Chapter 8

On Anthemius generally, see Procopius,De Aedific. i. 1; Agathias,Hist. v. 6-9;Gibbon’s Decline and Fall, cap. xl.

(T. L. H.)

ANTHESTERIA,one of the four Athenian festivals in honour of Dionysus, held annually for three days (11th-13th) in the month of Anthesterion (February-March). The object of the festival was to celebrate the maturing of the wine stored at the previous vintage, and the beginning of spring. On the first day, calledPithoigia(opening of the casks), libations were offered from the newly opened casks to the god of wine, all the household, including servants and slaves, joining in the festivities. The rooms and the drinking vessels in them were adorned with spring flowers, as were also the children over three years of age. The second day, namedChoës(feast of beakers), was a time of merrymaking. The people dressed themselves gaily, some in the disguise of the mythical personages in the suite of Dionysus, and paid a round of visits to their acquaintances. Drinking clubs met to drink off matches, the winner being he who drained his cup most rapidly. Others poured libations on the tombs of deceased relatives. On the part of the state this day was the occasion of a peculiarly solemn and secret ceremony in one of the sanctuaries of Dionysus in the Lenaeum, which for the rest of the year was closed. The basilissa (or basilinna), wife of the archon basileus for the time, went through a ceremony of marriage to the wine god, in which she was assisted by fourteen Athenian matrons, calledgeraerae, chosen by the basileus and sworn to secrecy. The days on which the Pithoigia and Choës were celebrated were both regarded asἀποφράδες(nefasti) andμιαραί(“defiled”), necessitating expiatory libations; on them the souls of the dead came up from the underworld and walked abroad; people chewed leaves of whitethorn and besmeared their doors with tar to protect themselves from evil. But at least in private circles the festive character of the ceremonies predominated. The third day was namedChytri(feast of pots, fromχύτρος, a pot), a festival of the dead. Cooked pulse was offered to Hermes, in his capacity of agod of the lower world, and to the souls of the dead. Although no performances were allowed at the theatre, a sort of rehearsal took place, at which the players for the ensuing dramatic festival were selected.

The name Anthesteria, according to the account of it given above, is usually connected withἄνθος(“flower,” or the “bloom” of the grape), but A.W. Verrall (Journal of Hellenic Studies, xx., 1900, p. 115) explains it as a feast of “revocation” (fromἀναθέσσασθαι, to “pray back” or “up”), at which the ghosts of the dead were recalled to the land of the living (cp.the Romanmundus patet). J.E. Harrison (ibid. 100, 109, andProlegomena), regarding the Anthesteria as primarily a festival of all souls, the object of which was the expulsion of ancestral ghosts by means of placation, explainsπιθοιγίαas the feast of the opening of the graves (πίθοςmeaning a large urn used for burial purposes),χόεςas the day of libations, andχύτροιas the day of the grave-holes (not “pots,” which isχύτροι), in point of time really anterior to theπιθοιγία. E. Rohde and M.P. Nilsson, however, take theχύτροιto mean “water vessels,” and connect the ceremony with the Hydrophoria, a libation festival to propitiate the dead who had perished in the flood of Deucalion.

See F. Hiller von Gartringen in Pauly-Wissowa’sRealencyclopadie(s.v.); J. Girard in Daremberg and Saglio,Dictionnaire des antiquités(s.v.“Dionysia”); and F.A. Voigt in Roscher’sLexikon der Mythologie(s.v.“Dionysos”); J.E. Harrison,Prolegomena to the Study of Greek Religion(1903); M.P. Nilsson,Studia de Dionysiis Atticis(1900) andGriechische Feste(1906); G.F. Schömann,Griechische Alterthümer, ii. (ed. J.H. Lipsius, 1902), p. 516; A. Mommsen,Feste der Stadt Athen(1898); E. Rohde,Psyche(4th ed., 1907), p. 237.

ANTHIM THE IBERIAN,a notable figure in the ecclesiastical history of Rumania. A Georgian by birth, he came to Rumania early in the second half of the 17th century, as a simple monk. He became bishop of Râmnicu in 1705, and in 1708 archbishop of Walachia. Taking a leading part in the political movements of the time, he came into conflict with the newly appointed Greek hospodars, and was exiled to Rumelia. But on his crossing the Danube in 1716 he was thrown into the water and drowned, as it is alleged, at the instigation of the prince of Walachia. He was a man of great talents and spoke and wrote many Oriental and European languages. Though a foreigner, he soon acquired a thorough knowledge of Rumanian, and was instrumental in helping to introduce that language into the church as its official language. He was a master printer and an artist of the first order. He cut the wood blocks for the books which he printed in Tirgovishtea, Râmnicu, Snagov and Bucharest. He was also the first to introduce Oriental founts of type into Rumania, and he printed there the first Arabic missal for the Christians of the East (Râmnicu, 1702). He also trained Georgians in the art of printing, and cut the type with which under his pupil Mihail Ishtvanovitch they printed the first Georgian Gospels (Tiflis, 1709). A man of great oratorical power, Anthim delivered a series of sermons (Didahii), and some of his pastoral letters are models of style and of language as well as of exact and beautiful printing. He also completed a wholecorpusof lectionaries, missals, gospels, &c.

