Chapter VI: Part 6
_History_.--The authentic history of the greater part of the country embraced in the Central Provinces does not begin till the 16th century A.D. By the people of northern India the country was known as Gondwana, after the savage tribes of Gonds by whom it was inhabited. The Mussulman invaders of the Deccan passed it by, not caring to enter its mountain fastnesses and impenetrable forests; though occasional inscriptions show that parts of it had fallen from time to time under the dominion of one or other of the great kingdoms of the north, e.g. of Asoka, of the Guptas of Maghada, or of the ancient Hindu kingdom of Vidarbha (Berar); and inscriptions and numerous discoveries of coins prove that, during the middle ages, the open spaces were occupied by a series of Rajput dynasties. Of these the most important was that of the Haihayas of Ratanpur, a family which, settled from time immemorial in the Nerbudda valley, had towards the close of the 10th century succeeded the Pandava dynasty of Maha Kosala (Chhattisgarh) and ruled, though from the 16th century onwards over greatly diminished territories, until its overthrow by the Mahrattas in 1745. The second ruler of this dynasty, Ratnaraja, was the founder of Ratanpur.
The inscriptional records cease abruptly in the 12th century, and no more is known of the country until the rise of the Gond dynasties from the 14th to the 16th centuries. The first of these is mentioned in 1398, when Narsingh Rai, raja of Kherla, is said by Ferishta to have ruled all the hills of Gondwana. He was finally overthrown and killed by Hoshang Shah, king of Malwa. The 16th century saw the establishment of a powerful Gond kingdom by Sangram Sah, who succeeded in 1480 as the 47th of the petty Gond rajas of Garha-Mandla, and extended his dominions so as to include Saugor and Damoh on the Vindhyan plateau, Jubbulpore and Narsinghpur in the Nerbudda valley, and Seoni on the Satpura highlands. Sangram Sah died in 1530; and the break up of his dominion began with the enforced cession to the Mogul emperor by Chandra Sah (1563-1575) of Saugor and Damoh and of that portion of his territories which afterwards formed the state of Bhopal.
About 200 years after Sangram Sah's time, Bakht Buland, the Gond chieftain of a principality seated at Deogarh in Chhindwara, having visited Delhi, set about introducing the civilization he had there admired. He founded the city of Nagpur, which his successor made his capital. The Deogarh kingdom, at its widest extent, embraced the modern districts of Betul, Chhindwara, Nagpur, with parts of Seoni, Bhandara and Balaghat. In the south of the province Chanda was the seat of another Gond dynasty, which first came into prominence in the 16th century. The three Gond principalities of Garha-Mandla, Deogarh and Chanda were nominally subject to the Mogul emperors. In addition to the acquisitions made in the north at the expense of Garha-Mandla, the Moguls, after the annexation of Berar, established governors at Paunar in Wardha and Kherla in Betul. Having thus hemmed in the Gond states, however, they made no efforts to assert any effective sovereignty over them; the Gond rajas for their part were content with practical independence within their own dominions. Under their peaceful rule their territories flourished, until the weakening of the Mogul empire and the rise of the predatory Bundela and Mahratta powers, with the organized forces of which their semi-barbarous feudal levies were unable to cope, brought misfortune upon them.
In the 17th century Chhatarsal, the Bundela chieftain, deprived the Mandla principality of part of the Vindhyan plateau and the Nerbudda valley. In 1733 the peshwa of Poona invaded Bundelkhand; and in 1735 the Mahrattas had established their power in Saugor. In 1742 the peshwa advanced to Mandla and exacted the payment of _chauth_ (tributary blackmail), and from this time until 1781, when the successors of Sangram Sah were finally overthrown, Garha-Mandla remained practically a Mahratta dependency. Meanwhile the other independent principalities of Gondwana had in turn succumbed. In 1743 Raghoji Bhonsla of Berar established himself at Nagpur, and by 1751 had conquered the territories of Deogarh, Chanda and Chhattisgarh. In 1741 Ratanpur had surrendered to the Mahratta leader Bhaskar Pant without a blow, and the ancient Rajput dynasty came to an end. In Chanda and Deogarh the Gond rajas were suffered by Raghoji Bhonsla and his successor to carry on a shadowy existence for a while, in order to give them an excuse for avoiding the claims of the peshwa as their overlord; though actually decisions in important matters were sought at Poona. Raghoji died in 1755, and in 1769 his son and successor, Janoji, was forced to acknowledge the peshwa's effective supremacy. The Nagpur state, however, continued to grow. In 1785 Mudhoji (d. 1788), Janoji's successor, bought from the Poona court the cession of Mandla and the upper Nerbudda valley, and between 1796 and 1798 this was followed by the acquisition of Hoshangabad and the larger part of Saugor and Damoh by Raghoji II. (d. 1816). Under this latter raja the Nagpur state covered practically the whole of the present Central Provinces and Berar, as well as Orissa and some of the Chota Nagpur states.
In 1803 Raghoji joined Sindhia against the British; the result was the defeat of the allies at Assaye and Argaon, and the treaty of Deogaon, by which Raghoji had to cede Cuttack, Sambalpur and part of Berar. Up to this time the rule of the Bhonsla rajas, rough warriors of peasant extraction, had been on the whole beneficent; but, soured by his defeat, Raghoji now set to work to recover some of his losses by a ruthless exploitation of the peasantry, and until the effective intervention of the British in 1818 the country was subjected to every kind of oppression. After Raghoji II.'s death in 1816 his imbecile son Parsaji was deposed and murdered by Mudhoji, known as Appa Sahib. In spite of a treaty signed with the British in this year, Mudhoji in 1817 joined the peshwa, but was defeated at Sitabaldi and forced to cede the rest of Berar to the nizam, and parts of Saugor and Damoh, with Mandla, Betul, Seoni and the Nerbudda valley, to the British. After a temporary restoration to the throne he was deposed, and Raghoji III., a grandchild of Raghoji II., was placed on the throne. During his minority, which lasted till 1840, the country was well administered by a British resident. In 1853, on the death of Raghoji III. without heirs, Nagpur lapsed to the British paramount power. Until the formation of the Central Provinces in 1861, Nagpur province, which consists of the present Nagpur division, Chhindwara and Chhatisgarh, was administered by a commissioner under the central government.
