Plate I.Fig. 1.—SECTION THROUGH RABBIT’S LIVER, INFECTED WITHCOCCIDIUM CUNICULI. (AFTER THOMA.)Fig. 2.—KLOSSIA HELICINA, FROM KIDNEY OF HELIX HORTENSIS.a, Portion of a section of the kidney showing normal epithelial cells containing concretions (c), and enlarged epithelial cells containing the parasite (k) in various stages; b, cyst of theKlossiacontaining sporoblasts; c, cyst with ripe spores, each enclosing four sporozoites and a patch of residual protoplasm. (From Wasielewski, after Balbiani.)Fig. 3.—THE LIFE-CYCLE OFCOCCIDIUM SCHUBERGI, SCHAUD. (PAR.LITHOBIUS FORFICATUS). (FROM MINCHIN, AFTER SCHAUDINN.)I.-IV represents the schizogony, commencing with infection of an epithelial cell by a sporozoite or merozoite. After stage IV the development may start again at stage I, as indicated by the arrows; or it may go on to the formation of gametocytes (V). V-VIII represents the sexual generation. The line of development, hitherto single (I-IV) becomes split into two lines—male (VI ♂, VII ♂, VIII ♂), and female (VI ♀, VII ♀, VIII ♀), culminating in the highly differentiated micro- and mega-gametes. By conjugation these two lines are again united. IX, X, show the formation of the zygote by fusion of the nuclei of the gametes. XI-XV, sporogony. H.C, host-cell; N, its nucleus; mz, merozoite; szt, schizont; ky, karyosome (or fragments of same); n.n, daughter-nuclei of schizont; pl.gr, plastinoid grains; ooc, oocyst; n.zyg, zygote-nucleus (segmentation-nucleus); sp.m, spore-membrane (sporocyst); rp, residual protoplasm of oocyst (“reliquat kystal”); rp.sp, residual protoplasm of spore (“reliquat sporal”); sp.z, sporozoite.Fig. 4.—PHASES OFCARYOTROPHA MESNILII, SIEDL. (PAR.POLYMNIA NEBULOSA).a, Young schizont in a cluster of spermatogonia; the host-cell (represented granulated) and two of its neighbours are greatly hypertrophied, with very large nuclei, and have fused into a single mass containing the parasite (represented clear, with a thick outline). The other spermatogonia are normal. b, Intracellular schizont divided up into schizontocytes (c), each schizontocyte giving rise to a cluster of merozoites arranged as a “corps en barillet”; spg, spermatogonia; h.c, host-cell; N, nucleus of host-cell or cells; n, nucleus of parasite; szc, schizontocyte; mz, merozoites; r.b, residual bodies of the schizontocytes. (From Minchin, after Siedlecki.)Plate II.Fig. 5.—SCHIZOGONY OFADELEA OVATA, A. SCHN. (PAR.LITHOBIUS FORFICATUS).a-c, ♀ generation; d-f, ♂ generation. a, Full-grown ♀ schizont (megaschizont), with a large nucleus (n) containing a conspicuous karyosome (ky). b, Commencement of schizogony; the nucleus has divided up to form a number of daughter-nuclei (d.n). The karyosome of stage a has broken up into a great number of daughter-karyosomes, each of which forms at first the centre of one of the star-shaped daughter-nuclei; but in a short time the daughter-karyosomes become inconspicuous. c, Completion of schizogony; the ♀ schizont has broken up into a number ofmegamerozoites(♀ mz) implanted on a small quantity of residual protoplasm (r.p.). Each ♀ merozoite has a chromatic nucleus (n) without a karyosome. d, Full-grown ♂ schizont (microschizont), with nucleus (n), karyosome (ky), and a number of characteristic pigment-granules (p.gr). e, Commencement of schizogony. The nucleus is dividing up into a number of daughter-nuclei (d.n), each with a conspicuous karyosome (ky). f, Completion of schizogony. The numerous micro-merozoites (♂ mz) have each a nucleus with a conspicuous karyosome (ky) at one pole, and the protoplasm contains pigment-granules (p.gr) near the nucleus, on the side farthest from the karyosome. (From Minchin, after Siedlecki.)Fig. 6.—ASSOCIATION AND CONJUGATION INADELEA OVATA.a, Young microgametocyte (♂ gamc.) attached to a megagametocyte (♀ gamc.). The nucleus of the microgametocyte gives rise to 4 daughter-nuclei (c) which become (d) 4 microgametes (♀ gam.). e, One of the microgametes penetrates the megagamete, which forms a fertilization-spindle composed of male and female chromatin (♂ and ♀ chr.). The other 3 microgametes and the residual protoplasm of the microgametocyte (r.p.) perish. The karyosome of the megagamete has disappeared, as such. f, Union of the chromatin of both elements, to produce the zygote-nucleus (n.zyg.). (From Minchin, after Siedlecki.)Fig. 7.—SPORES OF VARIOUS COCCIDIAN GENERA.a,Minchinia chitonis(E.R.L.), (par. (Chiton); b,Diaspora hydatidea, Léger (par.Polydesmus); c,Echinospora labbei, Léger (par.Lithobius mutabilis); d,Goussia motellae, Labbé; e,Diplospora(Hyaloklossia),lieberkuhni(Labbé), (par.Rana esculenta); f,Crystallospora crystalloides(Thél.), (par.Motella tricirrata). (From Minchin; b and c after Léger, the others after Labbé.)Fig. 8.—SPOROGONY AND SPORE-GERMINATION INBARROUSSIA ORNATA, A. SCH., FROM THE GUT OFNEPA CINERA.a, Oocyst with sporoblasts; b, oocyst with ripe spores; c, a spore highly magnified, showing the single sporozoite bent on itself; d, the spore has split along its outer coat or epispore, but the sporozoite is still enclosed in the endospore; e, the sporozoite, freed from the endospore, is emerging; f, the sporozoite has straightened itself out and is freed from its envelopes. (From Wasielewski, after A. Schneider.)
Plate I.
a, Portion of a section of the kidney showing normal epithelial cells containing concretions (c), and enlarged epithelial cells containing the parasite (k) in various stages; b, cyst of theKlossiacontaining sporoblasts; c, cyst with ripe spores, each enclosing four sporozoites and a patch of residual protoplasm. (From Wasielewski, after Balbiani.)
I.-IV represents the schizogony, commencing with infection of an epithelial cell by a sporozoite or merozoite. After stage IV the development may start again at stage I, as indicated by the arrows; or it may go on to the formation of gametocytes (V). V-VIII represents the sexual generation. The line of development, hitherto single (I-IV) becomes split into two lines—male (VI ♂, VII ♂, VIII ♂), and female (VI ♀, VII ♀, VIII ♀), culminating in the highly differentiated micro- and mega-gametes. By conjugation these two lines are again united. IX, X, show the formation of the zygote by fusion of the nuclei of the gametes. XI-XV, sporogony. H.C, host-cell; N, its nucleus; mz, merozoite; szt, schizont; ky, karyosome (or fragments of same); n.n, daughter-nuclei of schizont; pl.gr, plastinoid grains; ooc, oocyst; n.zyg, zygote-nucleus (segmentation-nucleus); sp.m, spore-membrane (sporocyst); rp, residual protoplasm of oocyst (“reliquat kystal”); rp.sp, residual protoplasm of spore (“reliquat sporal”); sp.z, sporozoite.
a, Young schizont in a cluster of spermatogonia; the host-cell (represented granulated) and two of its neighbours are greatly hypertrophied, with very large nuclei, and have fused into a single mass containing the parasite (represented clear, with a thick outline). The other spermatogonia are normal. b, Intracellular schizont divided up into schizontocytes (c), each schizontocyte giving rise to a cluster of merozoites arranged as a “corps en barillet”; spg, spermatogonia; h.c, host-cell; N, nucleus of host-cell or cells; n, nucleus of parasite; szc, schizontocyte; mz, merozoites; r.b, residual bodies of the schizontocytes. (From Minchin, after Siedlecki.)