See M. Gaster,Chrestomathie roumaine(1881), and “Gesch. d. rumänischen Litteratur,” in Grober,Grundriss d. rom. Philologie, vol. ii. (1899); and E. Picot,Notice sur Anthim d’Ivir(Paris, 1886).

(M. G.)

ANTHOLOGY.The term “anthology,” literally denoting a garland or collection of flowers, is figuratively applied to any selection of literary beauties, and especially to that great body of fugitive poetry, comprehending about 4500 pieces, by upwards of 300 writers, which is commonly known as theGreek Anthology.

Literary History of the Greek Anthology.—The art of occasional poetry had been cultivated in Greece from an early period,—less, however, as the vehicle of personal feeling, than as the recognized commemoration of remarkable individuals or events, on sepulchral monuments and votive offerings: Such compositions were termed epigrams,i.e.inscriptions. The modern use of the word is a departure from the original sense, which simply indicated that the composition was intended to be engraved or inscribed. Such a composition must necessarily be brief, and the restraints attendant upon its publication concurred with the simplicity of Greek taste in prescribing conciseness of expression, pregnancy of meaning, purity of diction and singleness of thought, as the indispensable conditions of excellence in the epigrammatic style. The term was soon extended to any piece by which these conditions were fulfilled. The transition from the monumental to the purely literary character of the epigram was favoured by the exhaustion of more lofty forms of poetry, the general increase, from the general diffusion of culture, of accomplished writers and tasteful readers, but, above all, by the changed political circumstances of the times, which induced many who would otherwise have engaged in public affairs to addict themselves to literary pursuits. These causes came into full operation during the Alexandrian era, in which we find every description of epigrammatic composition perfectly developed. About 60B.C., the sophist and poet, Meleager of Gadara, undertook to combine the choicest effusions of his predecessors into a single body of fugitive poetry. Collections of monumental inscriptions, or of poems on particular subjects, had previously been formed by Polemon Periegetes and others; but Meleager first gave the principle a comprehensive application. His selection, compiled from forty-six of his predecessors, and including numerous contributions of his own, was entitledThe Garland(Στέφανος); and in an introductory poem each poet is compared to some flower, fancifully deemed appropriate to his genius. The arrangement of his collection was alphabetical, according to the initial letter of each epigram.

In the age of the emperor Tiberius (or Trajan, according to others) the work of Meleager was continued by another epigrammatist, Philippus of Thessalonica, who first employed the term anthology. His collection, which included the compositions of thirteen writers subsequent to Meleager, was also arranged alphabetically, and contained an introductory poem. It was of inferior quality to Meleager’s. Somewhat later, under Hadrian, another supplement was formed by the sophist Diogenianus of Heracleia (2nd centuryA.D.), and Strato of Sardis compiled his elegant but taintedΜοῦσα Παιδική(Musa Puerilis) from his productions and those of earlier writers. No further collection from various sources is recorded until the time of Justinian, when epigrammatic writing, especially of an amatory character, experienced a great revival at the hands of Agathias of Myrina, the historian, Paulus Silentiarius, and their circle. Their ingenious but mannered productions were collected by Agathias into a new anthology, entitledThe Circle(Κύκλος); it was the first to be divided into books, and arranged with reference to the subjects of the pieces.

These and other collections made during the middle ages are now lost. The partial incorporation of them into a single body, classified according to the contents in 15 books, was the work of a certain Constantinus Cephalas, whose name alone is preserved in the single MS. of his compilation extant, but who probably lived during the temporary revival of letters under Constantine Porphyrogenitus, at the beginning of the 10th century. He appears to have merely made excerpts from the existing anthologies, with the addition of selections from Lucillius, Palladas, and other epigrammatists, whose compositions had been published separately. His arrangement, to which we shall have to recur, is founded on a principle of classification, and nearly corresponds to that adopted by Agathias. His principle of selection is unknown; it is only certain that while he omitted much that he should have retained, he has preserved much that would otherwise have perished. The extent of our obligations may be ascertained by a comparison between his anthology and that of the next editor, the monk Maximus Planudes (A.D.1320), who has not merely grievously mutilated the anthology of Cephalas by omissions, but has disfigured it by interpolating verses of his own. We are, however, indebted to him for the preservation of the epigrams on works of art, which seem to have been accidentally omitted from our only transcript of Cephalas.

