Various types of form are to be met with among the Haemosporidia. In one, characteristic of most (though not of absolutely all) parasites of warm-blooded Vertebrates, the trophozoites are of irregular amoeboid shape; hence this sectionComparative Morphology; variations in the life-cycle where known.is generally known as theHaemamoebidae. In another type, characteristic of the parasites of cold-blooded Vertebrates, the body possesses a definite, vermiform,i.e.gregariniform shape, which is retained during the intracorpuscular as well as during the free condition; this section comprises theHaemogregarinidae. Allied to this latter type of form are the trophozoites ofPiroplasma, which are normally pear-shaped; they differ, however, in being very minute, and, moreover, exhibit considerable polymorphism, rod-like (so-called bacillary) and ring-forms being of common occurrence. It is important to note that in a certain species ofHaemogregarina(fig. 3) the young trophozoites markedly resemblePiroplasmain their pyriform appearance; and a further point of agreement between the two forms is mentioned below. Lastly there is the Avian genusHalteridium, the trophozoites of which are characteristically bean-shaped or reniform. True Haemogregarines also differ in other slight points from “Haemamoebae.” Thus the young endoglobular trophozoite does not exhibit a ring (vacuolar) phase; and the cytoplasm never contains, at any period, the characteristic melanin pigment above noted. In some species ofHaemogregarinathe parasite, while intracorpuscular, becomes surrounded by a delicate membrane, the cytocyst; on entering upon an active, “free” period, the cytocyst is ruptured and left behind with the remains of the corpuscle. A very interesting cytological feature is the occurrence, in one or two Haemosporidia, of nuclear dimorphism,i.e.of a larger and smaller chromatic body, probably comparable to the trophic and kinetic nuclei of a Trypanosome, or of the “Leishman-Donovan” bodies. Schaudinn was the first to notice this character, inPiroplasma canis, and his observation has since been confirmed by Lühe.4Moreover, Brumpt has also noticed nuclear dimorphism in the ookinete of a species ofHaemogregarinain a leech (as the Invertebrate host)—a highly important observation.As regards the life-history, the endogenous (schizogonous) cycle is known in many cases. Sometimes schizogony takes the primitive form of simple binary (probably) longitudinal fission; this is the case inPiroplasma(fig. 4) and also inHaemogregarina bigeminajust referred to. From this result the pairs of individuals (“twins”) so often found in the corpuscles. In addition, however, at any rate inPiroplasma, it is probable that multiple division (more allied to ordinary schizogony) also takes place; such is the case, according to Laveran, inP. equi, and the occurrence at times of four parasites in a corpuscle, arranged in a cruciform manner, is most likely to be thus explained. Labbé has described schizogony inHalteridium danilewskyias taking place in a rather peculiar manner; the parasite becomes much drawn-out and halter-like, and the actual division is restricted to its two ends, two clumps of merozoites being formed, at first connected by a narrow strand of unused cytoplasm, which subsequently disappears. Some doubt, however, attaches to this account, as no one else appears to have seen the process. For the rest, schizogony takes place more or less in the customary way, allowing for variations in the mode of arrangement of the merozoites. It remains to be noted that inKaryolysus lacertarum, according to Labbé, two kinds of schizont are developed, which give rise, respectively, to micromerozoites and megamerozoites, in either case enclosed in a delicate cytocyst. This probably corresponds to an early sexual differentiation (such as is found among certain Coccidia (q.v.), the micromerozoites producing eventually micro-gametocytes, the others megagametocytes.From Lankester’sTreatise on Zoology.Fig. 4.