The territories in the north ceded in 1817 by the peshwa (parts of Saugor and Damoh) and in 1818 by Appa Sahib were in 1820 formed into the Saugor and Nerbudda Territories under an agent to the governor-general, and in 1835 were included in the newly formed North-West Provinces. In 1842, in consequence of a rising, they were again placed under the jurisdiction of an agent to the governor-general. Restored to the North-West Provinces in 1853, they were finally joined with the Nagpur province to constitute the new Central Provinces in 1861. On the 1st of October 1903 Berar also was placed under the administration of the commissioner of the Central Provinces (for history see BERAR). In 1905 the greater part of Sambalpur district, with the feudatory states of Bamra, Rairakhol, Sonpur, Patna and Kalahandi, were transferred to Bengal, while the feudatory states of Chang Bhakar, Korea, Surguja, Udaipur and Jashpur were transferred from Bengal to the Central Provinces.
During the decade 1891-1901 the Central Provinces suffered from famine more severely than any other part of India. The complete failure of the rain in the autumn of 1896 caused scarcity to develop suddenly into famine, which lasted until the end of 1897. The total number of persons in receipt of relief reached its maximum of nearly 700,000 in May 1897. The expenditure on relief alone was about a million sterling; and the total cost of the famine, including loss of revenue, amounted to nearly twice that amount. During 1897 the death-rate for the whole province rose to sixty-nine per thousand, or double the average, while the birth-rate fell to twenty-seven per thousand. The Central Provinces were stricken by another famine, yet more severe and widespread, caused by the complete failure of the rains in 1899. The maximum of persons relieved for the whole province was 1,971,000 in June 1900. In addition, about 68,000 persons were in receipt of relief in the native states. During the three years 1899-1902 the total expenditure on famine relief amounted to about four millions sterling. Berar also suffered from the famines of 1897 and 1900.
See _The Imperial Gazetteer of India_ (Oxford, 1908), x. 99, for list
of authorities.
CENTUMVIRI (_centum_, hundred; _vir_, man), an ancient court of civil jurisdiction at Rome, probably instituted by Servius Tullius.[1] Its antiquity is attested by the symbol and formula used in its procedure, the lance (_hasta_) as the sign of true ownership, the oath or wager (_sacramentum_), the ancient formula for recovery of property or assertion of liberty. It is probably alluded to in Livy's account of the Valerio-Horatian laws of 449 B.C. (Livy iii. 55, _Consules ... fecerunt sanciendo ut qui tribunis plebis, aedilibus, judicibus, decemviris nocuisset, ejus caput Jovi sacrum esset_). If the _judices_ here mentioned are the _centumviri_, it is clear that they formed a tribunal which represented the interests of the _plebs_. This is in accordance with Cicero's account (_de Orat._ i. 38. 173) of the sphere of their jurisdiction. He says this was mainly concerned with the property of which account was taken at the census; it was therefore in their power to make or unmake a citizen. They also decided questions concerning debt. Hence the _plebs_ had an interest in securing their decisions against undue influence. They were never regarded as magistrates, but merely as _judices_, and as such would be appointed for a fixed term of service by the magistrate, probably by the _praetor urbanus_. But in Cicero's time they were elected by the _Comitia Tributa_. They then numbered 105. Their original number is uncertain. It was probably increased by Augustus and in Pliny's time had reached 180. The office was probably open in quite early times to both patricians and plebeians. The term is also applied in the inscriptions of Veii to the municipal senates and Cures, which numbered 100 members.
AUTHORITIES.--Tigerström, _De Judicibus apud Romanos_ (Berlin, 1826);
Greenidge, _Legal Procedure of Cicero's Time_, pp. 40 ff., 58 ff., 182
ff., 264 (Oxford, 1901); Bethmann-Hollweg, _Der romische
Civilprozess_, ii. 53 ff. (Bonn, 1864); Pauly-Wissowa,
_Realencyclopadie_, iii. 1935 ff. (Wlassak). (A. M. Cl.)
FOOTNOTE:
[1] Mommsen (_Staatsrecht_, i³. 275, n. 4, ii³. 231, n. 1, 590 f.)
believed that the _Centumviri_ were instituted about 150 B.C.
CENTURION (Lat. _centurio_), in the ancient Roman army, an officer in command of a _centuria_, originally a body of a hundred infantry, later the sixtieth part of the normal legion. There were therefore in the legion sixty centurions, who, though theoretically subordinate to the six military tribunes, were the actual working officers of the legion. For the most part the centurions were promoted from the ranks: they were arranged in a complicated order of seniority; the senior centurion of the legion (_primus pilus_) was an officer of very high importance. Besides commanding the centuries of the legion, centurions were "seconded" for various kinds of special service, e.g. for staff employment, the command of auxiliaries. See further ROMAN ARMY.
CENTURIPE (formerly CENTORBI, anc. [Greek: Kentoripa] or _Centuripae_), a town of Sicily, in the province of Catania, situated 2380 ft. above sea-level in a commanding situation, 7 m. N. of the railway station of Catenanuova-Centuripe, which is 28 m. W. from Catania. Pop. (1901) 11,311. Thucydides mentions it as a city of the Sicels. It became an ally of the Athenians at the time of their expedition against Syracuse, and maintained its independence almost uninterruptedly (though it fell under the power of Agathocles) until the First Punic War. Cicero describes it, perhaps with some exaggeration, as being far the largest and richest city of Sicily, and as having a population of 10,000, engaged in the cultivation of an extensive territory. It was granted Latin rights before the rest of Sicily. It appears to have suffered much in the war against Sextus Pompeius, and not to have regained its former prosperity under the empire. Frederick II. entirely destroyed it in 1233, but it was soon rebuilt. Considerable remains of the ancient city walls and of buildings, mostly of the Roman period, still exist, and numerous antiquities, including some fine Hellenistic _terra-cottas_, have been discovered in casual excavations.