Plate II.
a-c, ♀ generation; d-f, ♂ generation. a, Full-grown ♀ schizont (megaschizont), with a large nucleus (n) containing a conspicuous karyosome (ky). b, Commencement of schizogony; the nucleus has divided up to form a number of daughter-nuclei (d.n). The karyosome of stage a has broken up into a great number of daughter-karyosomes, each of which forms at first the centre of one of the star-shaped daughter-nuclei; but in a short time the daughter-karyosomes become inconspicuous. c, Completion of schizogony; the ♀ schizont has broken up into a number ofmegamerozoites(♀ mz) implanted on a small quantity of residual protoplasm (r.p.). Each ♀ merozoite has a chromatic nucleus (n) without a karyosome. d, Full-grown ♂ schizont (microschizont), with nucleus (n), karyosome (ky), and a number of characteristic pigment-granules (p.gr). e, Commencement of schizogony. The nucleus is dividing up into a number of daughter-nuclei (d.n), each with a conspicuous karyosome (ky). f, Completion of schizogony. The numerous micro-merozoites (♂ mz) have each a nucleus with a conspicuous karyosome (ky) at one pole, and the protoplasm contains pigment-granules (p.gr) near the nucleus, on the side farthest from the karyosome. (From Minchin, after Siedlecki.)
a, Young microgametocyte (♂ gamc.) attached to a megagametocyte (♀ gamc.). The nucleus of the microgametocyte gives rise to 4 daughter-nuclei (c) which become (d) 4 microgametes (♀ gam.). e, One of the microgametes penetrates the megagamete, which forms a fertilization-spindle composed of male and female chromatin (♂ and ♀ chr.). The other 3 microgametes and the residual protoplasm of the microgametocyte (r.p.) perish. The karyosome of the megagamete has disappeared, as such. f, Union of the chromatin of both elements, to produce the zygote-nucleus (n.zyg.). (From Minchin, after Siedlecki.)
a,Minchinia chitonis(E.R.L.), (par. (Chiton); b,Diaspora hydatidea, Léger (par.Polydesmus); c,Echinospora labbei, Léger (par.Lithobius mutabilis); d,Goussia motellae, Labbé; e,Diplospora(Hyaloklossia),lieberkuhni(Labbé), (par.Rana esculenta); f,Crystallospora crystalloides(Thél.), (par.Motella tricirrata). (From Minchin; b and c after Léger, the others after Labbé.)
a, Oocyst with sporoblasts; b, oocyst with ripe spores; c, a spore highly magnified, showing the single sporozoite bent on itself; d, the spore has split along its outer coat or epispore, but the sporozoite is still enclosed in the endospore; e, the sporozoite, freed from the endospore, is emerging; f, the sporozoite has straightened itself out and is freed from its envelopes. (From Wasielewski, after A. Schneider.)
Comparing the life-cycle of other Coccidia with that just described, a greater or less degree of modification is frequently met with. In the process of schizogony two orders of division sometimes occur; the parent-schizont first divides up into a varying number of rounded daughter-schizonts (schizontocytes), each of which gives rise, in the usual manner, to a cluster of merozoites,3which thus constitute a second order of cells. Siedlecki (1902) has found this to be the case inCaryotropha mesnilii(fig. 4), and Woodcock (1904) has shown that it is most probably really the same process which Smith and Johnson (1902) mistook for sporogony when originally describing their Coccidian of the mouse,Klossiella. InCaryotropha, a perfectly similar state of affairs is seen in the formation of microgametes from the microgametocyte; this is additionally interesting as showing that this process is neither more nor less than male schizogony.
Coming to the sexual generation, considerable variation is met with as regards the period in the life-history when sexual differentiation first makes its appearance. Sexuality may become evident at the very beginning of schizogony, as,e.g.inAdelea ovata(Siedlecki, 1899), where the first-formed schizonts (those developed from the sporozoites) are differentiated into male and female (micro-and mega-schizonts) (see Plate II., fig. 5). Correspondingly, the merozoites, to which they give rise, are also different (micro-and mega-merozoites). In one or two cases sexuality appears even earlier in the cycle, and has thus been carried still farther back.
The Coccidia, as a whole, have not developed the phenomenon of association of the sexual individuals prior to gamete-formation which is so characteristic of Gregarines. Their method of endeavouring to secure successful sporulation, and thus the survival of the species, has been rather by the extreme specialization of the sexual process. In place of many female elements, which the primitive or ancestral forms may be assumed to have had,4there is always, save possibly for one exception,5only a single relatively huge megagamete formed, which offers a comparatively easy goal for one of the many microgametes. Nevertheless in the effort to render fertilization absolutely certain, a few Coccidia have acquired (secondarily) the power of associating; a state of things which enables those forms, moreover, to effect an economy in the number of male gametes, only three or four being developed. Instances are seen inAdelea mesnili(Perez, 1903),A. ovata(fig. 6), andKlossia helicina(Siedlecki, 1899). It is very interesting to note that, in the two last cases, unless this association of the microgametocyte with the megagametocyte occurs, neither can the former produce male elements (microgametes) nor can the female individual maturate and become ready for fertilization. (Concerning this question of association see alsoGregarines.)
In sporogony, great variation is seen with respect to the number of spores and sporozoites formed; and, as in Gregarines, these characters are largely used for purposes of classification, under which heading they are better considered. Usually, the spores (fig. 7) are quite simple in outline, and not produced into spines or processes; exceptions are found, however, in a few instances (e.g.Minchinia chitonis). In one case (Coccidium mitrarium), the oocyst itself, instead of being spherical, is curiously shaped like a mitre.
The life-history as a whole is invariably undergone in a single host,i.e.there is no alternation of true hosts.6Schaudinn, in his work on theCoccidiaofLithobius(1900), showed that the oocysts expelled with the faeces may be eaten by wood-lice (Oniscus), but when this happens they pass through the intestine of the wood-louse unaltered, the latter not being an intermediate host but merely a carrier.