—Development and schizogony ofPiroplasma bigeminumin the blood-corpuscles of the ox. (After Laveran and Nicolle.)a, Youngest form.b, Slightly older.candd. Division of the nucleus.eandf, Division of the body of the parasite.g,h,i,j, Various forms of the twin parasite.kandl, Doubly infected corpuscles.It has now been recognized for some time that the sexual (exogenous) part of the life-cycle of all theHaemamoebidaetakes place in an Invertebrate (Insectan) host, and is fundamentally similar to that above described in those cases where it has been followed. In contradistinction to the malarial parasites, this host, in the Avian forms (HaemoproteusandHalteridium)5is a species ofCulexand not ofAnopheles; in other words, gamete-formation, conjugation and subsequent sporozoite-formation in these cases will only go on in the former. On the other hand, in the case of the Haemogregarines, it was thought until quite lately that the entire life-history, including conjugation and sporogony, went on in the Vertebrate host; and only in 1902 Hintze described what purported to be the complete life-history ofLankesterella(Drepanidium)ranarumundergone in the frog. This view was rendered obsolete by the work of Siegel and Schaudinn, who demonstrated the occurrence of an alternation of hosts and of generations in the case ofHaemogregarina stepanovi, parasitic in a tortoise, and inKaryolysus lacertarum; the Invertebrate hosts, in which, in both cases, the sexual process is undergone, being respectively a leech (Placobdella) and a tick (Ixodes). With this discovery the main distinction (as supposed) between the Haemosporidia of warm and of cold-blooded Vertebrates vanished. It was further acknowledged by Schaudinn (under whom Hintzehad worked) that the latter had been misled by Coccidian cysts and spores, which he took for those ofLankesterella. The gametogony and sporogony ofHaemogregarina stepanoviin the leech agree in essential particulars with the process above described. The microgametes are extremely minute, and the sporozoites, which are developed in the salivary glands, where the motile ookinetes finally come to rest, are extremely “spirochaetiform”—the full significance of this latter fact being, perhaps, not appreciated.Christophers recently described some remarkable phases which he regarded as belonging to the cycle ofHaemogregarina gerbilli(one of the few Mammalian Haemogregarines known) in a louse (Haematopinus). In a private communication, however, the author states that he has probably mistaken phases in the development of an ordinary gregarine parasite in the louse for part of the life-cycle of this Haemogregarine.The Mammalian parasitePiroplasmais the one about whose life-history our knowledge is most vague. Besides the typical and generally occurring forms, others have also been observed in the blood, but it is doubtful how far these are to be looked upon as normal; for instance, Bowhill and Le Doux have described, in various species, a phase in which a long, slender pseudopodial-like outgrowth is present, with a swelling at the distal end. It is, moreover, quite uncertain which are the sexual forms, comparable to gametocytes. Doflein regards large pear-shaped forms as such (megagametocytes?), which become spherical when maturing; and Nocard and Motas have figured amoeboid, irregular forms, with the nucleus fragmented and possessing flagella-like processes (possibly microgametes?). The Invertebrate host is well known to be, in the case of all species, a tick; thus bovine piroplasmosis (P. bigeminum) in America is conveyed byRhipicephalus annulatus(Boophilus bovis), canine piroplasmosis (P. canis) in South Africa byHaemaphysalis leachi(and perhapsDermacentor reticulatus), and so on. The manner in which the infection is transmitted by the tick varies greatly. In some cases (e.g.P. bigeminumandP. canis) only the generation subsequent to that which receives the infection (by feeding on an infected ox) can transmit it back again to another ox; in other words, true hereditary infection of the ova in the mother-tick is found to occur. The actual period in the life of the daughter-tick at which it can convey the infection apparently varies. On the other hand, in the case of East African coast-fever, Theiler found that hereditary infection does not occur, the same generation transmitting the parasite (P. parvum) at different periods of life. Little is certainly known regarding the phases of the parasite which are passed through in the tick. Lignières has observed a kind of multiple fission in the stomach, several very minute bodies, consisting mostly of chromatin, being formed, which may