See F. Ansaldi, _I Monumenti dell' antica Centuripi_ (Catania, 1851);
P. Orsi in _Atti del Congresso Internazionale di Scienze Storiche_
(Rome, 1904), v. 177. (T. As.)
CENTURY (from Lat. _centuria_, a division of a hundred men), the name for a unit in the Roman army, originally amounting to one hundred men, and for one of the divisions into which the Roman people was separated for voting purposes (see COMITIA). The word is applied to any group of one hundred, and more particularly to a period of a hundred years, and to the successive periods of a hundred years, dating before or after the birth of Christ. The "Century-plant" is a name given to the Agave (q.v.), or American aloe, from the supposition that it flowered once only in every hundred years.
CEOS (Gr. [Greek: Keôs], mod. _Zea_ or _Tzia_), an island in the Aegean Sea, belonging to the group of the Cyclades and the eparchy of Syra, 14 m. off the coast of Attica. Its greatest length is about 15 m. and its breadth about 8 m. It rises gradually towards the centre, where it culminates in Mount Elias, 1864 ft. high. Among its natural productions are lemons, citrons, olives, wine and honey; it also exports a considerable quantity of valonia. There were formerly four towns of some importance in the island:--Iulis, about 3 m. from the north-west shore; Coressia, the harbour of Iulis, with a temple of Apollo Smintheus in the neighbourhood; Carthaea, in the south-east, with a temple of Apollo; and Poieëssa, in the south-west. Of these Iulis is represented by the town of Zea, and Carthaea by the village of 'S tais Polais; traces of the other two can still be made out. Iulis was the birthplace of the lyric poets Simonides and Bacchylides, the philosophers Prodicus and Ariston, and the physician Erasistratus; the excellence of its laws was so generally recognized that the title of Cean Laws passed into a proverb. One of them forbade a citizen to protract his life beyond sixty years. The people of Ceos fought on the Greek side at Artemisium and Salamis; they joined the Delian League and also the later Athenian alliance in 377 B.C. They revolted in 363-362, but were reduced again, and the Athenians established a monopoly of the ruddle, or red earth, which was one of the most valuable products of the island. In A.D. 1207 it was divided between four Italian adventurers; after forming part of the duchy of Naxos in 1537, it passed under Turkish rule in 1566. Silver coins of Carthaea and Coressia have been found dating from the 6th century B.C. (see NUMISMATICS: _Greek_, "Cyclades and Sporades"). The present population of the island is about 4000, of which the capital has about 2000.
See Pridik, _De Cei Insulae rebus_ (1892). (E. Gr.)
CEPHALIC INDEX, the term in use by anthropologists to express the percentage of breadth to length in any skull. The principle employed by Retzius is to take the longer diameter of a skull, the antero-posterior diameter, as 100; if the shorter or transverse diameter falls below 80 the skull may be classed as long (dolichocephalic), while if it exceeds 80 the skull is broad (brachycephalic) (see CRANIOMETRY).
CEPHALONIA (Ital. _Cefalonia_, ancient and modern official Greek _Cephallenia_, [Greek: Kephallênia]), an island belonging to the kingdom of Greece, and the largest of those known as the Ionian Islands, situated on the west side of the mainland, almost directly opposite the Gulf of Corinth. The name was traditionally derived from Cephalus, the Attic hero who was regarded as having colonized the island. The tradition, which is repeated by Aristotle, is probably due solely to the similarity of the names (see J.G. Frazer, _Pausanias_, i. 37, 6 note). Pop. (1907) 71,235. Its extreme length is 31 m., and its breadth varies from about 20 m. in the southern portion to 3 m. or less in the projecting part, which runs parallel with the island of Ithaca, at a distance of about 4 m. across the strait of Guiscardo or Viscaro. The whole island, with its area of 348 English sq. m., is covered with rocky hills of varying elevation, the main range running from north-west to south-east. The ancient Mount Aenos, now Elato, Monte Negro, or the Black Mountain (5315 ft.), frequently retains the snow for several months. It is not only the loftiest part of the sierra, but also the highest land in the whole Ionian group. The name "Black" was given from the darkness of the pine woods which still constitute the most striking feature in Cephalonian scenery, although their extent has been greatly curtailed by fire. The summit is called Megálo Sorós. The island is ill supplied with fresh water; there are few permanent streams except the Rakli, and springs are apt to fail in dry summers. In the western part of the island a gulf runs up from the south, a distance of about 7 m.; on its east side stands the chief town Argostoli, with about 10,000 inhabitants, and on its west side the rival city of Lixouri, with 6000. About a mile west of the town are the curious sea mills; a stream of sea water running down a chasm in the shore is made to turn the wheels. About 5 m. from Argostoli is the castle of St George, a building of Venetian origin, and the strongest fortification in the island. On an eminence east-south-east of Argostoli are the ruins of the ancient Cranii, and Lixouri is close to or upon those of Pale; while on the other side of the island are the remains of Samos on the bay of the same name, of Proni or Pronni, farther south above the vale of Rakli and its blossoming oleanders, and of an unknown city near the village of Scala. The ruins of this city include Roman baths, a brick-built temple, rock-cut tombs, and tessellated pavements; and Cranii, Proni and Samos are remarkable for stretches of Cyclopean and Hellenic walls, partly of the most irregular construction, and partly preserving almost unimpaired the results of the most perfect skill. The inhabitants of Cephalonia have all along been extremely active; and no slight amount of toil has been expended in the construction of terraces on the steep sides of the hills. Owing to the thinness of the population, however, but a small proportion of the soil is under cultivation, and the quantity of grain grown in the island is comparatively meagre. The staple is the currant, in the production of which the island surpasses Zante. The fruit is smaller than that of the Morea, and has a peculiar flavour; it finds a market mainly in Holland, Belgium and Germany. The grape vine also is grown, and the manufacture of wine is a rising industry. The olive crop is of considerable importance, and the culture of cotton in the low grounds has been successfully attempted. Manufactures are few and undeveloped, but lace from the aloe fibre, Turkey carpets and basket-work are produced by the villagers, and boats are built at both the principal towns. Of all the seven Ionian islands Cephalonia and Zante are most purely Greek, and the inhabitants display great mental activity.
