in which he so absolutely surpasses the brutes.

The distinction does not seem to lie principally in the range and delicacy of direct sensation, as may be judged from such well-known facts as man's inferiority to the eagle in sight, or to the dog in scent. At the same time, it seems that the human sensory organs may have in various respects acuteness beyond those of other creatures. But, beyond a doubt, man possesses, and in some way possesses by virtue of his superior brain, a power of co-ordinating the impressions of his senses, which enables him to understand the world he lives in, and by understanding to use, resist, and even in a measure rule it. No human art shows the nature of this human attribute more clearly than does language. Man shares with the mammalia and birds the direct expression of the feelings by emotional tones and interjectional cries; the parrot's power of articulate utterance almost equals his own; and, by association of ideas in some measure, some of the lower animals have even learnt to recognize words he utters. But, to use words in themselves unmeaning, as symbols by which to conduct and convey the complex intellectual processes in which mental conceptions are suggested, compared, combined, and even analysed, and new ones created--this is a faculty which is scarcely to be traced in any lower animal. The view that this, with other mental processes, is a function of the brain, is remarkably corroborated by modern investigation of the disease of aphasia, where the power of thinking remains, but the power is lost of recalling the word corresponding to the thought, and this mental defect is found to accompany a diseased state of a particular locality of the brain (see APHASIA). This may stand among the most perfect of the many evidences that, in Professor Bain's words, "the brain is the principal, though not the sole organ of mind." As the brains of the vertebrate animals form an ascending scale, more and more approaching man's in their arrangement, the fact here finds its explanation, that lower animals perform mental processes corresponding in their nature to our own, though of generally less power and complexity. The full evidence of this correspondence will be found in such works as Brehm's _Thierleben_; and some of the salient points are set forth by Charles Darwin, in the chapter on "Mental Powers," in his _Descent of Man_. Such are the similar effects of terror on man and the lower animals, causing the muscled to tremble, the heart to palpitate, the sphincters to be relaxed, and the hair to stand on end. The phenomena of memory, as to both persons and places, is strong in animals, as is manifest by their recognition of their masters, and their returning at once to habits of which, though disused for many years, their brain has not lost the stored-up impressions. Such facts as that dogs "hunt in dreams," make it likely that their minds are not only sensible to actual events, present and past, but can, like our minds, combine revived sensations into ideal scenes in which they are actors,--that is to say, they have the faculty of imagination. As for the reasoning powers in animals, the accounts of monkeys learning by experience to break eggs carefully, and pick off bits of shell, so as not to lose the contents, or of the way in which rats or martens after a while can no longer be caught by the same kind of trap, with innumerable similar facts, show in the plainest way that the reason of animals goes so far as to form by new experience a new hypothesis of cause and effect which will henceforth guide their actions. The employment of mechanical instruments, of which instances of monkeys using sticks and stones furnish the only rudimentary traces among the lower animals, is one of the often-quoted distinctive powers of man. With this comes the whole vast and ever-widening range of inventive and adaptive art, where the uniform hereditary instinct of the cell-forming bee and the nest-building bird is supplanted by multiform processes and constructions, often at first rude and clumsy in comparison to those of the lower instinct, but carried on by the faculty of improvement and new invention into ever higher stages. "From the moment," writes A.R. Wallace (_Natural Selection_), "when the first skin was used as a covering, when the first rude spear was formed to assist in the chase, when fire was first used to cook his food, when the first seed was sown or shoot planted, a grand revolution was effected in nature, a revolution which in all the previous ages of the earth's history had had no parallel; for a being had arisen who was no longer necessarily subject to change with the changing universe,--a being who was in some degree superior to nature, inasmuch as he knew how to control and regulate her action, and could keep himself in harmony with her, not by a change in body, but by an advance of mind."

As to the lower instincts tending directly to self-preservation, it is acknowledged on all hands that man has them in a less developed state than other animals; in fact, the natural defencelessness of the human being, and the long-continued care and teaching of the young by the elders, are among the commonest themes of moral discourse. Parental tenderness and care for the young are strongly marked among the lower animals, though so inferior in scope and duration to the human qualities; and the same may be said of the mutual forbearance and defence which bind together in a rudimentary social bond the families and herds of animals. Philosophy seeking knowledge for its own sake; morality, manifested in the sense of truth, right, and virtue; and religion, the belief in and communion with superhuman powers ruling and pervading the universe, are human characters, of which it is instructive to trace, if possible, the earliest symptoms in the lower animals, but which can there show at most only faint and rudimentary signs of their wondrous development in mankind. That the tracing of physical and even intellectual continuity between the lower animals and our own race, does not necessarily lead the anthropologist to lower the rank of man in the scale of nature, may be shown by citing A.R. Wallace. Man, he considers, is to be placed "apart, as not only the head and culminating point of the grand series of organic nature, but as in some degree a new and distinct order of being."