The order Coccidiidea is divided into four families, characterized by the number of sporocysts (if any) found in the oocyst.Classification.Fam.Asporocystidae, Léger. No sporozoites are formed in the oocyst, the sporozoites being unenclosed (gymnospores).Genus,Légerella, Mesnil. This genus actually conforms to Aimé Schneider’s original definition ofEimeria, which was founded on what were really the schizogonous generations of other forms, then thought to be distinct. In view of the great confusion attending the use of this name, however, Mesnil (1900) has suggested the new one here adopted. Two species known,L. novaandL. testiculi, both from different species ofGlomeris, a Myriapod; the former inhabits the Malpighian tubules, the latter the testis.Fam.Disporocystidae, Léger. The oocyst contains 2 spores.Genus 1.Cyclospora, A. Schneider. Spores dizoic,i.e.with two sporozoites.C. glomericola, from the intestinal epithelium ofGlomeris, andC. caryolytica, from the intestinal epithelium of the mole, intranuclear.Genus 2.Diplospora, Labbé. Spores tetrazoic.D. lacazei, from many birds, is the best-known species; and others have been described from different Sauropsida.D. lieberkühniis an interesting form occurring in the kidneys of the frog, which it reaches by way of the circulation.Genus 3.Isospora, Schn. Spores polyzoic. Founded forI. rara, parasitic in the black slug (Limax cinereo-niger). Many authors consider that Schneider was mistaken in attributing many sporozoites to this form, and would unite with it the genusDiplospora.Fam.Tetrasporocystidae, Léger. The oocyst contains 4 spores.Genus 1.Coccidium,7Leuckart. The spores are dizoic and the sporocysts rounded or oval. A very large number of species are known, mostly from Vertebrate hosts.C. cuniculi(=C. oviforme) from the rabbit (intestine and diverticula), but also occurring sometimes in other domestic animals; C. falciformis, from the mouse;C. faureifrom sheep; andC. schubergi, fromLithobius(a centipede), are among the best-known forms. All of them may cause disastrous epidemics of coccidiosis.Genus 2.Paracoccidium, Laveran and Mesnil. This genus is distinguished fromCoccidiumby the fact that the sporocysts become dissolved up in the oocyst, thus leaving the 8 sporozoites unenclosed, recalling the condition inLégerella.P. prevoti, unique species, from the frog’s intestine.Genus 3.Crystallospora, Labbé. Spores also dizoic, but having the form of a double pyramid.C. crystalloidesfrom a fish,Motella tricirrata.Genus 4.Angeiocystis, Brasil. Apparently 6 sporozoites, but the only species,A. audouiniae, has only been briefly described; from a Polychaete (Audouinia).Fam.Polysporocystidae, Léger. The oocyst contains numerous spores.There are several genera with monozoic spores, characterized by variations in the form and structure of the sporocysts,e.g.Barroussia, Schn. (fig. 8),Echinospora, Léger, andDiaspora, Léger; most of these forms are from Myriapods.GenusAdelea, Schn. Dizoic spores; sporocysts round or oval, plain. Several species are included in this well-known genus, among them beingA. ovata,A. mesnili,A. dimidiata; most of them are parasitic in Insects or Myriapods.GenusMinchinia, Labbé. Dizoic spores; the sporocysts are produced at each pole into a long filament.M. chitonis, from the liver ofChiton(Mollusca).GenusKlossia, Schn. The spores are tetrazoic (or perhaps polyzoic).K. helicinafrom the kidney of various land-snails is the best-known form. Usually said to have 5 to 6 spores, but Mesnil considers that the normal number is 4, as is the case in another species,K. soror.GenusCaryotropha, Siedlecki. Many spherical spores (about 20)each with 12 sporozoites.C. mesnilii, unique species, from the spermatogonial (testis) cells ofPolymnia(a Polychaete). An interesting point in the schizogony is the formation of schizontocytes (see above).A Coccidian parasitic in the kidneys of the mouse has been described by Smith and Johnson (1902) and named by themKlossiella, on the ground that it possessed many spores, each with about 20 sporozoites. Woodcock has shown, however, that the authors were in all probability dealing with a similar modification of schizogony to that which obtains inCaryotropha. The sporogony of this form (and hence its systematic position) remains at present, therefore, quite unknown.There are several doubtful or insufficiently known genera,e.g.Bananella,Goussia,Hyaloklossia,Gonobia,PfeifferellaandRhabdospora, many of which probably represent only schizogonous generations of other forms. (For information concerning these see Labbé, 1897.)Lastly it remains to mention the extremely interesting forms parasitic in Cephalopods. For some years these have provided a fruitful source of discussion to systematists. Here it may be stated simply that their systematic position and nomenclature were thought to have been finally settled by the researches of Jacquemet (1903) and Lühe (1902) in the following terms:—GenusEucoccidium. Lühe (syn.LégerinaJacq.), Coccidia possessing polysporous oocysts and lacking schizogony, parasitic in Cephalopods. Two well-known species:E. eberthi(Labbé), (=BenedeniaseuKlossia e.seuoctopiana), parasitic inSepia, which is tri- or tetra-zoic; andE. octopianum(Schn.), (syn.BenedeniaseuKlossia o.) fromOctopus, which is polyzoic, having 10 to 12 sporozoites. In both forms cysts containing megaspores and megasporozoites, and others containing microspores and microsporozoites are found, considered as representing sexual differentiation thrown back to the very earliest stages of the life-cycle.Quite recently much additional light has been thrown upon our knowledge of these parasites, including a new one,E. jacquemeti. Moroff (1906) has shown that not one but many megagametes are formed, and fertilized by the microgametes. For this reason he regards them as Gregarines rather than Coccidia. Further, Léger and Duboscq (1906) have found that the characteristic coelomic parasites (Aggregata) of Crustacea, generally regarded as gymnosporous Gregarines (i.e.Gregarines in which the sporozoites are naked) constitute in reality nothing more or less than a schizogonous generation of these Cephalopodan parasites, which have thus an alternation of true hosts. The ripe sporocysts from the Cephalopod are eaten by a particular crab (e.g.PortunusorInachus, according to the parasite), the sporozoites are liberated and traverse the mucous membrane of the intestine, coming to rest in the surrounding lymphatic layer. Here a large “cyst” is formed, projecting into the body-cavity, the contents of which give rise to a great number of merozoites. On the crab being devoured by the right species of Cephalopod, the merozoites doubtless give rise to the sexual generation again.As the nameAggregatais much the older, and as, moreover, there is no longer any reason to retain that ofEucoccidium, these parasites must in future receive the former generic appellation. With regard to the various specific names, however, they remain quite unsettled until the life-history is properly worked out in different cases (see alsoGregarines).It seems to the writer a much more open question than Moroff and Léger and Duboscq apparently suppose, whether these parasites are to be relegated to the Gregarines. For undoubtedly they have many Coccidian features, and on the other hand they differ in many ways from Gregarines. The chief feature of agreement with the latter order is the possession of many female gametes. As already said, there can be little doubt that this was the condition in the Coccidian ancestor, and it is by no means impossible that one or two forms existing at the present day remain primitive in that respect. On the other hand, the advanced character of the parasitism (the parasites remaining intracellular up to and including gamete-formation); the entire lack of the characteristic feature of association; the schizogony, which is only a very rare occurrence in Gregarines, and which, in the present case, strongly suggests the process inCaryotrophaandKlossiella; and, last but not least, the varying number of the sporozoites (3 in one form, 10-15 in others), which is very different from the almost constant number (8) in Gregarines, are all characters in which these forms agree with Coccidia and not with Gregarines. Having regard to these points, the writer is inclined, for the present, to considerAggregataas an offshoot rather from the Coccidian than from the Gregarine branch of the Ectosporan tree.Bibliography.