The Planudean (in seven books) was the only recension of the anthology known at the revival of classical literature, and was first published at Florence, by Janus Lascaris, in 1494. It long continuedto be the only accessible collection, for although the Palatine MS., the sole extant copy of the anthology of Cephalas, was discovered in the Palatine library at Heidelberg, and copied by Saumaise (Salmasius) in 1606, it was not published until 1776, when it was included in Brunck’sAnalecta Veterum Poetarum Graecorum. The MS. itself had frequently changed its quarters. In 1623, having been taken in the sack of Heidelberg in the Thirty Years’ War, it was sent with the rest of the Palatine Library to Rome as a present from Maximilian I. of Bavaria to Gregory XV., who had it divided into two parts, the first of which was by far the larger; thence it was taken to Paris in 1797. In 1816 it went back to Heidelberg, but in an incomplete state, the second part remaining at Paris. It is now represented at Heidelberg by a photographic facsimile. Brunck’s edition was superseded by the standard one of Friedrich Jacobs (1794-1814, 13 vols.), the text of which was reprinted in a more convenient form in 1813-1817, and occupies three pocket volumes in the Tauchnitz series of the classics. The best edition for general purposes is perhaps that of Dubner in Didot’sBibliotheca(1864-1872), which contains the Palatine Anthology, the epigrams of the Planudean Anthology not comprised in the former, an appendix of pieces derived from other sources, copious notes selected from all quarters, a literal Latin prose translation by Boissonade, Bothe, and Lapaume and the metrical Latin versions of Hugo Grotius. A third volume, edited by E. Cougny, was published in 1890. The best edition of the Planudean Anthology is the splendid one by van Bosch and van Lennep (1795-1822). There is also a complete edition of the text by Stadlmuller in the Teubner series.

Arrangement.—The Palatine MS., the archetype of the present text, was transcribed by different persons at different times, and the actual arrangement of the collection does not correspond with that signalized in the index. It is as follows: Book 1. Christian epigrams; 2. Christodorus’s description of certain statues; 3. Inscriptions in the temple at Cyzicus; 4. The prefaces of Meleager, Philippus, and Agathias to their respective collections; 5. Amatory epigrams; 6. Votive inscriptions; 7. Epitaphs; 8. The epigrams of Gregory of Nazianzus; 9. Rhetorical and illustrative epigrams; 10. Ethical pieces; 11. Humorous and convivial; 12. Strata’s Musa Puerilis; 13. Metrical curiosities; 14. Puzzles, enigmas, oracles; 15. Miscellanies. The epigrams on works of art, as already stated, are missing from theCodex Palatinus, and must be sought in an appendix of epigrams only occurring in the Planudean Anthology. The epigrams hitherto recovered from ancient monuments and similar sources form appendices in the second and third volumes of Dübner’s edition.