serve for endogenous reproduction. Koch has published an account of certain curious forms ofP. bigeminum, in which the body is produced into many stiff, ray-like processes, giving the appearance of a star; according to him fusion of such forms takes place, and the resulting zygote becomes rounded, perhaps transitional to the pear-shaped forms.The classification and nomenclature of the Haemosporidia are in a very unsettled condition. For an account of the various systems and modifications hitherto adopted, the article of Minchin (see underSporozoa:Bibliography) should be consulted.Classification.With the realization that the life-history in the case of the “Haemamoebae” and the Haemogregarines is fundamentally similar in type, the chief reason for grouping them as distinct suborders has disappeared. It is most convenient to regard them as separate, but closely allied families, thePlasmodidae(“Haemamoebidae”) and theHaemogregarinidae. ThePiroplasmata, on the other hand, constitute another family, which is better placed in a distinct section or sub-order. In addition there are, as already noted, two or three genera whose systematic position must be considered as quite uncertain. One is the well-knownHalteridiumof Labbé, parasitic in various birds; the type-species isH. danilewskyi(Gt. and Fel.). Another is the much-debated parasite of white blood-corpuscles (leucocytes), originally described in birds by Danilewsky under the name ofLeucocytozoon, a form of which has been recently observed in Mammals.In conclusion, the chief members of the above-mentioned families may be enumerated.Fam.Plasmodidae(“Haemamoebidae”).GenusLaverania, Gr. and Fel. (syn.Haemamoenas, Ross), for L.malariae, Gr. and Fel. (synn. L. s.Plasmodium, s. “Haemamoeba,” &c.,praecoxs.immaculatum, &c.), the parasite of pernicious malaria. GenusPlasmodium, March. and Celli (syn. “Haemamoeba”) forP. vivaxandP. malariae, the tertian and quartan parasite, respectively. There is also a form known in apes,P. kochi. GenusHaemoproteus, Kruse (syn.Proteosoma), forH. danilewskyi(syn.Proteosoma grassi,Plasmodium praecox, &c.), parasitic in numerous birds. Recently, another form has been described, from reptiles, which Castellani and Willey have termedHaemocystidium simondi.Remarks.—The distinguishing characters of the malarial parasites have been mentioned above. Some authorities would includeLaveraniain the genusPlasmodium, as differing only specifically from the other two forms. It has, moreover, been suggested by Sergent that all three are merely different phases of the same parasite, predominating at different seasons; this idea cannot be regarded, however, as in any way proved so far. From what is known of the morphology and mode of manifestation of these forms, the differences betweenLaveraniaand the two species ofPlasmodiumare considerably more pronounced than those betweenP. vivaxandP. malariae; if the latter are to be considered as distinct species, the first-named is probably generically distinct. Lühe, it may be noted, in his recent comprehensive account of the Haematozoa, also takes this view. Lastly, whatever be the correct solution of the above problem, there is certainly not sufficient justification for including the Avian genusHaemoproteus, as also only a species ofPlasmodium, which is done by some. Its different Vertebrate habitat, and also the fact that its Insectan definitive host is Culex and notAnopheles, differentiate it sharply fromLaveraniaandPlasmodium.From Lankester’sTreatise on Zoology.Fig. 5.—Haemoproteus danilewskyi, Kruse (parasite of various birds). × about 1200.a,b,canddfrom the chaffinch;dandefrom the lark. (After Labbé.)a, Young trophozoite in a blood-corpuscle,bandc, Older trophozoite.dande, Sporulation.d, Precocious sporulation with few merozoites.e, Sporulation of a full-grown schizont, with numerous merozoites.f, Gametocyte.N, Nucleus of blood-corpuscle.n, Nucleus of parasite.p, Pigment.mz, Merozoites.r.p, Residual protoplasm.From Lankester’sTreatise on Zoology.Fig. 6.