—The following are some of the important papers dealing with the order:—G. Bonnet-Eymard, “Sur l’Évolution de l’Eimeria nova, Schneider,”C.R. Soc. Biol.52, p. 659, 1900; L. Brasil, “Sur une Coccidie nouvelle, &c.,”C.R.Ac. Sci.139, p. 645, 1904; L. Cuénot, “Légerella testiculin. sp., &c.,”Arch. Zool. exp.(N. et R.), (3) 10, p. 49, 6 figs., 1902; M. Jacquemet, “Sur la systématique des Coccidies des Céphalopodes,”Arch. Protistenk.2 p., 190, 1903; A. Labbé, “Recherches zoologiques, cytologiques et biologiques sur les Coccidies,”Arch. zool. exp.(3), 4, p. 517, 3 pls., 1897; A. Laveran, “Sur les modes de réproduction d’Isospora lacazei,”C.R. Soc. Biol. 50, p. 1139, 1898; A. Laveran and F. Mesnil, “Sur deux Coccidies intestinales de laRana esculenta,” op. cit. 54, p. 857, 9 figs., 1902; A. Laveran and F. Mesnil, “Sur la Coccidie trouvée dans le rein de laRana esculenta, &c.,”C.R.Ac. Sci.135, p. 82, 10 figs., 1902; A. Laveran and F. Mesnil, “Sur quelques Protozoaires parasites d’une tortue, &c.”t. c.p. 609, 14 figs., 1902; L. Léger, “Sur une nouvelle Coccidie à microgamètes ciliés,” op. cit., 127, p. 418, 1898; L. Léger, “Sur la morphologie et le développement des microgamètes des Coccidies,”Arch. zool. exp.(N. et R.) (3), 6, 1898; L. Léger, “Essai sur la classification des Coccidies, &c.,”Ann. Mus. Nat. Hist., Marseille (2), Bull. i. p. 71, 4 pls., 1898; L. Léger “Sur la présence d’une Coccidie coelomique chez Olocrates, &c.,”Arch. zool. exp. (N. et R.) (3), 8, p. i., 1900; L. Léger, “Sur le genreEimeriaet la classification des Coccidies,”C.R. Soc. Biol.52, p. 575, 1900; L. Léger and O. Duboscq, “Recherches sur les Myriapodes de Corse et leurs parasites,”Arch. zool. exp.(4), 1, p. 307, 24 figs., 1903; L. Léger and O. Duboscq, “Sur l’évolution des Grégarines gymnosporées des Crustacés,”C.R.Ac. Sci.142, p. 1225, 1906; L. Léger and O. Duboscq, “L’Évolution d’uneAggregatade la seiche chez lePortunus depurator,”C.R. Soc. Biol. 6o, p. 1001, 1906; M. Lühe, “Über Geltung und Bedeutung der GattungsnamenEimeriaundCoccidium,” C.B. Bakter (1) 31 Orig, p. 771, 1902; C.B. Bakter, “Die Coccidien-Literatur der letzten vier Jahre,”Zool. Centrlbl.10, 45 pp., 1903; F. Mesnil, “Sur la conservation du nom génériqueEimeria, &c.,”C.R. Soc. Biol.52, p. 603, 1900; F. Mesnil, “Les Travaux récents sur les Coccidies,”Bull. Inst. Pasteur, i. pp. 473, 505, 1903; R. Metzner, “Untersuchungen an Coccidium cuniculi,”Arch. Protistenk.2, p. 13, pl. ii. 1903; G. Moussu and G. Marotel, “La Coccidiose du mouton et son parasite,”Arch. Parasitol.6, p. 82, 10 figs., 1902; T. Moroff, “Sur l’évolution des prétendues Coccidies des Céphalopodes,”C.R.Ac. Sci.142, p. 652, 1906; C. Perez, “Le Cycle évolutif de l’Adelea mesnili, &c.,”Arch. Protistenk.2, p. 1, pl. 1, 1903; F. Schaudinn, “Untersuchungen über den Generationswechsel bei Coccidien,”Zool. Jahrbücher(Anat.) 13, p. 197, 4 pls., 1900; F. Schaudinn, “Studien über krankheitserregende Protozoen—I.Cyclospora caryolytica, &c.,”Arb. kais. Gesundh.-amte, 18, p. 378, 2 pls., 1902; M. Siedlecki, “Réproduction sexuée ... chez ...Coccidium proprium,”C.R. Soc. Biol.50, p. 664, figs., 1898; M. Siedlecki, “Étude cytologique ... de la Coccidie de la seiche, &c.,”Ann. Inst. Pasteur, 12, p. 799, 3 pls., 1898; M. Siedlecki, “Étude cytologique ... de Adelea ovata,” op. cit. 13, p. 169, 3 pls., 1899; M. Siedlecki, “Cycle évolutif de laCaryotropha mesnilii, &c.,”Bull. Ac. Cracovie, p. 561, 5 figs., 1902; T. Smith and H. P. Johnson, “On a Coccidian (Klossiella muris, gen. et spec. nov.), &c.,”J. exp. Med. 6, p. 303, 3 pls., 1902; H.M. Woodcock, “Notes on Sporozoa, I. OnKlossiella muris, &c.,”Q.J. micr. Sci.48, p. 153, 2 figs., 1904.
The order Coccidiidea is divided into four families, characterized by the number of sporocysts (if any) found in the oocyst.
Classification.Fam.Asporocystidae, Léger. No sporozoites are formed in the oocyst, the sporozoites being unenclosed (gymnospores).
Genus,Légerella, Mesnil. This genus actually conforms to Aimé Schneider’s original definition ofEimeria, which was founded on what were really the schizogonous generations of other forms, then thought to be distinct. In view of the great confusion attending the use of this name, however, Mesnil (1900) has suggested the new one here adopted. Two species known,L. novaandL. testiculi, both from different species ofGlomeris, a Myriapod; the former inhabits the Malpighian tubules, the latter the testis.
Fam.Disporocystidae, Léger. The oocyst contains 2 spores.
Genus 1.Cyclospora, A. Schneider. Spores dizoic,i.e.with two sporozoites.C. glomericola, from the intestinal epithelium ofGlomeris, andC. caryolytica, from the intestinal epithelium of the mole, intranuclear.
Genus 2.Diplospora, Labbé. Spores tetrazoic.D. lacazei, from many birds, is the best-known species; and others have been described from different Sauropsida.D. lieberkühniis an interesting form occurring in the kidneys of the frog, which it reaches by way of the circulation.
Genus 3.Isospora, Schn. Spores polyzoic. Founded forI. rara, parasitic in the black slug (Limax cinereo-niger). Many authors consider that Schneider was mistaken in attributing many sporozoites to this form, and would unite with it the genusDiplospora.
Fam.Tetrasporocystidae, Léger. The oocyst contains 4 spores.
Genus 1.Coccidium,7Leuckart. The spores are dizoic and the sporocysts rounded or oval. A very large number of species are known, mostly from Vertebrate hosts.C. cuniculi(=C. oviforme) from the rabbit (intestine and diverticula), but also occurring sometimes in other domestic animals; C. falciformis, from the mouse;C. faureifrom sheep; andC. schubergi, fromLithobius(a centipede), are among the best-known forms. All of them may cause disastrous epidemics of coccidiosis.
Genus 2.Paracoccidium, Laveran and Mesnil. This genus is distinguished fromCoccidiumby the fact that the sporocysts become dissolved up in the oocyst, thus leaving the 8 sporozoites unenclosed, recalling the condition inLégerella.P. prevoti, unique species, from the frog’s intestine.
Genus 3.Crystallospora, Labbé. Spores also dizoic, but having the form of a double pyramid.C. crystalloidesfrom a fish,Motella tricirrata.
Genus 4.Angeiocystis, Brasil. Apparently 6 sporozoites, but the only species,A. audouiniae, has only been briefly described; from a Polychaete (Audouinia).
Fam.Polysporocystidae, Léger. The oocyst contains numerous spores.
There are several genera with monozoic spores, characterized by variations in the form and structure of the sporocysts,e.g.Barroussia, Schn. (fig. 8),Echinospora, Léger, andDiaspora, Léger; most of these forms are from Myriapods.
GenusAdelea, Schn. Dizoic spores; sporocysts round or oval, plain. Several species are included in this well-known genus, among them beingA. ovata,A. mesnili,A. dimidiata; most of them are parasitic in Insects or Myriapods.
GenusMinchinia, Labbé. Dizoic spores; the sporocysts are produced at each pole into a long filament.M. chitonis, from the liver ofChiton(Mollusca).
GenusKlossia, Schn. The spores are tetrazoic (or perhaps polyzoic).K. helicinafrom the kidney of various land-snails is the best-known form. Usually said to have 5 to 6 spores, but Mesnil considers that the normal number is 4, as is the case in another species,K. soror.
GenusCaryotropha, Siedlecki. Many spherical spores (about 20)each with 12 sporozoites.C. mesnilii, unique species, from the spermatogonial (testis) cells ofPolymnia(a Polychaete). An interesting point in the schizogony is the formation of schizontocytes (see above).