Style and Value.—One of the principal claims of the Anthology to attention is derived from its continuity, its existence as a living and growing body of poetry throughout all the vicissitudes of Greek civilization. More ambitious descriptions of composition speedily ran their course, and having attained their complete development became extinct or at best lingered only in feeble or conventional imitations. The humbler strains of the epigrammatic muse, on the other hand, remained ever fresh and animated, ever in intimate union with the spirit of the generation that gave them birth. To peruse the entire collection, accordingly, is as it were to assist at the disinterment of an ancient city, where generation has succeeded generation on the same site, and each stratum of soil enshrines the vestiges of a distinct epoch, but where all epochs, nevertheless, combine to constitute an organic whole, and the transition from one to the other is hardly perceptible. Four stages may be indicated:—1. The Hellenic proper, of which Simonides of Ceos (c.556-469B.C.), the author of most of the sepulchral inscriptions on those who fell in the Persian wars, is the characteristic representative. This is characterized by a simple dignity of phrase, which to a modern taste almost verges upon baldness, by a crystalline transparency of diction, and by an absolute fidelity to the original conception of the epigram. Nearly all the pieces of this era are actualbona fideinscriptions or addresses to real personages, whether living or deceased; narratives, literary exercises, and sports of fancy are exceedingly rare. 2. The epigram received a great development in its second or Alexandrian era, when its range was so extended as to include anecdote, satire, and amorous longing; when epitaphs and votive inscriptions were composed on imaginary persons and things, and men of taste successfully attempted the same subjects in mutual emulation, or sat down to compose verses as displays of their ingenuity. The result was a great gain in richness of style and general interest, counterbalanced by a falling off in purity of diction and sincerity of treatment. The modification—a perfectly legitimate one, the resources of the old style being exhausted—had its real source in the transformation of political life, but may be said to commence with and to find its best representative in the playful and elegant Leonidas of Tarentum, a contemporary of Pyrrhus, and to close with Antipater of Sidon, about 140B.C.(or later). It should be noticed, however, that Callimachus, one of the most distinguished of the Alexandrian poets, affects the sternest simplicity in his epigrams, and copies the austerity of Simonides with as much success as an imitator can expect. 3. By a slight additional modification in the same direction, the Alexandrian passes into what, for the sake of preserving the parallelism with eras of Greek prose literature, we may call the Roman style, although the peculiarities of its principal representative are decidedly Oriental. Meleager of Gadara was a Syrian; his taste was less severe, and his temperament more fervent than those of his Greek predecessors; his pieces are usually erotic, and their glowing imagery sometimes reminds us of the Song of Solomon. The luxuriance of his fancy occasionally betrays him into far-fetched conceits, and the lavishness of his epithets is only redeemed by their exquisite felicity. Yet his effusions are manifestly the offspring of genuine feeling, and his epitaph on himself indicates a great advance on the exclusiveness of antique Greek patriotism, and is perhaps the first clear enunciation of the spirit of universal humanity characteristic of the later Stoic philosophy. His gaiety and licentiousness are imitated and exaggerated by his somewhat later contemporary, the Epicurean Philodemus, perhaps the liveliest of all the epigrammatists; his fancy reappears with diminished brilliancy in Philodemus’s contemporary, Zonas, in Crinagoras, who wrote under Augustus, and in Marcus Argentarius, of uncertain date; his peculiar gorgeousness of colouring remains entirely his own. At a later period of the empire anothergenre, hitherto comparatively in abeyance, was developed, the satirical. Lucillius, who flourished under Nero, and Lucian, more renowned in other fields of literature, display a remarkable talent for shrewd, caustic epigram, frequently embodying moral reflexions of great cogency, often lashing vice and folly with signal effect, but not seldom indulging in mere trivialities, or deformed by scoffs at personal blemishes. This style of composition is not properly Greek, but Roman; it answers to the modern definition of epigram, and has hence attained a celebrity in excess of its deserts. It is remarkable, however, as an almost solitary example of direct Latin influence on Greek literature. The same style obtains with Palladas, an Alexandrian grammarian of the 4th century, the last of the strictly classical epigrammatists, and the first to be guilty of downright bad taste. His better pieces, however, are characterized by an austere ethical impressiveness, and his literary position is very interesting as that of an indignant but despairing opponent of Christianity. 4. The fourth or Byzantine style of epigrammatic composition was cultivated by thebeaux-espritsof the court of Justinian. To a great extent this is merely imitative, but the circumstances of the period operated so as to produce a species of originality. The peculiarly ornate andrecherchédiction of Agathias and his compeers is not a merit in itself, but, applied for the first time, it has the effect of revivifying an old form, and many of their new locutions are actual enrichments of the language. The writers, moreover, were men of genuine poetical feeling, ingenious in invention, and capable of expressing emotion with energy and liveliness; the colouring of their pieces is sometimes highly dramatic.

It would be hard to exaggerate the substantial value of the Anthology, whether as a storehouse of facts bearing on antique manners, customs and ideas, or as one among the influences which have contributed to mould the literature of the modern world. The multitudinous votive inscriptions, serious and sportive, connote the phases of Greek religious sentiment, from pious awe to irreverent familiarity and sarcastic scepticism; the moral tone of the nation at various periods is mirrored with corresponding fidelity; the sepulchral inscriptions admit us intothe inmost sanctuary of family affection, and reveal a depth and tenderness of feeling beyond the province of the historian to depict, which we should not have surmised even from the dramatists; the general tendency of the collection is to display antiquity on its most human side, and to mitigate those contrasts with the modern world which more ambitious modes of composition force into relief. The constant reference to the details of private life renders the Anthology an inexhaustible treasury for the student of archaeology; art, industry and costume receive their fullest illustration from its pages. Its influence on European literatures will be appreciated in proportion to the inquirer’s knowledge of each. The further his researches extend, the greater will be his astonishment at the extent to which the Anthology has been laid under contribution for thoughts which have become household words in all cultivated languages, and at the beneficial effect of the imitation of its brevity, simplicity, and absolute verbal accuracy upon the undisciplined luxuriance of modern genius.