—Haemogregarina stepanovi, Danilewsky (par.EmysandCistudo), phases of the schizogony. (a-eandjafter Laveran;f-iafter Börner.) × 1000 to 1200 diameters.a, Blood-corpuscle with young trophozoite.b, Older trophozoite.c, Full-grown trophozoite, ready to leave the corpuscle.dande, Trophozoites free in the blood-plasma, showing changes of form.f-i, Trophozoites, still within the blood-corpuscle (not drawn), showing the structure of the nucleus, the coarse chromatoid granules in the protoplasm and the manner in which the parasite grows into theU-shaped Haemogregarine without increase of body-mass.j, Commencement of sporulation; the nucleus has divided into eight nuclei, and the body of the parasite is beginning to divide up into as many merozoites within a blood-corpuscle.N, Nucleus of the blood-corpuscle.n, Nucleus of the parasite.Fam.Haemogregarinidae.—The different genera are characterized chiefly by their size relative to the blood-corpuscles, and their disposition in the latter. Here, again, it has been suggested to unite the various types all in one genus,Haemogregarina, but this seems at least premature when it is remembered how little is known in most cases of the life-cycle, which may prove to exhibit important divergences.GenusHaemogregarina, Danilewsky (syn.Danilewskya, Labbé).The body of the parasite exceeds the blood-corpuscle in length, when adult, and is bent upon itself, like aU. A very great number of species are known, mostly from reptiles and fishes; among them may be mentionedH. stepanovi(fig. 6), fromEmysandCistudo, whose sexual-cycle in a leech has been worked out by Siegel (see above),H. delagei, fromRaja,H. bigemina, from blennies, andH. simondi, from soles. Recently one or two Mammalian forms have been observed,H. gerbilli, from an Indian rat (Gerbillus), andH. jaculi, from the jerboa.GenusLankesterella, Labbé (syn.Drepanidium, Lankester). The parasite is not more than three-quarters the length of the corpuscle.L. ranarumfromRanais the type-species; another, recently described by Fantham, isL. tritonis, from the newt.From Lankester’sTreatise on Zoology.Fig. 7.—Karyolysus lacertarum(Danil.), in the blood-corpuscles of Lacerta muralis, showing the effects of the parasite upon the nucleus of the corpuscle. Incanddthe nucleus is broken up. N, Nucleus of the corpuscle;n, nucleus of the parasite, seen as a number of masses of chromatin, not enclosed by a distinct membrane. (After Marceau.)GenusKaryolysus, Labbé. The parasite does not exceed the corpuscle in length; the forms included in this genus, moreover, although not actually intranuclear, have a marked karyolytic and disintegrating action upon the nucleus of the corpuscle. The type-species is the well-knownK. lacertarum, of lizards; another isK. (Haemogregarina) viperini, fromTropidonotus.In the section of thePiroplasmatathere is only the genusPiroplasma, Patton (synn.Babesia, Starcovici,Pyrosoma, Smith and Kilborne), the principal species of which are as follows:P. bigeminum, the cause of Texas cattle-fever, tick-fever (Rinder-malaria) of South Africa, andP. bovis, causing haemoglobinuria of cattle in Southern Europe; there is some uncertainty as to whether these two are really distinct;P. canis,P. ovisandP. equiassociated, respectively, with those animals. Lately, a very small form,P. parvum, has been described by Theiler in Rhodesia, which causes East-African coast-fever; and another,P. muris, has been observed in white rats by Fantham.Bibliography.—(The older literature is enumerated in most treatises on Sporozoa—see bibliography underSporozoa). P. Argutinsky, “Malariastudien,”Arch. mikr. Anat.59, p. 315, pls. 18-21 (1901), andop. cit.61, p. 331, pl. 18 (1902); A. Balfour, “Haemogregarine of Mammals,”J. Trop. Med.8, p. 241, 8 figs. (1905); C. A. Bentley, “Leucocytozoan of the Dog,”B.M.J.(1905), 1, pp. 988 and 1078; N. Berestneff, “Über einen neuen Blutparasiten der indischen Frösche,”Arch. Protistenk.2, p. 343, pl. 8 (1903); “Über das’Leucocytozoan’ danilewskyi,”op. cit.3, p. 376, pl. 15 (1904); A. Billet, “Contribution à l’étude du paludisme et de son hématozoaire en Algérie,”Ann. Inst. Pasteur, 16, p. 186 (1902); (Notes on various Haemogregarines).C. R. Soc. Biol.56, pp. 482, 484, 607 and 741 (1904); C. Börner, “Untersuchungen über Hämosporidien,”Zeitschr. wiss. Zool.69, p. 398, 1 pl. (1901); T. Bowhill, “Equine piroplasmosis,” &c.,J. Hyg.5, p. 7, pls. 1-3 (1905); Bowhill and C. le Doux, “Contribution to the Study of ’Piroplasmosis canis,’”op. cit.4, p. 217, pl. 11 (1904); E. Brumpt and C. Lebailly, “Description de quelques nouvelles espèces de trypanosomes et d’hémogrégarines,” &c., C. R. Ac. Sci. 139, p. 613 (1904); A. Castellani and A. Willey, “Observations on the Haematozoa of Vertebrates in Ceylon,”Spolia Zeylan. 