In the Homeric poems Cephalonia is generally supposed to be mentioned under the name of Same, and its inhabitants, among the subjects of Ulysses, to be designated Cephallenes (see, however, under ITHACA). In the Persian War they took but little part; in the Peloponnesian they sided with the Athenians. The town of Pale was vainly besieged by Philip of Macedon in 218 B.C., because it had supported the Aetolian cause. In 189 B.C. all the cities surrendered to the Romans, but Same afterwards revolted, and was only reduced after a siege of four months. The island was presented by Hadrian to Athens, but it appears again at a later date as "free and autonomous." After the division of the Roman empire, it continued attached to Byzantium till 1082, when it was captured by Robert Guiscard, who died, however, before he could repress the revolt of 1085. In 1204 it was assigned to Gaius, prince of Tarentum, who accepted the protection of Venice in 1215; and after 1225 it was held along with Santa Maura and Zante by a succession of five counts of the Tocco family at Naples. Formally made over to Venice in 1350 by the prince of Tarentum, it was afterwards captured by the Turks in 1479; but the Hispanico-Venetian fleet under Benedetto Pessaro and Gonsalvo of Cordova effected their expulsion in 1500, and the island continued in Venetian possession till the fall of the republic. For some time it was administered for the French government, but in 1809 it was taken by the British under Cuthbert, Lord Collingwood. Till 1813 it was in the hands of Major de Bosset, a Swiss in the British service, who displayed an industry and energy in the repression of injustice and development of civilization only outdone by the despotic vigour of Sir Charles Napier, who held the same office for the nine years from 1818 to 1827. During the British protectorate the island made undoubted advances in material prosperity, but was several times the scene of political disturbances. It retained longer than the sister islands traces of feudal influence exerted by the landed proprietors, but has been gradually becoming more democratic. Under the Venetians it was divided into eight districts, and an elaborate system of police was in force; since its annexation to Greece it has been broken up into twenty demarchies, each with its separate jurisdiction and revenues, and the police system has been abolished.
AUTHORITIES.--A special treatise on the antiquities of Cephalonia was
written by Petrus Maurocenus. See Holland's _Travels_ (1815); Ansted's
_Ionian Islands_ (1863); Viscount Kirkwall's _Four Years in Ionian
Islands_ (1864); Wiebel's _Die Insel Kephalonia_; parliamentary
papers. Riemann, _Recherches archéologiques sur les Iles Ioniennes_
(Paris, 1879-1880); Partsch, _Kephallenia und Ithaka_ (1890); see also
CORFU; IONIAN ISLANDS. (E. Gr.)
CEPHALOPODA, the fifth of the classes into which the zoological phylum Mollusca is divided (see MOLLUSCA). The Cephalopoda are mainly characterized by the concrescence of the foot and head. The foot grows forward on each side so as to surround the mouth, the two upgrowths meeting on the dorsal side of the head--whence the name Cephalopoda. The perioral portion of the foot is drawn out into paired arm-like processes; these may be beset with sheathed tentacles or with suckers or hooks, or both. The epipodia are expanded into a pair of muscular lobes right and left, which are bent round towards one another so that their free margins meet and constitute a short tube--the siphon or funnel. The hind-foot is either very small or absent. A distinctive feature of the Cephalopoda is their bilateral symmetry and the absence of anything like the torsion of the visceral mass seen in the Anisopleurous Gastropoda.
The anus, although it may be a little displaced from the median line,
is approximately median and posterior. The mantle-skirt is deeply
produced posteriorly, forming a large sub-pallial chamber around the
anus. By the side of the anus are placed the single or paired
apertures of the nephridia, the genital apertures (paired only in
_Nautilus_, in female Octopoda, female _Ommatostrephes_ and male
_Eledone_), and the paired ctenidia. The visceral hump or dome is
elevated, and may be very much elongated in a direction almost at
right angles to the primary horizontal axis of the foot.
A shell is frequently, but not invariably, secreted on the visceral
hump and mantle-skirt. The shell is usually light in substance or
lightened by air-chambers in correlation with the free-swimming habits
of the Cephalopoda. It may be external or internal, that is, enclosed
in folds of the mantle. Very numerous minute pigmented sacs, capable
of expansion and contraction, and known as chromatophores, are usually
present in the integument. The sexes are separate.
The ctenidia are well developed as paired gill-plumes, serving as the
efficient branchial organs (figs. 4, 24),
The vascular system is very highly developed; the heart consists of a
pair of auricles and a ventricle (figs. 12, 28). Branchial hearts are
formed on the afferent vessels of the branchiae. It is not known to
what extent the minute subdivision of the arteries extends, or whether
there is a true capillary system.
The pericardium is extended so as to form a very large sac, passing
among the viscera dorsalwards and sometimes containing the ovary or
testis--the viscero-pericardial sac--which opens to the exterior
either directly or through the renal organs. It has no connexion with
the vascular system. The renal organs are always paired sacs, the
walls of which invest the branchial afferent vessels (figs. 28, 29).
They open each by a pore into the viscero-pericardial sac, except in
_Nautilus_. The anal aperture is median and raised on a papilla. Jaws
(fig. 6, e) and a radula (fig. 9) are well developed. The jaws have
the form of powerful beaks, either horny or calcified (_Nautilus_),
and are capable of inflicting severe wounds.
Cerebral, pleural and pedal ganglia are present, but the connectives
are shortened and the ganglia concentrated and fused in the cephalic
region. Large special ganglia (optic, stellate and supra-buccal) are
developed. Sense-organs are highly developed; the eye exhibits a very
special elaboration of structure in the Dibranchiata, and a remarkable
archaic form in the nautilus. Otocysts are present in all. The typical
osphradium is not present, except in _Nautilus_, but other organs are
present in the cephalic region, to which an olfactory function is
ascribed both in _Nautilus_ and in the other Cephalopoda.