Translations, Imitations, &c.—The best versions of the Anthology ever made are the Latin renderings of select epigrams by Hugo Grotius. They have not been printed separately, but will be found in Bosch and Lennep’s edition of the PlanudeanAnthology, in the Didot edition, and in Dr Wellesley’sAnthologia Polyglotta. The number of more or less professed imitations in modern languages is infinite, that of actual translations less considerable. French and Italian, indeed, are ill adapted to this purpose, from their incapacity of approximating to the form of the original, and their poets have usually contented themselves with paraphrases or imitations, often exceedingly felicitous. F.D. Dehèque’s French prose translation, however (1863), is most excellent and valuable. The German language alone admits of the preservation of the original metre—a circumstance advantageous to the German translators, Herder and Jacobs, who have not, however, compensated the loss inevitably consequent upon a change of idiom by any added beauties of their own. Though unfitted to reproduce the precise form, the English language, from its superior terseness, is better adapted to preserve the spirit of the original than the German; and the comparative ill success of many English translators must be chiefly attributed to the extremely low standard of fidelity and brevity observed by them. Bland, Merivale, and their associates (1806-1813), are often intolerably diffuse and feeble, from want, not of ability, but of taking pains. Archdeacon Wrangham’s too rare versions are much more spirited; and John Sterling’s translations of the inscriptions of Simonides deserve high praise. Professor Wilson (Blackwood’s Magazine, 1833-1835) collected and commented upon the labours of these and other translators, with his accustomed critical insight and exuberant geniality, but damaged his essay by burdening it with the indifferent attempts of William Hay. In 1849 Dr Wellesley, principal of New Inn Hall, Oxford, published hisAnthologia Polyglotta, a most valuable collection of the best translations and imitations in all languages, with the original text. In this appeared some admirable versions by Goldwin Smith and Dean Merivale, which, with the other English renderings extant at the time, will be found accompanying the literal prose translation of thePublic School Selections, executed by the Rev. George Burges for Bohn’s Classical Library (1854). This is a useful volume, but the editor’s notes are worthless. In 1864 Major R.G. Macgregor published an almost complete translation of the Anthology, a work whose stupendous industry and fidelity almost redeem the general mediocrity of the execution.Idylls and Epigrams, by R. Garnett (1869, reprinted 1892 in the Cameo series), includes about 140 translations or imitations, with some original compositions in the same style. Recent translations (selections) are: J.W. Mackail,Select Epigrams from the Greek Anthology(with text, introduction, notes, and prose translation), 1890, revised 1906, a most charming volume; Graham R. Tomson (Mrs Marriott Watson),Selections from the Greek Anthology(1889); W.H.D. Rouse,Echo of Greek Song(1899); L.C. Perry,From the Garden of Hellas(New York, 1891); W.R. Paton,Love Epigrams(1898). An agreeable little volume on the Anthology, by Lord Neaves, is one of Collins’s series ofAncient Classics for Modern Readers. The earl of Cromer, with all the cares of Egyptian administration upon him, found time to translate and publish an elegant volume of selections (1903). Two critical contributions to the subject should be noticed, the Rev. James Davies’s essay on Epigrams in theQuarterly Review(vol. cxvii.), especially valuable for its lucid illustration of the distinction between Greek and Latin epigram; and the brilliant disquisition in J.A. Symonds’sStudies of the Greek Poets(1873; 3rd ed., 1893).

Latin Anthology.—TheLatin Anthologyis the appellation bestowed upon a collection of fugitive Latin verse, from the age of Ennius to aboutA.D.1000, formed by Peter Burmann the Younger. Nothing corresponding to the Greek anthology is known to have existed among the Romans, though professional epigrammatists like Martial published their volumes on their own account, and detached sayings were excerpted from authors like Ennius and Publius Syrus, while thePriapeïawere probably but one among many collections on special subjects. The first general collection of scattered pieces made by a modern scholar was Scaliger’sCatalecta veterum Poetarum(1573), succeeded by the more ample one of Pithoeus,Epigrammata et Poemata e Codicibus et Lapidibus collecta(1590). Numerous additions, principally from inscriptions, continued to be made, and in 1759-1773 Burmann digested the whole into hisAnthologia veterum Latinorum Epigrammatum et Poematum. This, occasionally reprinted, was the standard edition until 1869, when Alexander Riese commenced a new and more critical recension, from which many pieces improperly inserted by Burmann are rejected, and his classified arrangement is discarded for one according to the sources whence the poems have been derived. The first volume contains those found in MSS., in the order of the importance of these documents; those furnished by inscriptions following. The first volume (in two parts) appeared in 1869-1870, a second edition of the first part in 1894, and the second volume,Carmina Epigraphica(in two parts), in 1895-1897, edited by F. Bücheler. AnAnthologiae Latinae Supplementa, in the same series, followed. Having been formed by scholars actuated by no aesthetic principles of selection, but solely intent on preserving everything they could find, the Latin anthology is much more heterogeneous than the Greek, and unspeakably inferior. The really beautiful poems of Petronius and Apuleius are more properly inserted in the collected editions of their writings, and more than half the remainder consists of the frigid conceits of pedantic professional exercises of grammarians of a very late period of the empire, relieved by an occasional gem, such as the apostrophe of the dying Hadrian to his spirit, or the epithalamium of Gallienus. The collection is also, for the most part, too recent in date, and too exclusively literary in character, to add much to our knowledge of classical antiquity. The epitaphs are interesting, but the genuineness of many of them is very questionable.

(R. G.)