2, p. 78, 1 pl. (1904), andQ. J. Micr. Sci.49, p. 383, pl. 24 (1905); S. R. Christophers, “Haemogregarina gerbilli,”Sci. Mem. India, 18, 15 pp., 1 pl. (1905); H. B. Fantham, “Lankesterella tritonis, n. sp.,” &c.,Zool. Anz.29, p. 257, 17 figs. (1905); “Piroplasma muris,” &c.,Q. J. Micr. Sci.50, p. 493, pl. 28 (1906); C. Graham-Smith, “A new Form of Parasite found in the Red Blood-Corpuscles of Moles,”J. Hyg.5, p. 453, pls. 13 and 14 (1905); R. Hintze, “Lebensweise und Entwickelung vonLankesterella minima,”Zool. Jahrb. Anat.15, p. 693, pl. 36 (1902); S. James, “On a Parasite found in the White Blood-Corpuscles of Dogs,”Sci. Mem. India, 14, 12 pp. 1 pl. (1905); R. Koch, “Vorläufige Mitteilungen über die Ergebnisse einer Forschungsreise nach Ostafrika,”Deutsch. med. Wochenschr., 1905, p. 1865, 24 figs.; A. Labbé, “Recherches sur les parasites endoglobulaires du sang des vertébrés,”Arch. zool. exp.(3) ii. p. 55, 10 pls. (1894); A. Laveran, “Sur quelques hémogrégarines des ophidiens,”C. R. Ac. Sci.135, p. 1036, 13 figs. (1902); “Sur uneHaemamoebad’une mésange (Parus major),”C. R. Soc. Biol.54, p. 1121, 10 figs. (1902); “Sur la piroplasmose bovine bacilliforme,”C. R. Ac. Sci.138, p. 648, 18 figs. (1903); “Contribution à l’étude deHaemamoeba ziemanni,”C. R. Soc. Biol.55, p. 620, 7 figs. (1903); “Sur une hémogrégarine des gerboises,”C. R. Ac. Sci.141, p. 295, 9 figs. (1905); (On different Haemogregarines)C. R. Soc. Biol.59, pp. 175, 176, with figs. (1905); “Haemocytozoa. Essai de classification,”Bull. Inst. Pasteur, 3, p. 809 (1905); Laveran and F. Mesnil, “Sur les hématozoaires des poissons marins,”C. R. Ac. Sci.135, p. 567 (1902); “Sur quelques protozoaires parasites d’une tortue d’Asie,”t.c.p. 609, 14 figs. (1902); Laveran and Nègre, “Sur un protozoaire parasite deHyalomma aegyptium,”C. R. Soc. Biol.58, p. 964, 6 figs. (1905); (for various earlier papers by these authors, reference should be made to the C. R. Ac. Sci. and C R. Soc. Biol. for previous years); C. Lebailly (On Piscine Haemogregarines)C. R. Ac. Sci.139, p. 576 (1904), andC. R. Soc. Biol.59, p. 304 (1905); J. Lignières, “Sur la ‘Tristeza,’”Ann. Inst. Pasteur, 15, p. 121, pl. 6 (1901); “La Piroplasmose bovine; nouvelles recherches,” &c.,Arch. parasit.7, p. 398, pl. 4 (1903); M. Lühe, “Die im Blute schmarotzenden Protozoen,” in Mense’sHandbuch der Tropenkrankheiten(Leipzig, 1906), 3, 1; F. Marceau, “Note sur leKaryolysus lacertarum,”Arch. parasitol.4, p. 135, 46 figs. (1901); W. MacCallum, “On the Haematozoan Infection of Birds,”J. Exp. Med.3, p. 117, pl. 12 (1898); G. Mauser, “Die Malaria perniciosa,”Centrbl. Bakter.(1) 32, Orig. p. 695, 3 pls. (1902); C. Nicolle (On various Reptilian Haemogregarines),C. R. Soc. Biol.56, pp. 330, 608 and 912, with figs. (1904); Nicolle and C. Comte, “Sur le rôle ... deHyalomma... dans l’infection hémogrégarinienne,”op. cit.58, p. 1045 (1905); Norcard and Motas, “Contribution à l’étude de la piroplasmose canine,”Ann. Inst. Pasteur, 16, p. 256, pls. 5 and 6 (1902); G. Nuttall and G. Graham-Smith, “Canine piroplasmosis,”J. Hygiene, p. 237, pl. 9 (1905); F. Schaudinn, “Der Generationswechsel der Coccidien und Hämosporidien,”Zool. Centrbl.6, p. 675 (1899); “Studien über krankheitserregende Protozoen—II.Plasmodium vivax,”Arb. Kais. Gesundheitsamte, 19, p. 169, pls. 4-6 (1902); E. and E. Sergent (On different Haemogregarines),C. R. Soc. Biol.56, pp. 130, 132 (1904),op. cit.58, pp. 56, 57, 670 (1905); J. Siegel, “Die geschlechtliche Entwickelung vonHaemogregarina,” &c.,Arch. Protistenk.2, p. 339, 7 figs. (1903); P. L. Simond, “Contribution à l’étude des hématozoaires endoglobulaires des reptiles,”Ann. Inst. Pasteur, 15, p. 319, 1 pl. (1901); T. Smith and F. Kilborne, “Investigations into the Nature, Causation and Prevention of Texas Cattle Fever,”Rep. Bureau Animal Industry, U.S.A., 9 and 10, p. 177, pls. (1893); A. Theiler, “ThePiroplasma bigeminumof the Immune Ox,”J. Army Med. Corps, 3, pp. 469, 599, 1 pl. (1904); J. Vassal, “Sur une hématozoaire endoglobulaire nouveau d’un mammifère,”Ann. Inst. Pasteur, 19, p. 224, pl. 10 (1905); L. B. Wilson and W. Chowning, “Studies inPiroplasmosis hominis,”J. Infect. Diseases, 1, p. 31, 2 pls. (1904).