Hermaphroditism is unknown in Cephalopoda; male and female individuals
always being differentiated. The genital aperture and duct is
sometimes single, when it is the left; sometimes the typical pair is
developed right and left of the anus. The males of nearly all
Cephalopoda have been shown to be characterized by a peculiar
modification of the arm-like processes or lobes of the fore-foot,
connected with the copulative function. The term hectocotylization is
applied to this modification (see figs. 6, 24). Elaborate
spermatophores or sperm-ropes are formed by all Cephalopoda, and very
usually the female possesses special capsule-forming and nidamental
glands for providing envelopes to the eggs (fig. 4, g.n.). The egg is
large, and the development is much modified by the presence of an
excessive amount of food-material diffused in the protoplasm of the
egg-cell. Trochosphere and veliger stages of development are
consequently not recognizable.
The Cephalopoda are divisible into two orders, Tetrabranchiata and Dibranchiata, the names of which (due to Sir R. Owen) describe the number of gill-plumes present; but in fact there are several characters, of as great importance as those derived from the gills, by which the members of these two orders are separated from one another.
ORDER 1. TETRABRANCHIATA (= Schizosiphona, Tentaculifera).
_Characters_.--The inrolled lateral margins of the epipodia are not fused, but form a siphon by apposition (fig. 4). The circumoral lobes of the fore-foot carry numerous retractile tentacles, not suckers (fig. 6). There are two pairs of ctenidial gills (hence Tetrabranchiata), and two pairs of renal organs, consequently four renal apertures (fig. 4). The viscero-pericardial chamber opens by two independent apertures to the exterior, and not into the renal sacs. There are two oviducts (right and left) in the female, and two sperm-ducts in the male, the left duct in both sexes being rudimentary. A large external shell, either coiled or straight, is present, and is not enclosed by reflections of the mantle-skirt. The shell consists of a series of chambers, the last-formed of which is occupied by the body of the animal, the hinder ones (successively deserted) containing gas (fig. 1). The pair of cephalic eyes are hollow chambers (fig. 14. A), opening to the exterior by minute orifices (pin-hole camera), and devoid of refractive structures. A pair of osphradia are present at the base of the gills (fig. 4, _olf_). Salivary glands are wanting. An ink-sac is _not_ present. Branchial hearts are _not_ developed on the branchial afferent vessels.
a, Visceral hump.
b, Portion of the free edge of the mantle-skirt reflected on to the
shell,--the edge of the mantle-skirt can be traced downwards and
forwards around the base of the mid-foot or siphon i.
l, l, Superficial origin of the retractor muscle of the mid-foot
(siphon), more or less firmly attached to the shell, of which a small
piece (s) is seen between the letters l, l.
s, (farther back) points to the siphuncular pedicle, which is broken
off short and not continued, as in the perfect state, through the
whole length of the siphuncle of the shell, also marked s and s'.
o, points to the right eye.
t, is placed near the extremities of the contracted tentacles of the
outer or annular lobe of the fore-foot--the jointed tentacles are seen
protruding a little from their long cylindrical sheaths.
v, The dorsal "hood" formed by an enlargement in this region of the
annular lobe of the fore-foot (m in figs. 2, 3).
V, A swelling of the mantle-skirt, indicating the position on its
inner face of the nidamental gland (see fig. 4, g.n.).]
_Visceral Hump and Shell._--The visceral hump of _Nautilus_ (if we exclude from consideration the fine siphuncular pedicle which it trails, as it were, behind it) is very little, if at all, affected by the coiled form of the shell which it carries, since the animal always slips forward in the shell as it grows, and inhabits a chamber which is practically cylindrical (fig. 1). Were the deserted chambers thrown off instead of being accumulated behind the inhabited chamber as a coiled series of air-chambers, we should have a more correct indication in the shell of the extent and form of the animal's body. Amongst Gastropods it is not very unusual to find the animal slipping forward in its shell as growth advances and leaving an unoccupied chamber in the apex of the shell. This may indeed become shut off from the occupied cavity by a transverse septum, and a series of such septa may be formed, but in no Gastropod are these apical chambers known to contain a gas during the life of the animal in whose shell they occur. A further peculiarity of the nautilus shell and of that of the allied extinct _Ammonites_, _Scaphites_, _Orthoceras_, &c., and of the living _Spirula_, is that the series of deserted air-chambers is traversed by a cord-like pedicle extending from the centro-dorsal area of the visceral hump to the smallest and first-formed chamber of the series. No structure comparable to this siphuncular pedicle is known in any other Mollusca. The siphuncle does not communicate with the coelomic cavity; it is a simple vascular process of the mantle, whose cavity consists of a venous sinus, and whose wall contains a ramification of the pallial artery. There appears to be no doubt that the deserted chambers of the nautilus shell contain in the healthy living animal a gas which serves to lessen the specific gravity of the whole organism. This gas is said to be of the same composition as the atmosphere, with a larger proportion of nitrogen. With regard to its origin we have only conjectures. Each septum shutting off an air-containing chamber is formed during a period of quiescence, probably after the reproductive act, when the visceral mass of the nautilus may be slightly shrunk, and gas is secreted from the dorsal integument so as to fill up the space previously occupied by the animal. A certain stage is reached in the growth of the animal when no new chambers are formed. The whole process of the loosening of the animal in its chamber and of its slipping forward when a new septum is formed, as well as the mode in which the air-chambers may be used as a hydrostatic apparatus, and the relation to this use, if any, of the siphuncular pedicle, is involved in obscurity, and is the subject of much ingenious speculation. In connexion with the secretion of gas by the animal, besides the parallel cases ranging from the protozoon _Arcella_ to the physoclistic fishes, from the hydroid _Siphonophora_ to the insect-larva _Corethra_, we have the identical phenomenon observed in the closely allied _Sepia_ when recently hatched. Here, in the pores of the internal rudimentary shell, gas is observable, which has necessarily been liberated by the tissues which secrete the shell, and not derived from any external source (Huxley).