A Coccidian parasitic in the kidneys of the mouse has been described by Smith and Johnson (1902) and named by themKlossiella, on the ground that it possessed many spores, each with about 20 sporozoites. Woodcock has shown, however, that the authors were in all probability dealing with a similar modification of schizogony to that which obtains inCaryotropha. The sporogony of this form (and hence its systematic position) remains at present, therefore, quite unknown.
There are several doubtful or insufficiently known genera,e.g.Bananella,Goussia,Hyaloklossia,Gonobia,PfeifferellaandRhabdospora, many of which probably represent only schizogonous generations of other forms. (For information concerning these see Labbé, 1897.)
Lastly it remains to mention the extremely interesting forms parasitic in Cephalopods. For some years these have provided a fruitful source of discussion to systematists. Here it may be stated simply that their systematic position and nomenclature were thought to have been finally settled by the researches of Jacquemet (1903) and Lühe (1902) in the following terms:—
GenusEucoccidium. Lühe (syn.LégerinaJacq.), Coccidia possessing polysporous oocysts and lacking schizogony, parasitic in Cephalopods. Two well-known species:E. eberthi(Labbé), (=BenedeniaseuKlossia e.seuoctopiana), parasitic inSepia, which is tri- or tetra-zoic; andE. octopianum(Schn.), (syn.BenedeniaseuKlossia o.) fromOctopus, which is polyzoic, having 10 to 12 sporozoites. In both forms cysts containing megaspores and megasporozoites, and others containing microspores and microsporozoites are found, considered as representing sexual differentiation thrown back to the very earliest stages of the life-cycle.
Quite recently much additional light has been thrown upon our knowledge of these parasites, including a new one,E. jacquemeti. Moroff (1906) has shown that not one but many megagametes are formed, and fertilized by the microgametes. For this reason he regards them as Gregarines rather than Coccidia. Further, Léger and Duboscq (1906) have found that the characteristic coelomic parasites (Aggregata) of Crustacea, generally regarded as gymnosporous Gregarines (i.e.Gregarines in which the sporozoites are naked) constitute in reality nothing more or less than a schizogonous generation of these Cephalopodan parasites, which have thus an alternation of true hosts. The ripe sporocysts from the Cephalopod are eaten by a particular crab (e.g.PortunusorInachus, according to the parasite), the sporozoites are liberated and traverse the mucous membrane of the intestine, coming to rest in the surrounding lymphatic layer. Here a large “cyst” is formed, projecting into the body-cavity, the contents of which give rise to a great number of merozoites. On the crab being devoured by the right species of Cephalopod, the merozoites doubtless give rise to the sexual generation again.
As the nameAggregatais much the older, and as, moreover, there is no longer any reason to retain that ofEucoccidium, these parasites must in future receive the former generic appellation. With regard to the various specific names, however, they remain quite unsettled until the life-history is properly worked out in different cases (see alsoGregarines).
It seems to the writer a much more open question than Moroff and Léger and Duboscq apparently suppose, whether these parasites are to be relegated to the Gregarines. For undoubtedly they have many Coccidian features, and on the other hand they differ in many ways from Gregarines. The chief feature of agreement with the latter order is the possession of many female gametes. As already said, there can be little doubt that this was the condition in the Coccidian ancestor, and it is by no means impossible that one or two forms existing at the present day remain primitive in that respect. On the other hand, the advanced character of the parasitism (the parasites remaining intracellular up to and including gamete-formation); the entire lack of the characteristic feature of association; the schizogony, which is only a very rare occurrence in Gregarines, and which, in the present case, strongly suggests the process inCaryotrophaandKlossiella; and, last but not least, the varying number of the sporozoites (3 in one form, 10-15 in others), which is very different from the almost constant number (8) in Gregarines, are all characters in which these forms agree with Coccidia and not with Gregarines. Having regard to these points, the writer is inclined, for the present, to considerAggregataas an offshoot rather from the Coccidian than from the Gregarine branch of the Ectosporan tree.
Bibliography.—The following are some of the important papers dealing with the order:—G. Bonnet-Eymard, “Sur l’Évolution de l’Eimeria nova, Schneider,”C.R. Soc. Biol.52, p. 659, 1900; L. Brasil, “Sur une Coccidie nouvelle, &c.,”C.R.Ac. Sci.139, p. 645, 1904; L. Cuénot, “Légerella testiculin. sp., &c.,”Arch. Zool. exp.(N. et R.), (3) 10, p. 49, 6 figs., 1902; M. Jacquemet, “Sur la systématique des Coccidies des Céphalopodes,”Arch. Protistenk.2 p., 190, 1903; A. Labbé, “Recherches zoologiques, cytologiques et biologiques sur les Coccidies,”Arch. zool. exp.(3), 4, p. 517, 3 pls., 1897; A. Laveran, “Sur les modes de réproduction d’Isospora lacazei,”C.R. Soc. Biol. 50, p. 1139, 1898; A. Laveran and F. Mesnil, “Sur deux Coccidies intestinales de laRana esculenta,” op. cit. 54, p. 857, 9 figs., 1902; A. Laveran and F. Mesnil, “Sur la Coccidie trouvée dans le rein de laRana esculenta, &c.,”C.R.Ac. Sci.135, p. 82, 10 figs., 1902; A. Laveran and F. Mesnil, “Sur quelques Protozoaires parasites d’une tortue, &c.”t. c.p. 609, 14 figs., 1902; L. Léger, “Sur une nouvelle Coccidie à microgamètes ciliés,” op. cit., 127, p. 418, 1898; L. Léger, “Sur la morphologie et le développement des microgamètes des Coccidies,”Arch. zool. exp.(N. et R.) (3), 6, 1898; L. Léger, “Essai sur la classification des Coccidies, &c.,”Ann. Mus. Nat. Hist., Marseille (2), Bull. i. p. 71, 4 pls., 1898; L. Léger “Sur la présence d’une Coccidie coelomique chez Olocrates, &c.,”Arch. zool. exp. (N. et R.) (3), 8, p. i., 1900; L. Léger, “Sur le genreEimeriaet la classification des Coccidies,”C.R. Soc. Biol.52, p. 575, 1900; L. Léger and O. Duboscq, “Recherches sur les Myriapodes de Corse et leurs parasites,”Arch. zool. exp.(4), 1, p. 307, 24 figs., 1903; L. Léger and O. Duboscq, “Sur l’évolution des Grégarines gymnosporées des Crustacés,”C.R.Ac. Sci.142, p. 1225, 1906; L. Léger and O. Duboscq, “L’Évolution d’uneAggregatade la seiche chez lePortunus depurator,”C.R. Soc. Biol. 6o, p. 1001, 1906; M. Lühe, “Über Geltung und Bedeutung der GattungsnamenEimeriaundCoccidium,” C.B. Bakter (1) 31 Orig, p. 771, 1902; C.B. Bakter, “Die Coccidien-Literatur der letzten vier Jahre,”Zool. Centrlbl.10, 45 pp., 1903; F. Mesnil, “Sur la conservation du nom génériqueEimeria, &c.,”C.R. Soc. Biol.52, p. 603, 1900; F. Mesnil, “Les Travaux récents sur les Coccidies,”Bull. Inst. Pasteur, i. pp. 473, 505, 1903; R. Metzner, “Untersuchungen an Coccidium cuniculi,”Arch. Protistenk.2, p. 13, pl. ii. 1903; G. Moussu and G. Marotel, “La Coccidiose du mouton et son parasite,”Arch. Parasitol.6, p. 82, 10 figs., 1902; T. Moroff, “Sur l’évolution des prétendues Coccidies des Céphalopodes,”C.R.Ac. Sci.142, p. 652, 1906; C. Perez, “Le Cycle évolutif de l’Adelea mesnili, &c.,”Arch. Protistenk.2, p. 1, pl. 1, 1903; F. Schaudinn, “Untersuchungen über den Generationswechsel bei Coccidien,”Zool. Jahrbücher(Anat.) 13, p. 197, 4 pls., 1900; F. Schaudinn, “Studien über krankheitserregende Protozoen—I.Cyclospora caryolytica, &c.,”Arb. kais. Gesundh.-amte, 18, p. 378, 2 pls., 1902; M. Siedlecki, “Réproduction sexuée ... chez ...Coccidium proprium,”C.R. Soc. Biol.50, p. 664, figs., 1898; M. Siedlecki, “Étude cytologique ... de la Coccidie de la seiche, &c.,”Ann. Inst. Pasteur, 12, p. 799, 3 pls., 1898; M. Siedlecki, “Étude cytologique ... de Adelea ovata,” op. cit. 13, p. 169, 3 pls., 1899; M. Siedlecki, “Cycle évolutif de laCaryotropha mesnilii, &c.,”Bull. Ac. Cracovie, p. 561, 5 figs., 1902; T. Smith and H. P. Johnson, “On a Coccidian (Klossiella muris, gen. et spec. nov.), &c.,”J. exp. Med. 6, p. 303, 3 pls., 1902; H.M. Woodcock, “Notes on Sporozoa, I. OnKlossiella muris, &c.,”Q.J. micr. Sci.48, p. 153, 2 figs., 1904.