Various types of form are to be met with among the Haemosporidia. In one, characteristic of most (though not of absolutely all) parasites of warm-blooded Vertebrates, the trophozoites are of irregular amoeboid shape; hence this sectionComparative Morphology; variations in the life-cycle where known.is generally known as theHaemamoebidae. In another type, characteristic of the parasites of cold-blooded Vertebrates, the body possesses a definite, vermiform,i.e.gregariniform shape, which is retained during the intracorpuscular as well as during the free condition; this section comprises theHaemogregarinidae. Allied to this latter type of form are the trophozoites ofPiroplasma, which are normally pear-shaped; they differ, however, in being very minute, and, moreover, exhibit considerable polymorphism, rod-like (so-called bacillary) and ring-forms being of common occurrence. It is important to note that in a certain species ofHaemogregarina(fig. 3) the young trophozoites markedly resemblePiroplasmain their pyriform appearance; and a further point of agreement between the two forms is mentioned below. Lastly there is the Avian genusHalteridium, the trophozoites of which are characteristically bean-shaped or reniform. True Haemogregarines also differ in other slight points from “Haemamoebae.” Thus the young endoglobular trophozoite does not exhibit a ring (vacuolar) phase; and the cytoplasm never contains, at any period, the characteristic melanin pigment above noted. In some species ofHaemogregarinathe parasite, while intracorpuscular, becomes surrounded by a delicate membrane, the cytocyst; on entering upon an active, “free” period, the cytocyst is ruptured and left behind with the remains of the corpuscle. A very interesting cytological feature is the occurrence, in one or two Haemosporidia, of nuclear dimorphism,i.e.of a larger and smaller chromatic body, probably comparable to the trophic and kinetic nuclei of a Trypanosome, or of the “Leishman-Donovan” bodies. Schaudinn was the first to notice this character, inPiroplasma canis, and his observation has since been confirmed by Lühe.4Moreover, Brumpt has also noticed nuclear dimorphism in the ookinete of a species ofHaemogregarinain a leech (as the Invertebrate host)—a highly important observation.
As regards the life-history, the endogenous (schizogonous) cycle is known in many cases. Sometimes schizogony takes the primitive form of simple binary (probably) longitudinal fission; this is the case inPiroplasma(fig. 4) and also inHaemogregarina bigeminajust referred to. From this result the pairs of individuals (“twins”) so often found in the corpuscles. In addition, however, at any rate inPiroplasma, it is probable that multiple division (more allied to ordinary schizogony) also takes place; such is the case, according to Laveran, inP. equi, and the occurrence at times of four parasites in a corpuscle, arranged in a cruciform manner, is most likely to be thus explained. Labbé has described schizogony inHalteridium danilewskyias taking place in a rather peculiar manner; the parasite becomes much drawn-out and halter-like, and the actual division is restricted to its two ends, two clumps of merozoites being formed, at first connected by a narrow strand of unused cytoplasm, which subsequently disappears. Some doubt, however, attaches to this account, as no one else appears to have seen the process. For the rest, schizogony takes place more or less in the customary way, allowing for variations in the mode of arrangement of the merozoites. It remains to be noted that inKaryolysus lacertarum, according to Labbé, two kinds of schizont are developed, which give rise, respectively, to micromerozoites and megamerozoites, in either case enclosed in a delicate cytocyst. This probably corresponds to an early sexual differentiation (such as is found among certain Coccidia (q.v.), the micromerozoites producing eventually micro-gametocytes, the others megagametocytes.
a, Youngest form.
b, Slightly older.
candd. Division of the nucleus.
eandf, Division of the body of the parasite.
g,h,i,j, Various forms of the twin parasite.
kandl, Doubly infected corpuscles.