The coiled shell of _Nautilus_, and of the majority of extinct Tetrabranchiata, is peculiar in its relation to the body of the animal, inasmuch as the curvature of the coil proceeding from the centro-dorsal area is towards the head or forwards, instead of away from the head and backwards as in other discoid coiled shells such as _Planorbis_; the coil is in fact absolutely reversed in the two cases. Such a shell is said to be exogastric. But in some extinct forms, e.g. _Phragmoceras_, _Cyrtoceras_, _Ptenoceras_, the shell is coiled towards the ventral side, when it is termed endogastric. Amongst the extinct allies of the nautilus (Tetrabranchiata) we find shells of a variety of shapes, open coils such as _Scaphites_, leading on to perfectly cylindrical shells with chamber succeeding chamber in a straight line (_Orthoceras_), whence again we may pass to the corkscrew spires formed by the shell of _Turrilites_. In some extinct genera, e.g. _Gomphoceras_, among the Nautiloidea the aperture of the shell is contracted and the edge of the aperture is lobed. In these cases the animal was probably able only to protrude its appendages and not its whole head. The ventral part of the aperture corresponding to the funnel is separated from the dorsal part by a constriction. Hence it is possible to distinguish the ventral and dorsal sides of the shell and to decide whether it was exogastric or endogastric. The direction of the coil of the shell cannot be determined by the position of the siphuncle, which traverses the septa centrally, ventrally or dorsally. Contracted shell apertures occur also in Ammonitoidea, the condition reaching an extreme in _Morphoceras_, where the original aperture is subdivided by the ingrowth of the sides, so that only five small separate apertures remain. Of these the central probably corresponded to the mouth, two lateral to the eyes, and the remaining two to the pedal appendages.
_Head, Foot, Mantle-skirt and Sub-pallial Chamber._--In the pearly
nautilus the ovoid visceral hump is completely encircled by the free
flap of integument known as mantle-skirt (figs. 2, 3, d, e). In the
antero-dorsal region this flap is enlarged so as to be reflected a
little over the coil of the shell which rests on it. In the
postero-ventral region the flap is deepest, forming an extensive
sub-pallial chamber, at the entrance of which e is placed in fig. 3. A
view of the interior of the sub-pallial chamber, as seen when the
mantle-skirt is retroverted and the observer faces in the direction
indicated by the reference line passing from e in fig. 3, is given in
fig. 4. With this should be compared the similar view of the
sub-pallial chamber of the Dibranchiate _Sepia_. It should be noted as
a difference between _Nautilus_ and the Dibranchiates that in the
former the nidamental gland (in the female) lies on that surface of
the pallial chamber formed by the dependent mantle-flap (fig. 4, g.n.;
fig. 1, V), whilst in the latter it lies on the surface formed by the
body-wall; in fact in the former the base of the fold forming the
mantle-skirt comprises in its area a part of what is unreflected
visceral hump in the latter.
m, The dorsal "hood" formed by the enlargement of the outer or
annular lobe of the fore-foot, and corresponding to the sheaths of
two tentacles (g, g in fig. 6).
n, Tentacular sheaths of lateral portion of the annular lobe.
u, The left eye.
b, The nuchal plate, continuous at its right and left posterior
angles with the root of the mid-foot, and corresponding to the
nuchal cartilage of Sepia.
c, Visceral hump.
d, The free margin of the mantle-skirt, the middle letter d points
to that portion of the mantle-skirt which is reflected over a part
of the shell as seen in fig. 1, b; the cup-like fossa to which b and
d point in the present figure is occupied by the coil of the shell.
g.a. points to the lateral continuation of the nuchal plate b to
join the root of the mid-foot or siphon.]
e, points to the concave margin of the mantle-skirt leading into the
sub-pallial chamber.
g, The mid-foot or siphon.
k, The superficial origin of its retractor muscles closely applied
to the shell and serving to hold the animal in its place.
l, The siphuncular pedicle of the visceral hump broken off short.
v, v, The superior and inferior ophthalmic tentacles.]
The apertures of the two pairs of renal sacs, of the
viscero-pericardial sac, of the genital ducts, and of the anus, are
shown in position on the body-wall of the pallial chamber of
_Nautilus_ in figs. 4, 5. There are nine apertures in all, one median
(the anus) and four paired. Besides these apertures we notice _two_
pairs of gill-plumes which are undoubtedly typical ctenidia, and a
short papilla (the osphradium) between each anterior and posterior
gill-plume (see figs. 4, 5, and explanation). As compared with this in
a Dibranchiate, we find (fig. 25) only four apertures, viz. the median
anus with adjacent orifice of the ink-sac, the single pair of renal
apertures, and one asymmetrical genital aperture (on the left side)
except in female Octopoda and a few others, where the genital ducts
and their apertures are paired. No viscero-pericardial pores are
present on the surface of the pallial chamber, since in the
Dibranchiata the viscero-pericardial sac opens by a pore into each
nephridium instead of directly to the surface. A single pair of
ctenidia (gill-plumes) is present instead of the two pairs in
_Nautilus_. The existence of two pairs of ctenidia and of two pairs of
renal sacs in _Nautilus_, placed one behind the other, is highly
remarkable. The interest of this arrangement is in relation to the
general morphology of the Mollusca, for it is impossible to view this
repetition of organs in a linear series as anything else than an
instance of metameric segmentation, comparable to the segmentation of
the ringed worms and Arthropods. The only other example which we have
of this metamerism in the Mollusca is presented by the Chitons. There
we find not two pairs of ctenidia merely, but sixteen pairs (in some
species more) accompanied by a similar metamerism of the dorsal
integument, which carries eight shells. In _Chiton_ the renal organs
are not affected by the metamerism as they are in _Nautilus_. It is
impossible on the present occasion to discuss in the way which their
importance demands the significance of these two instances among
Mollusca of incomplete or partial metamerism; but it would be wrong to
pass them by without insisting upon the great importance which the
occurrence of these isolated instances of metameric segmentation in a
group of otherwise unsegmented organisms possesses, and the light
which they may be made to throw upon the nature of metameric
segmentation in general.