ANTHON, CHARLES(1797-1867), American classical scholar, was born in New York city on the 19th of November 1797. After graduating with honours at Columbia College in 1815, he began the study of law, and in 1819 was admitted to the bar, but never practised. In 1820 he was appointed assistant professor of Greek and Latin in his old college, full professor ten years later, and at the same time headmaster of the grammar school attached to the college, which post he held until 1864. He died at New York on the 29th of July 1867. He produced for use in colleges and schools a large number of classical works, which enjoyed great popularity, although his editions of classical authors were by no means in favour with schoolmasters, owing to the large amount of assistance, especially translations, contained in the notes.

ANTHONY, SAINT,the first Christian monk, was born in Egypt about 250. At the age of twenty he began to practise an ascetical life in the neighbourhood of his native place, and after fifteen years of this life he withdrew into solitude to a mountain by the Nile, called Pispir, now Der el Memun, opposite Arsinoë in the Fayum. Here he lived strictly enclosed in an old fort for twenty years. At last in the early years of the 4th century he emerged from his retreat and set himself to organize the monastic life of the crowds of monks who had followed him and taken up their abode in the caves around him. After a time, again in pursuit of more complete solitude, he withdrew to the mountain by the Red Sea, where now stands the monastery that bears his name (Der Mar Antonios). Here he died about the middle of the 4th century. HisLifestates that on two occasions he went to Alexandria, to strengthen the Christians in the Diocletian persecution and to preach against Arianism. Anthony is recognized as the first Christian monk and the first organizer and father of Christian monachism (seeMonasticism). Certain letters and sermons are attributed to him, but their authenticity is more than doubtful. The monastic rule which bears his name was not written by him, but was compiled out of these writingsand out of discourses and utterances put into his mouth in theLifeand theApophthegmata Patrum. According to this rule live a number of Coptic Syrian and Armenian monks to this day. The chief source of information about St Anthony is theLife, attributed to St Athanasius. This attribution, as also the historical character of the book, and even the very existence of St Anthony, were questioned and denied by the sceptical criticism of thirty years ago; but such doubts are no longer entertained by critical scholars.

The GreekVitais among the works of St Athanasius; the almost contemporary Latin translation is among Rosweyd’sVitae Patrum(Migne,Patrol. Lat. lxxiii.); an English translation is in the Athanasius volume of the “Nicene and Post-Nicene Library.” Accounts of St Anthony are given by Card. Newman,Church of the Fathers(Historical Sketches) and Alban Butler,Lives of the Saints(Jan. 17). Discussions of the historical and critical questions raised will be found in E.C. Butler’sLausiac History of Palladius(1898, 1904), Part I. pp. 197, 215-228; 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. Lepître,St Antoine de Padoue(Paris, 1902, inLes Saintsseries: good bibliography; Eng. trans. by Edith Guest, London, 1902), and by Léopold de Chérancé,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 theAnnales Minorumof 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 publishedThe 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’sLife 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 theAnthozoa, 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 orcorallum, as a good example of the structure of a typical Anthozoan polyp or zooid. The individual animal or zooid ofActinia equinahas the form of a column fixed by one extremity, called thebase, to a rock or other object, and bearing at the opposite extremity a crown oftentacles. The tentacles surround an area known as theperistome, 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 thestomodaeum, which in its turn opens below into the general body-cavity orcoelenteron. 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 thesulcusandsulculus, 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 asmesenteries, 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 amesenterial filament. The reproductiveorgans or gonads are borne on the mesenteries, the germinal cells being derived from the inner layer or endoderm.

m, Mesentery.

t, Tentacles.

st, Stomodaeum.

sc, Sulcus.

r, Rotteken’s muscle.

s, Stoma.

lm, Longitudinal muscle.

d, Diagonal Muscle.

go, Gonads.

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 aregland cells, sense cells, andcnidoblasts. The last-named are specially numerous on the tentacles and on some other regions of the body, and produce the well-known “thread cells,” ornematocysts, 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. InActinia equinathe 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.

The Anthozoa are divisible into two sub-classes, sharply marked off from one another by definite anatomical characters. These are theAlcyonariaand theZoantharia. 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.