(H. M. Wo.)
1A curious organism, parasitic in a gregarine, has lately been described by Dogiel as a coccidian, and termedHyalosphaera.2It is important to note that in schizogony there is never any cyst or cyst-membrane formed around the parasite.3The merozoites are frequently arranged like the staves of a barrel—whence the termbarillet, which is frequently used.4InCyclospora, Schaudinn (1902) has noted certain abnormal cases of the persistence and further multiplication of the “reduction-nuclei” of the female element (i.e.the nuclear portions given off during maturation), followed by multiple fertilization. This occurrence points strongly to the conclusion that there were originally many female gametes (cf. also the sporoblasts of Gregarines).5The remarkable forms parasitic in Cephalopods (of late known asEucoccidium), if still ranked with the Coccidia, furnish an exception (see below).6Again with the exception ofEucoccidium.7Purists in systematic nomenclature maintain that this name should be relinquished in favour ofEimeria, since the latter was the first legitimate generic name given to a Coccidian. But one reason against the use ofEimeriahas been stated already (it should be used forE. (Légerella) nova, if anywhere); and in addition, the wordCoccidiumand its important derivatives are now so universally established that it would be little short of ridiculous to displace them.
1A curious organism, parasitic in a gregarine, has lately been described by Dogiel as a coccidian, and termedHyalosphaera.
2It is important to note that in schizogony there is never any cyst or cyst-membrane formed around the parasite.
3The merozoites are frequently arranged like the staves of a barrel—whence the termbarillet, which is frequently used.
4InCyclospora, Schaudinn (1902) has noted certain abnormal cases of the persistence and further multiplication of the “reduction-nuclei” of the female element (i.e.the nuclear portions given off during maturation), followed by multiple fertilization. This occurrence points strongly to the conclusion that there were originally many female gametes (cf. also the sporoblasts of Gregarines).
5The remarkable forms parasitic in Cephalopods (of late known asEucoccidium), if still ranked with the Coccidia, furnish an exception (see below).
6Again with the exception ofEucoccidium.
7Purists in systematic nomenclature maintain that this name should be relinquished in favour ofEimeria, since the latter was the first legitimate generic name given to a Coccidian. But one reason against the use ofEimeriahas been stated already (it should be used forE. (Légerella) nova, if anywhere); and in addition, the wordCoccidiumand its important derivatives are now so universally established that it would be little short of ridiculous to displace them.
COCCULUS INDICUS, the commercial name for the dried fruits ofAnamirta Cocculus(natural order Menispermaceae), a large climbing shrub, native to India. It contains a bitter poisonous principle,picrotoxin, used in small doses to control the night sweats of phthisis. It was formerly known as Levant nut and Levant shell, owing to the fact that it was brought to Europe by way of the Levant.
COCHABAMBA, a central department of Bolivia, occupying the eastern angle of the great Bolivian plateau, bounded N. by the department of El Beni, E. by Santa Cruz, S. by Chuquisaca and Potosi, and W. by Potosi, Oruro and La Paz. Area, 23,328 sq. m.; pop. (1900) 328,163. Its average elevation is between 8000 and 10,000 ft., and its mean temperature ranges from 50° to 60° F., making it one of the best climatic regions in South America. The rainfall is moderate and the seasons are not strongly marked, the difference being indicated by rainfall rather than by temperature. The rainy season is from November to February. Cochabamba is essentially an agricultural department, although its mineral resources are good and include deposits of gold, silver and copper. Its temperate climate favours the production of wheat, Indian corn, barley and potatoes, and most of the fruits and vegetables of the temperate zone. Coca, cacáo, tobacco and most of the fruits and vegetables of the tropics are also produced. Its forest products include rubber and cinchona. Lack of transportation facilities, however, have been an insuperable obstacle to the development of any industry beyond local needs except those of cinchona and rubber. Sheep and cattle thrive in this region, and an experiment with silkworms gave highly successful results. The population is chiefly of the Indian andmestizotypes, education is in a backward state, and there are no manufactures other than those of the domestic stage, the natives making many articles of wearing apparel and daily use in their own homes. Rough highways andmule-paths are the only means of communication, but a projected railway from Cochabamba (city) to Oruro, 132 m., promises to bring this isolated region into touch with the commercial world. The department is divided into nine provinces, but there is no effective local government outside the municipalities. The capital is Cochabamba; other important towns are Punata, Tarata, Totora, Mizque and Sacába.
COCHABAMBA, a city of Bolivia, capital of the department of the same name and of the province of Cercado, situated on the Rocha, a small tributary of the Guapay river, in lat. 17° 27′ S. and long. 65° 46′ W. Pop. (1900) 21,886, mostly Indians andmestizos. The city stands in a broad valley of the Bolivian plateau, 8400 ft. above sea-level, overshadowed by the snow-clad heights of Tunari and Larati, 291 m. north-north-west of Sucre and 132 m. east-north-east of Oruro, with both of which places it is connected by rough mountain roads. A subsidized stage-coach line runs to Oruro. A contract for a railway between the two cities was made in 1906, connecting with the Antofagasta and Arica lines. The climate is mild and temperate, and the surrounding country fertile and cultivated. Cochabamba is often described as the most progressive city of Bolivia, but it has been held back by its isolated situation. The warehouses of the city are well supplied with foreign goods, and trade is active in spite of high prices. The city is provided with telegraphic communication via Oruro, and enjoys a large part of the Amazon trade through some small river ports on tributaries of the Mamoré. The city is regularly laid out, and contains many attractive residences surrounded by gardens. It is an episcopal city (since 1847), containing many churches, four conventual establishments, and a missionary college of the “Propaganda Fide” for the conversion of Indians. The city has a university and two colleges, but they are poorly equipped and receive very little support from the government. Cochabamba was founded in the 16th century, and for a time was called Oropesa. It took an active part in the “war of independence,” the women distinguishing themselves in an attack on the Spanish camp in 1815, and some of them being put to death in 1818 by the Spanish forces. In 1874 the city was seized and partly destroyed by Miguel Aguirre, but in general its isolated situation has been a protection against the disorders which have convulsed Bolivia since her independence.