a, Muscular band passing from the mid-foot to the integument.
b, The valve on the surface of the funnel, partially concealed by
the inrolled lateral margin of the latter.
c, The mantle-skirt retroverted.
an, The median anus.
x, Post-anal papilla of unknown significance.
g.n., Nidamental gland.
r.ov, Aperture of the right oviduct.
l.ov, Aperture of the rudimentary left oviduct (pyriform sac of
Owen).
neph.a, Aperture of the left anterior renal sac.
neph.p, Aperture of the left posterior renal sac.
viscper, Left aperture of the viscero-pericardial sac.
olf, The left osphradium placed near the base of the anterior
gill-plume.
The four gill-plumes (ctenidia) are not lettered.]
The foot and head of _Nautilus_ are in the adult inextricably grown
together, the eye being the only part belonging primarily to the head
which projects from the all-embracing foot. The fore-foot or front
portion of the foot has the form of a number of lobes carrying
tentacles and completely surrounding the mouth (figs. 2, 3). The
epipodia incline towards each other posteriorly so as to form an
incomplete siphon (fig. 4), a condition which is completed and
rendered permanent in the tubular funnel of Dibranchiata. The
epipodial nature of the funnel is well seen in young embryos, in which
this organ is situated laterally and posteriorly between the mantle
and the foot.
The lobes of the fore-foot of _Nautilus_ and of the other Cephalopoda
require further description. It has been doubted whether these lobes
were rightly referred (by T.H. Huxley) to the fore-foot, and it has
been maintained by some zoologists (H. Grenadier, H. von Jhering)
that they are truly processes of the head. It appears to be impossible
to doubt that the lobes in question are the fore-portion of the foot,
when their development is examined (see fig. 35), further, when the
fact is considered that they are innervated by the pedal ganglion. The
fore-foot of _Nautilus_ completely surrounds the buccal cone (fig. 6,
e), so as to present an appearance with its expanded tentacles similar
to that of the disk of a sea-anemone (_Actinia_). A.G. Bourne, of
University College, prepared from actual specimens the drawings of
this part in the male and female _Nautilus_ reproduced in fig. 6, and
restored the parts to their natural form when expanded. The drawings
show very strikingly the difference between male and female. In the
females (lower figure), we observe in the centre of the disk the
buccal cone e carrying the beak-like pair of jaws which project from
the finely papillate buccal membrane. Three tentaculiferous lobes of
the fore-foot are in immediate contact with this buccal cone; they are
the right and left (c, c) inner lobes, and the inferior inner lobe
(d)--called inferior because it really lies ventralwards of the mouth.
This inner inferior lobe is clearly a double one, representing a right
and left inner inferior lobe fused into one. A lamellated organ on its
surface, known as Owen's organ, probably olfactory in function (n),
marks the separation of the constituent halves of this double lobe.
Each half carries a group of fourteen tentacles. The right and the
left inner lobes (c, c) each carry twelve tentacles. External to these
three lobes the muscular substance of the mouth-embracing foot is
raised into a wide ring, which becomes especially thick and large in
the dorsal region where it is notably modified in form, offering a
concavity into which the coil of the shell is received, and furnishing
a protective roof to the retracted mass of tentacles. This part of the
external annular lobe of the fore-foot is called the "hood" (figs. 2,
3, m). The median antero-posterior line traversing this hood exactly
corresponds to the line of concrescence of the two halves of the
fore-foot, which primitively grew forward one on each side of the
head, and finally fused together along this line in front of the
mouth. The tentacles carried by the great annular lobe are nineteen on
each side, thirty-eight in all. They are called "digital," and are
somewhat larger than the "labial" tentacles carried on the three inner
lobes. The dorsalmost pair of tentacles (marked g in fig. 6) are the
only ones which actually belong to that part of the disk which forms
the great dorsal hood m. The hood is, in fact, to a large extent
formed by the enlarged sheaths of these two tentacles. All the
tentacles of the circumoral disk are set in remarkable tubular
sheaths, into which they can be drawn. The sheaths of some of those
belonging to the external or annular lobe are seen in fig. 3, marked
n. The sheaths are muscular as well as the tentacles, and are simply
tubes from the base of which the solid tentacle grows. The functional
significance of this sheathing arrangement is as obscure as its
morphological origin. With reference to the latter, it appears highly
probable that the tubular sheath represents the cup of a sucker such
as is found on the fore-foot of the Dibranchiata. In any case, it
seems to the writer impossible to doubt that each tentacle, and its
sheath on a lobe of the circumoral disk of Nautilus, corresponds to a
sucker on such a lobe of a Dibranchiate. W. Keferstein follows Sir R.
Owen in strongly opposing this identification, and in regarding such
tentacle as the equivalent of a whole lobe or arm of a Decapod or
Octopod Dibranch. The details of these structures, especially in the
facts concerning the hectocotylus and spadix, afford the most
conclusive reasons for dissenting from Owen's view. On the ventral
side an extensive part of the internal surface of the muscular ring is
laminated, forming the so-called "organ of Valenciennes," peculiar to
the female and serving for the attachment of the spermatophores. We
have so far enumerated in the female nautilus ninety tentacles. Four
more remain which have a very peculiar position, and almost lead to
the suggestion that the eye itself is a modified tentacle. These
remaining tentacles are placed one above (before) and one below
(behind) each eye, and bring up the total to ninety-four (fig. 3 v,
v).
a, The shell.
b, The _outer_ ring-like expansion (annular lobe) of the circumoral
muscular mass of the fore-foot, carrying nineteen tentacles on each
side--posteriorly this is enlarged to form the "hood" (marked v in
fig. 1 and m in figs. 2 and 3). giving off the pair of tentacles
marked g in the present figure.