To regard the intellectual functions of the brain and nervous system as alone to be considered in the psychological comparison of man with the lower animals, is a view satisfactory to those thinkers who hold materialistic views. According to this school, man is a machine, no doubt the most complex and wonderfully adapted of all known machines, but still neither more nor less than an instrument whose energy is provided by force from without, and which, when set in action, performs the various operations for which its structure fits it, namely, to live, move, feel, and think. This view, however, always has been strongly opposed by those who accept on theological grounds a spiritualistic doctrine, or what is, perhaps, more usual, a theory which combines spiritualism and materialism in the doctrine of a composite nature in man, animal as to the body and in some measure as to the mind, spiritual as to the soul. It may be useful, as an illustration of one opinion on this subject, to continue here the citation of Dr Prichard's comparison between man and the lower animals:--

"If it be inquired in what the still more remarkable difference consists, it is by no means easy to reply. By some it will be said that man, while similar in the organization of his body to the lower tribes, is distinguished from them by the possession of an immaterial soul, a principle capable of conscious feeling, of intellect and thought. To many persons it will appear paradoxical to ascribe the endowment of a soul to the inferior tribes in the creation, yet it is difficult to discover a valid argument that limits the possession of an immaterial principle to man. The phenomena of feeling, of desire and aversion, of love and hatred, of fear and revenge, and the perception of external relations manifested in the life of brutes, imply, not only through the analogy which they display to the human faculties, but likewise from all that we can learn or conjecture of their particular nature, the superadded existence of a principle distinct from the mere mechanism of material bodies. That such a principle must exist in all beings capable of sensation, or of anything analogous to human passions and feelings, will hardly be denied by those who perceive the force of arguments which metaphysically demonstrate the immaterial nature of the mind. There may be no rational grounds for the ancient dogma that the souls of the lower animals were imperishable, like the soul of man: this is, however, a problem which we are not called upon to discuss; and we may venture to conjecture that there may be immaterial essences of divers kinds, and endowed with various attributes and capabilities. But the real nature of these unseen principles eludes our research: they are only known to us by their external manifestations. These manifestations are the various powers and capabilities, or rather the habitudes of action, which characterize the different orders of being, diversified according to their several destinations."

Dr Prichard here puts forward distinctly the time-honoured doctrine which refers the mental faculties to the operation of the soul. The view maintained by a distinguished comparative anatomist, Professor St George Mivart, in his _Genesis of Species_, ch. xii., may fairly follow. "Man, according to the old scholastic definition, is 'a rational animal' (_animal rationale_), and his animality is distinct in nature from his rationality, though inseparably joined, during life, in one common personality. Man's animal body must have had a different source from that of the spiritual soul which informs it, owing to the distinctness of the two orders to which those two existences severally belong." The two extracts just given, however, significant in themselves, fail to render an account of the view of the human constitution which would probably, among the theological and scholastic leaders of public opinion, count the largest weight of adherence. According to this view, not only life but thought are functions of the animal system, in which man excels all other animals as to height of organization: but beyond this, man embodies an immaterial and immortal spiritual principle which no lower creature possesses, and which makes the resemblance of the apes to him but a mocking simulance. To pronounce any absolute decision on these conflicting doctrines is foreign to our present purpose, which is to show that all of them count among their adherents men of high rank in science.

II. _Origin of Man._--Opinion as to the genesis of man is divided between the theories of creation and evolution. In both schools, the ancient doctrine of the contemporaneous appearance on earth of all species of animals having been abandoned under the positive evidence of geology, it is admitted that the animal kingdom, past and present, includes a vast series of successive forms, whose appearances and disappearances have taken place at intervals during an immense lapse of ages. The line of inquiry has thus been directed to ascertaining what formative relation subsists among these species and genera, the last link of the argument reaching to the relation between man and the lower creatures preceding him in time. On both the theories here concerned it would be admitted, in the words of Agassiz (_Principles of Zoology_, pp. 205-206), that "there is a manifest progress in the succession of beings on the surface of the earth. This progress consists in an increasing similarity of the living fauna, and, among the vertebrates especially, in their increasing resemblance to man." Agassiz continues, however, in terms characteristic of the creationist school: "But this connexion is not the consequence of a direct lineage between the faunas of different ages. There is nothing like parental descent connecting them. The fishes of the Palaeozoic age are in no respect the ancestors of the reptiles of the Secondary age, nor does man descend from the mammals which preceded him in the Tertiary age. The link by which they are connected is of a higher and immaterial nature; and their connexion is to be sought in the view of the Creator himself, whose aim in forming the earth, in allowing it to undergo the successive changes which geology has pointed out, and in creating successively all the different types of animals which have passed away, was to introduce man upon the surface of our globe. Man is the end towards which all the animal creation has tended from the first appearance of the first Palaeozoic fishes." The evolutionist, on the contrary (see EVOLUTION), maintains that different successive species of animals are in fact connected by parental descent, having become modified in the course of successive generations. The result of Charles Darwin's application of this theory to man may be given in his own words (_Descent of Man_,