COCHEM, a town of Germany, in the Prussian Rhine province on the Mosel, and 30 m. W. of Coblenz by the railway to Trier, which above the town enters the longest tunnel (2½ m.) in Germany. Pop. 3500. It is romantically situated in the deep and winding valley of the Mosel, at the foot of a hill surrounded by a feudal castle dating from 1051, which has been restored in its former style. There is a considerable trade in wines.
COCHERY, LOUIS ADOLPHE(1819-1900), French statesman, was born at Paris. After studying law he soon entered politics, and was on the staff of the ministry of justice after the revolution of February 1848. From thecoup d’étatof 1851 to May 1869 he devoted himself to journalism. Then, elected deputy by the department of the Loiret, he joined the group of the Left Centre, and was a supporter of the revolution of the 4th of September 1870. His talent in finance won him a distinguished place in the chamber. From 1879 till 1885 he was minister of posts and telegraphs, and in January 1888 he was elected to the senate. He died in 1900.
His son,Georges Charles Paul, born in 1855, was in his father’s department from 1879 till 1885, deputy from 1885, five times president of the Budget Commission, minister of finance (1895-1898) and vice-president of the chamber (1898-1902), and again finance minister in the Briand Cabinet, 1909.
COCHIN, DENYS MARIE PIERRE AUGUSTIN(1851- ), French politician, was born at Paris. He studied law, was elected to the chamber of deputies in 1893, and gradually became one of the leaders and principal orators of the Conservative party. He opposed the project of the income-tax in 1894, the revision of the Dreyfus case in 1899, and the separation of the church and state in 1905. He is known as an author by his works,L’Évolution de la vie(1895);Le Monde extérieur(1895);Contre les barbares(1899);Ententes et ruptures(1905).
COCHIN, a feudatory state of southern India, in political subordination to Madras, with an area of 1361 sq. m. It is bounded on the N. by British Malabar, on the E. by British Malabar, Coimbatore and Travancore, on the S. by Travancore, and on the W. by British Malabar and the Arabian Sea. Isolated from the main territory, and situated to the north-east of it, lies the major portion of the Chittoretaluk, entirely surrounded by British territory. The whole state may be divided into three well-defined regions or zones: (1) the eastern zone, consisting of broken forested portions of the Western Ghats, which, gradually decreasing in height, merge into (2) the central belt, comprising the uplands and plains that dip towards the lagoons or “backwaters” along the coast (seeCochin, town), beyond which lies (3) the western zone, forming the littoral strip. The low belt which borders on the seas and the backwaters is by nature flat and swampy, but has in the course of ages become enriched by the work of man. On leaving the seaboard, an undulating country is found, diversified with grassy flats, naked hills and wooded terraces, intersected by numerous torrents and rapids, and profusely dotted with homesteads, orchards and cultivated fields, up to the very foot of the Ghats. Here the landscape, now on a grander scale, embraces great forests which form a considerable source of wealth. Of the total area of the state the forests and lagoons cover nearly 605 and 16 sq. m. respectively.
In 1901 the population was 812,025, showing an increase of 12% in the decade. More than one-fifth are Christians, mostly Syrians and Roman Catholics. The revenue is estimated at £153,000, subject to a tribute of £13,000. During recent years the financial condition of the state has been flourishing. The principal products are rice, cocoanuts, timber, cardamoms, pepper and a little coffee. Salt is manufactured along the coast. The capital is Ernakulam, but the raja resides at Tripunthora. The principal commercial centre is Mattancheri, adjoining the British town of Cochin. The chief means of communication is by boat along the backwaters; but in 1902 a metre-gauge line was constructed by the Madras railway at the expense of the state to connect Ernakulam with Shoranur.
History.—What is now the native state of Cochin formed, until about the middle of the 9th centuryA.D., part of the ancient Chera or Kerala kingdom (seeKerala). Its port of Kodungalur (Kranganur, the ancient Muziris), at the mouth of the Periyar, was from early times one of the chief centres for the trade between Europe and India; and it was at Malankara, near Kodungalur, that the apostle Thomas is traditionally said to have landed. The history of Cochin is, however, like that of the Kerala kingdom generally, exceedingly obscure previous to the arrival of the Portuguese. The rajas of Cochin, who are of pure Kshatriya blood, claim descent from the Chera king Cheraman Perumal, the last of his race to rule the vast tract from Gokarn in North Kanara to Cape Comorin. About the middle of the 9th century this king, according to tradition, resigned his kingdom, embraced Islam, and went on pilgrimage to Arabia, where he died. Towards the end of the century the Chera kingdom was overrun and dismembered by the Cholas. It was in 1498 that Vasco da Gama reached the Malabar coast; and in 1502 the Portuguese were allowed to settle in the town of Cochin, where they built a fort and began to organize trade with the surrounding country. By the end of the century their influence had become firmly established, largely owing to the effective aid they had given to the rajas of Cochin in their wars with the Zamorin of Calicut. The Syrian Christians, forming at that time a large proportion of the population, now felt the weight of Portuguese ascendancy; in 1599 Menezes, the archbishop of Goa, held a synod at Udayamperur (Diamper), a village 12 m. south-east of Cochin, at which their tenets were pronounced heretical and their service-books purged of all Nestorian phrases. In 1663, however, Portuguese domination came to an end with the capture of Cochin by the Dutch, whose ascendancy continued for about a hundred years. In 1776 Hyder Ali of Mysore invaded thestate and forced the raja to acknowledge his suzerainty and pay tribute. In 1791 Tippoo, son of Hyder Ali, ceded the sovereignty to the British, who entered into a treaty with the raja by which he became their vassal and paid an annual tribute of a lakh of rupees. On the 17th of October 1809, in consequence of an attempt of the hereditary chief minister Paliyath Achan, in 1808, to raise an insurrection against the British without his master’s knowledge, a fresh treaty was made, by which the raja undertook to hold no correspondence with any foreign state and to admit no foreigners to his service without the sanction of the British government, which, while undertaking to defend the raja’s territories against all enemies, reserved the right to dismantle or to garrison any of his fortresses. In 1818 the tribute, raised to 2½ lakhs in 1808, was permanently fixed at 2 lakhs. Since then, under the rule of the rajas, the state has greatly advanced in prosperity, especially under that of H. H. Sir Sri Rama Varma (b. 1852), who succeeded in 1895, was made a K.C.S.I. in 1897, and G.C.S.I. in 1903.
COCHIN, a town of British India, in the district of Malabar, Madras. Pop. (1901) 19,274. The town lies at the northern extremity of a strip of land about 12 m. in length, but in few places more than a mile in breadth, which is nearly insulated by inlets of the sea and estuaries of streams flowing from the Western Ghats. These form the Cochin backwaters, which consist of shallow lagoons lying behind the beach-line and below its level. In the monsoon the Cochin backwaters are broad navigable channels and lakes; in the hot weather they contract into shallows in many places not 2 ft. deep. The town of Cochin is about a mile in length by half a mile in breadth. Its first European possessors were the Portuguese. Vasco da Gama founded a factory in 1502, and Albuquerque built a fort, the first European fort in India, in 1503. The British made a settlement in 1634, but retired when the Dutch captured the town in 1663. Under the Dutch the town prospered, and about 1778 an English traveller described it as a place of great trade, “a harbour filled with ships, streets crowded with merchants, and warehouses stored with goods from every part of Asia and Europe, marked the industry, the commerce, and the wealth of the inhabitants.” In 1795 Cochin was captured from the Dutch by the British, and in 1806 the fortifications and public buildings were blown up by order of the authorities. The explosion destroyed much private property, and for a long time seriously affected the prosperity of the town. Considerable sea-borne trade is still carried on. A lighthouse stands on the ruins of the old fort. The chief exports are cocoanut products, for the preparation of which there are factories, and tea; and the chief import is rice. Cochin is the only port south of Bombay in which large ships can be built.