c, The right and left inner lobes of the fore-foot, each carrying
twelve tentacles in the female, in the male subdivided into p, the
"spadix" or hectocotylus on the left side, and q, the "anti-spadix,"
a group of four tentacles on the right side--it is thus seen that
the subdivided right and left inner lobes of the male correspond to
the undivided right and left inner lobes of the female.
d, The inner inferior lobe of the fore-foot, a bilateral structure
in the female carrying two groups, each of fourteen tentacles,
separated from one another by a lamellated organ n, supposed to be
olfactory in function--in the male the inner inferior lobe of the
fore-foot is very much reduced, and has the form of a paired group
of lamellae (d in the upper figure).
e, The buccal cone, rising from the centre of the three inner lobes,
and fringing the protruded calcareous beaks or jaws with a series of
minute papillae.
f, The tentacles of the outer circumoral lobe or annular lobe of the
fore-foot projecting from their sheaths.
g, The two most posterior tentacles of this series belonging to that
part of the annular lobe which forms the hood (m in figs. 2 and 3).
i, Superior ophthalmic tentacle.
k, Inferior ophthalmic tentacle.
l, Eye.
m, Paired laminated organ on each side of the base of the inner
inferior lobe (d) of the female.
n, Olfactory lamellae upon the inner inferior lobe (in the female).
o, The siphon (mid-foot).
p, The spadix (in the male), the hectocotylized portion of the left
inner lobe of the fore-foot representing four modified tentacles,
eight being left unmodified.
q, The anti-spadix (in the male), being four of the twelve tentacles
of the right inner lobe of the fore-foot isolated from the remaining
eight, and representing on the right side the differentiated spadix
of the left side. The four tentacles of the anti-spadix are set,
three on one base and one on a separate base.]
In the adult male nautilus we find the following important differences
in the tentaculiferous disk as compared with the female (see upper
drawing in fig. 6). The inner inferior lobe is rudimentary, and
carries no tentacles. It is represented by three groups of lamellae
(d), which are not fully exposed in the drawing. The right and left
inner lobes are subdivided each into two portions. The right shows a
larger portion carrying eight tentacles, and smaller detached groups
(q) of four tentacles, of which three have their sheaths united whilst
one stands alone. These four tentacles may be called the
"anti-spadix." The left inner lobe shows a similar larger portion
carrying eight tentacles, and a curious conical body behind it
corresponding to the anti-spadix. This is the "spadix." It carries no
tentacles, but is terminated by imbricated lamellae. These lamellae
appear to represent the four tentacles of the anti-spadix of the right
internal lobe, and are generally regarded as corresponding to that
modification of the sucker-bearing arms of male Dibranchiate
Siphonopods to which the name "hectocotylus" is applied. The spadix is
in fact the hectocotylized portion of the fore-foot of the male
nautilus. The hectocotylized arm or lobe of male Dibranchiata is
connected with the process of copulation, and in the male nautilus the
spadix has probably a similar significance, though it is not possible
to suggest how it acts in this relation. It is important to observe
that the modification of the fore-foot in the male as compared with
the female nautilus is not confined to the existence of the spadix.
The anti-spadix and the reduction of the inner inferior lobe are also
male peculiarities. The external annular lobe in the male does not
differ from that of the female; it carries nineteen tentacles on each
side. The four ophthalmic tentacles are also present. Thus in the male
nautilus we find altogether sixty-two tentacles, the thirty-two
additional tentacles of the female being represented by lamelliform
structures.
_Musculature, Fins and, Cartilaginous Skeleton._--Without entering
into a detailed account of the musculature of _Nautilus_, we may point
out that the great muscular masses of the fore-foot and of the
mid-foot (siphon) are ultimately traceable to a large transverse mass
of muscular tissue, the ends of which are visible through the
integument on the right and left surfaces of the body dorsal of the
free flap of the mantle-skirt (fig. 1, l, l, and fig. 3, k). These
muscular areae have a certain adhesion to the shell, and serve both to
hold the animal in its shell and as the fixed supports for the various
movements of the tentaculiferous lobes and the siphon. They are to be
identified with the ring-like area of adhesion by which the
foot-muscle of the limpet is attached to the shell of that animal. In
the Dibranchs a similar origin of the muscular masses of the fore-foot
and mid-foot from the sides of the shell--modified, as this is, in
position and relations--can be traced.
a, Simple cells.
b, Dividing cells.
c, Canaliculi.
d, An empty cartilage capsule with its pores.
e, Canaliculi in section.]
In _Nautilus_ there are no fin-like expansions of the integument,
whereas such occur in the Decapod Dibranchs along the sides of the
visceral hump (figs. 15, 16). As an exception among Octopoda lateral
fins occur in _Pinnoctopus_ (fig. 38, A), and in _Cirrhoteuthis_ (fig.
38, D).
In _Nautilus_ there is a curious plate-like expansion of integument in
the mid-dorsal region just behind the hood, lying between that
structure and the portion of mantle-skirt which is reflected over the
shell. This is shown in fig. 2, b. If we trace out the margin of this
plate we find that it becomes continuous on each side with the sides
of the funnel. In _Sepia_ and other Decapods (not in Octopods) a
closely similar plate exists in an exactly corresponding position (see
b in figs. 10, 26). In _Sepia_ a cartilaginous development occurs here
immediately below the integument forming the so-called "nuchal plate,"
drawn in fig. 8, D. The morphological significance of this nuchal
lamella, as seen both in _Nautilus_ and in _Sepia_, is not obvious.
Cartilage having the structure shown in fig. 7 occurs in various
regions of the body of Cephalopoda. In all Glossophorous Mollusca the
lingual apparatus is supported by internal skeletal pieces, having the
character of cartilage; but in the Cephalopoda such cartilage has a
wider range.
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Encyclopaedia Britannica, 11th Edition, "Celtes, Konrad" to "Ceramics"Chapter VI: Part 6
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