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. Thesemuscle-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).Nearly all the Alcyonaria are colonial. Four solitary species have been described, viz.Haimea funebrisandH. hyalina, Hartea elegans, andMonoxenia 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, calledsolenia. 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 genusCornularia, 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 whichzooids arise at intervals. InClavulariaand 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 calledstolons, and a stolon may be simple,i.e.contain only one solenium, as inCornularia, or may be complex and built up of many solenia, as inClavularia. 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—theStolonifera, Alcyonacea, Pseudaxonia, Axifera, Stelechotokea, andCornothecalia.Fig.5.A. Skeleton of a young colony ofTubipora purpurea. st, Stolon;p, platform.B. Diagrammatic longitudinal section of a corallite, showing two platforms,pand cup-shaped tabulae,t. (After S.J. Hickson.)Fig.6.—Portion of a colony ofCoralinum rubrum, showing expanded and contracted zooids. In the lower part of the figure the cortex has been cut away to show theaxis, ax, and the longitudinal canals,lc, surrounding it.In the orderStolonirerathe 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 genusTubiporathese horizontal solenia unite to form a series of horizontal platforms (fig. 5). The order comprises the familiesCornulamdae, Syringopordae, Tubipondae, andFavositidae. In the first-named, the zooids are united only by their bases and the skeleton consists of loose spicules. In theTubipondaethe 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 atabula. 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. InFavosites hemisphaericaa number of radial spines, projecting into the cavity of the corallite, give it the appearance of a madreporarian coral.In the orderAlcyonaceathe 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 familiesXeniidae, AlcyonidaeandNephthyidae.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.Fig.7.—The sea-fan (Gorgonia cavolinii).Fig.8.A. Colony ofPennatula phosphoreafrom 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 orderPseudaxoniathe 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—theBriareidae, Sclerogorgidae, Melitodidae, andCorallidae. 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 theMelitodidaeandCorallidaeit forms a dense axis, which in theMelitodidaeconsists of alternate calcareous and horny joints. The precious red coral of commerce,Corallium rubrum(fig. 6), a member of the familyCorallidae, 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 rubrumhas been the subject of a beautifully-illustrated memoir by de Lacaze-Duthiers, which should be consulted for details of anatomy.TheAxiferacomprise 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. InGorgoniathe axis remains horny throughout life, but in many forms it is further strengthened by a deposit of calcareous matter In the familyIsidinaethe axis consists of alternate segments of horny and calcareous substance, the latter being amorphous. The order contains six families—theDasygorgidae, Isidae, Primnoidae, Muriceidae, Plexauridae, andGorgoniaae.In the orderStelechotokeathe 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 sectionAsiphonaceathe 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, theTelestidaeand theCoelogorgidae. The second section comprises thePennatulaceaor sea-pens, which are remarkable from the fact that the colony is not fixed by the base to a rock or otherobject, 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.Fig.9.A, Portion of the surface of a colony ofHeliopora coeruleamagnified, 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 orderCoenothecaliais 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, calledcalices, 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 thecoenenchymal 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 ortabulae, 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 coeruleais found between tide-marks on the shore platforms of coral islands. The order was more abundantly represented in Palaeozoic times by theHeliolitidaefrom the Upper and Lower Silurian and the Devonian, and by theThecidaefrom the Wenlock limestone. InHeliolites porosusthe colonies had the form of spheroidal masses; the calices were furnished with twelve pseudosepta, and the coenenchymal tubes were more or less regularly hexagonal.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 ofHalcampa.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 genusEdwardsia(fig. 10, A, B). This is a small solitary Zoantharian which lives embedded in sand. Its body is divisible into three portions, an uppercapitulumbearing the mouth and tentacles, a medianscapuscovered 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 ofEdwardsiaare 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 anentocoele; the space enclosed between two mesenteries of adjacent couples is called anexocoele. 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 arevis-à-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.Fig.11.—A, Diagram showing the sequence of mesenterial development in an Actinian. B, Diagrammatic transverse section ofGonactinia prolifera.While the mesenterial couples belonging to the second and each successive cycle are formed simultaneously, those of the first cycleare 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 thesulcularfaces. 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. InActinia equinathe 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.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 asActinia 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 orderZoanthideacomprises 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 orderCerianthideacomprises 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 genusCerianthusfloat 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.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 ofArachnactis brachiolata, the larva ofCerianthus, with seven tentacles.D, Transverse section of an older larva. The numerals indicate the order of development of the mesenteries.The orderAntipathideais 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 inLeiopathes glaberrima, a genus with twelve mesenteries, there are distinct indications of an Edwardsia stage.Fig.14.A, Portion of a colony ofAntipathes dichotoma.B, Single zooid and axis of the same magnified.m, Mouth;mfmesenterial filament;ax, axis.C, Transverse section through the oral cone ofAntipathella minor, st, Stomodaeum;ov, ovary.There are, in addition to these groups, several genera of Actinians whose mesenterial arrangement differs from the normal type. Ofthese perhaps the most interesting isGonactinia 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, theActiniidea, 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.

Nearly all the Alcyonaria are colonial. Four solitary species have been described, viz.Haimea funebrisandH. hyalina, Hartea elegans, andMonoxenia 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, calledsolenia. 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 genusCornularia, 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 whichzooids arise at intervals. InClavulariaand 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 calledstolons, and a stolon may be simple,i.e.contain only one solenium, as inCornularia, or may be complex and built up of many solenia, as inClavularia. 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—theStolonifera, Alcyonacea, Pseudaxonia, Axifera, Stelechotokea, andCornothecalia.