COCHIN-CHINA,1a French colony in the extreme south of French Indo-China. The term formerly included the whole Annamese empire—Tongking, Annam, and Lower Cochin-China, but it now comprises only the French colony, which corresponds to Lower Cochin-China, and consists of the six southern provinces of the Annamese empire annexed by France in 1862 and 1867. Cochin-China is bounded W. by the Gulf of Siam, N.W. and N. by Cambodia, E. by Annam, and S.E. by the China Sea. Except along part of the north-west frontier, where the canal of Vinh-Thé divides it from Cambodia, its land-limits are conventional. Its area is about 22,000 sq. m.
In 1901 the population numbered 2,968,529, of whom 4932 were French (exclusive of French troops, who numbered 2537), 2,558,301 Annamese, 231,902 Cambodians, 92,075 Chinese, 42,940 savages (Min Huong), the rest being Asiatics of other nationalities, together with a few Europeans other than French.
Geography.—Cochin-China consists chiefly of an immense plain, flat and monotonous, traversed by the Mekong and extending from Ha-Tien in the west to Baria in the east, and from Bien-Hoa in the north-east to the southern point of the peninsula of Ca-Mau in the south-west. The last spurs of the mountains of Annam, which come to an end at Cape St Jacques, extend over parts of the provinces of Tay-Ninh, Bien-Hoa and Baria in the north-east and east of the colony, but nowhere exceed 2900 ft. in height; low hills are found in the north-western province of Chau-Doc. Cochin-China is remarkable for the abundance of its waterways. The Mekong divides at Pnom-Penh in Cambodia into two arms, the Fleuve supérieur and the Fleuve inférieur, which, pursuing a course roughly parallel from north-west to south-east, empty into the China Sea by means of the numerous channels of its extensive delta. From June to October the inundations of the Mekong cover most of the country, portions of which, notably the Plaine des Joncs in the north and a large tract of the peninsula of Ca-Mau, are little else than marshes. Besides a great number of small coastal streams there are four other rivers of secondary importance, all of which water the east of the colony, viz. the Don-Nai, which rising in the Annamese mountains flows west, then abruptly south, reaching the sea to the west of Cape St Jacques; the Saigon river, which flowing from north-west to south-east passes Saigon, the capital of the colony, 12 m. below which it unites with the Don-Nai; and the two Vaicos, which join the Don-Nai close to its mouth. These rivers flow into the sea through numerous winding channels, forming a delta united by canals to that of the Mekong. The waterways of Cochin-China communicate by means of natural or artificial channels (arroyos), facilitating transport and aiding in the uniform distribution of the inundation to which the country owes its fertility. Canals from Chau-Doc to Ha-Tien and from Long Xuyen to Rach-Gia join the Mekong with the Gulf of Siam. East of Cape St Jacques the mountains of Annam come down close to the sea; west of that point, as far as the southern headland of Ca-Mau, the coast-line of Cochin-China runs north-east to south-west for about 160 m. in a straight line broken only by the mouths of the Don-Nai and Mekong. From Cape Ca-Mau to Rach-Gia it runs north for a distance of 120 m., then north-west as far as Ha-Tien, where the boundary line between it and Cambodia meets the sea.
Climate and Fauna.—The climate of the country is warm, humid, and very trying to Europeans. The wet season, during which heavy rain falls almost daily, lasts from April to October, coinciding with the south-west monsoon. The hottest period lasts from the middle of April to the middle of June, the thermometer during that time often reaching 94° F., and never descending below 86°. The forest regions of Cochin-China harbour the tiger, panther, leopard, tiger-cat, ichneumon, wild boar, deer, buffalo, rhinoceros and elephant, as well as many varieties of monkeys and rats. Of birds some species of parrakeet, the “mandarin” blackbird, and the woodcock are not found in the rest of Indo-China. Duck, teal, cranes and other aquatic birds abound in the delta. Venomous reptiles are numerous, and the Mekong contains crocodiles.
Agriculture and Industries.—The cultivation of the rice-fields, which cover large extents of the plains of Cochin-China, is by far the chief industry of the colony. Pepper is grown in considerable quantities in the districts of Ha-Tien and Bien-Hoa, and sugarcanes, coffee, cotton, tobacco and jute are also produced. The buffalo, used both for transport and in the rice-fields, and swine, the flesh of which forms an important element in the native diet, are the principal domestic animals. Oxen and cows are of secondary importance and the climate is unsuitable for sheep; horses of a small breed are used to some extent. The chief industrial establishments are those for the decortication of rice at Saigon and Cholon; they are in the hands of the Chinese, by whom most of the trade in the colony is conducted. Sugar-making, the distillation of rice-spirit, silk-weaving, fishing and the preparation of a fish-sauce (nuoc-mam) made from decayed fish, and the manufacture of salt from sea-water and of lime are carried on in many localities.
Commerce.—Rice is the chief article of export, dried or salted fish, pepper and cotton ranking next in order of value. Imports include woven goods, metals, ironware, machinery, tea, wines and spirits, mineral oils, opium, paper, and arms and powder. The ports of Saigon and Mytho are accessible to the largest vessels, and are connected by a railway (seeIndo-China, French). The roadsteads of Rach-Gia, Ca-Mau, and Ha-Tien can accommodate only vessels of low tonnage. In 1905 exportsreached a value of £3,816,000, and imports a value of £4,834,000 (not including treasure and transit trade).
Government and Administration.—Cochin-China is administered by a lieutenant-governor under the authority of the governor-general of Indo-China. He is assisted by theconseil colonialnumbering sixteen members, six of whom are French citizens elected by the French, six natives elected by the natives, the other four being members of the chamber of commerce of Saigon and theconseil privé. Theconseil colonial, besides its advisory functions, discusses and votes the budget, determines the nature of the taxes, has supreme control over the tariffs, and extensive powers in the administration of colonial domains. Theconseil privéis a deliberative body under the presidency of the lieutenant-governor, composed of colonial officials together with two native members. The colony is divided into four circumscriptions (Saigon, My-Tho, Vinh-Long, Bassac), at the head of each of which is an inspector of native affairs. These are subdivided into twenty provinces, each administered by an administrator of native affairs by whose side is the provincial council consisting of natives and occupied with the discussion of ways and means and questions of public works. The provinces are divided into cantons and subdivided into communes. The commune forms the basis of the native social system. Its assembly of notables or municipal council forms a sort of oligarchy, the members of which themselves elect individuals from among the more prominent inhabitants to fill vacancies. The notables elect the provincial councillors in the proportion, usually, of one to every canton, and their delegates elect the chief of the canton, who voices the wishes of the natives to the government. Local administration,e.g.supervision of markets, policing, land-transfer, &c., are carried on by a mayor and two assistants, to whom the municipal council delegates its powers. The same body draws up the list of males liable to the poll-tax and of the lands liable to land-tax, these being the chief sources of revenue. There are French tribunals of first instance in nine of the chief towns of the colony, and in four of these there are criminal courts. These administer justice in accordance both with French law and, in the case of natives, with Annamese law, which has been codified for the purpose. Saigon has two chambers of the court of appeal of French Indo-China and a tribunal of commerce. Primary instruction is given in some six hundred schools. Cochin-China is represented in the French chamber by a deputy. The capital is Saigon (q.v.); of the other towns, Cholon (q.v.), My-Tho, Vinh-Long and Chau-Doc are of importance.
In 1904 the budget receipts amounted to £495,241 (as compared with £474,545 in 1899). To this sum the land and poll-tax and other direct taxes contributed £374,630. The main heads of expenditure, of which the total was £467,328, were as follows:—