A. Skeleton of a young colony ofTubipora purpurea. st, Stolon;p, platform.

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

In the orderStolonirerathe 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 genusTubiporathese horizontal solenia unite to form a series of horizontal platforms (fig. 5). The order comprises the familiesCornulamdae, Syringopordae, Tubipondae, andFavositidae. In the first-named, the zooids are united only by their bases and the skeleton consists of loose spicules. In theTubipondaethe 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 atabula. 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. InFavosites hemisphaericaa number of radial spines, projecting into the cavity of the corallite, give it the appearance of a madreporarian coral.

In the orderAlcyonaceathe 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 familiesXeniidae, AlcyonidaeandNephthyidae.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.

A. Colony ofPennatula phosphoreafrom 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 orderPseudaxoniathe 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—theBriareidae, Sclerogorgidae, Melitodidae, andCorallidae. 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 theMelitodidaeandCorallidaeit forms a dense axis, which in theMelitodidaeconsists of alternate calcareous and horny joints. The precious red coral of commerce,Corallium rubrum(fig. 6), a member of the familyCorallidae, 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 rubrumhas been the subject of a beautifully-illustrated memoir by de Lacaze-Duthiers, which should be consulted for details of anatomy.

TheAxiferacomprise 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. InGorgoniathe axis remains horny throughout life, but in many forms it is further strengthened by a deposit of calcareous matter In the familyIsidinaethe axis consists of alternate segments of horny and calcareous substance, the latter being amorphous. The order contains six families—theDasygorgidae, Isidae, Primnoidae, Muriceidae, Plexauridae, andGorgoniaae.

In the orderStelechotokeathe 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 sectionAsiphonaceathe 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, theTelestidaeand theCoelogorgidae. The second section comprises thePennatulaceaor sea-pens, which are remarkable from the fact that the colony is not fixed by the base to a rock or otherobject, 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.

A, Portion of the surface of a colony ofHeliopora coeruleamagnified, 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 orderCoenothecaliais 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, calledcalices, 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 thecoenenchymal 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 ortabulae, 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 coeruleais found between tide-marks on the shore platforms of coral islands. The order was more abundantly represented in Palaeozoic times by theHeliolitidaefrom the Upper and Lower Silurian and the Devonian, and by theThecidaefrom the Wenlock limestone. InHeliolites porosusthe colonies had the form of spheroidal masses; the calices were furnished with twelve pseudosepta, and the coenenchymal tubes were more or less regularly hexagonal.

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 ofHalcampa.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 genusEdwardsia(fig. 10, A, B). This is a small solitary Zoantharian which lives embedded in sand. Its body is divisible into three portions, an uppercapitulumbearing the mouth and tentacles, a medianscapuscovered 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 ofEdwardsiaare 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 anentocoele; the space enclosed between two mesenteries of adjacent couples is called anexocoele. 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 arevis-à-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.

Fig.11.—A, Diagram showing the sequence of mesenterial development in an Actinian. B, Diagrammatic transverse section ofGonactinia prolifera.

While the mesenterial couples belonging to the second and each successive cycle are formed simultaneously, those of the first cycleare 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 thesulcularfaces. 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. InActinia equinathe 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.

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 asActinia 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 orderZoanthideacomprises 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 orderCerianthideacomprises 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 genusCerianthusfloat 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.

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 ofArachnactis brachiolata, the larva ofCerianthus, with seven tentacles.

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

The orderAntipathideais 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 inLeiopathes glaberrima, a genus with twelve mesenteries, there are distinct indications of an Edwardsia stage.

A, Portion of a colony ofAntipathes dichotoma.

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

C, Transverse section through the oral cone ofAntipathella minor, st, Stomodaeum;ov, ovary.

There are, in addition to these groups, several genera of Actinians whose mesenterial arrangement differs from the normal type. Ofthese perhaps the most interesting isGonactinia 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, theActiniidea, 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.

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 orsepta, 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 ascalicoblasts. The skeleton or corallum of a typical solitary coral—the common Devonshire cup-coralCaryophyllia smithii(fig. 15) is a good example—exhibits the followings parts:—(1) Thebasal plate, between the zooid and the surface of attachment. (2) Thesepta, radial plates of calcite reaching from the periphery nearly or quite to the centre of the coral-cup or calicle. (3) Thethecaor wall, which in many corals is not an independent structure, but is formed by the conjoined thickened peripheral ends of the septa. (4) Thecolumella, 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) Thecostae, 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:—Dissepimentsare 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 calledendotheca.Synapticulaeare calcareous bars uniting adjacent septa.Tabulaeare 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 ofMadrepora—only two septa are fully developed, the remainder being very feebly represented.

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