It has now been recognized for some time that the sexual (exogenous) part of the life-cycle of all theHaemamoebidaetakes place in an Invertebrate (Insectan) host, and is fundamentally similar to that above described in those cases where it has been followed. In contradistinction to the malarial parasites, this host, in the Avian forms (HaemoproteusandHalteridium)5is a species ofCulexand not ofAnopheles; in other words, gamete-formation, conjugation and subsequent sporozoite-formation in these cases will only go on in the former. On the other hand, in the case of the Haemogregarines, it was thought until quite lately that the entire life-history, including conjugation and sporogony, went on in the Vertebrate host; and only in 1902 Hintze described what purported to be the complete life-history ofLankesterella(Drepanidium)ranarumundergone in the frog. This view was rendered obsolete by the work of Siegel and Schaudinn, who demonstrated the occurrence of an alternation of hosts and of generations in the case ofHaemogregarina stepanovi, parasitic in a tortoise, and inKaryolysus lacertarum; the Invertebrate hosts, in which, in both cases, the sexual process is undergone, being respectively a leech (Placobdella) and a tick (Ixodes). With this discovery the main distinction (as supposed) between the Haemosporidia of warm and of cold-blooded Vertebrates vanished. It was further acknowledged by Schaudinn (under whom Hintzehad worked) that the latter had been misled by Coccidian cysts and spores, which he took for those ofLankesterella. The gametogony and sporogony ofHaemogregarina stepanoviin the leech agree in essential particulars with the process above described. The microgametes are extremely minute, and the sporozoites, which are developed in the salivary glands, where the motile ookinetes finally come to rest, are extremely “spirochaetiform”—the full significance of this latter fact being, perhaps, not appreciated.
Christophers recently described some remarkable phases which he regarded as belonging to the cycle ofHaemogregarina gerbilli(one of the few Mammalian Haemogregarines known) in a louse (Haematopinus). In a private communication, however, the author states that he has probably mistaken phases in the development of an ordinary gregarine parasite in the louse for part of the life-cycle of this Haemogregarine.
The Mammalian parasitePiroplasmais the one about whose life-history our knowledge is most vague. Besides the typical and generally occurring forms, others have also been observed in the blood, but it is doubtful how far these are to be looked upon as normal; for instance, Bowhill and Le Doux have described, in various species, a phase in which a long, slender pseudopodial-like outgrowth is present, with a swelling at the distal end. It is, moreover, quite uncertain which are the sexual forms, comparable to gametocytes. Doflein regards large pear-shaped forms as such (megagametocytes?), which become spherical when maturing; and Nocard and Motas have figured amoeboid, irregular forms, with the nucleus fragmented and possessing flagella-like processes (possibly microgametes?). The Invertebrate host is well known to be, in the case of all species, a tick; thus bovine piroplasmosis (P. bigeminum) in America is conveyed byRhipicephalus annulatus(Boophilus bovis), canine piroplasmosis (P. canis) in South Africa byHaemaphysalis leachi(and perhapsDermacentor reticulatus), and so on. The manner in which the infection is transmitted by the tick varies greatly. In some cases (e.g.P. bigeminumandP. canis) only the generation subsequent to that which receives the infection (by feeding on an infected ox) can transmit it back again to another ox; in other words, true hereditary infection of the ova in the mother-tick is found to occur. The actual period in the life of the daughter-tick at which it can convey the infection apparently varies. On the other hand, in the case of East African coast-fever, Theiler found that hereditary infection does not occur, the same generation transmitting the parasite (P. parvum) at different periods of life. Little is certainly known regarding the phases of the parasite which are passed through in the tick. Lignières has observed a kind of multiple fission in the stomach, several very minute bodies, consisting mostly of chromatin, being formed, which may serve for endogenous reproduction. Koch has published an account of certain curious forms ofP. bigeminum, in which the body is produced into many stiff, ray-like processes, giving the appearance of a star; according to him fusion of such forms takes place, and the resulting zygote becomes rounded, perhaps transitional to the pear-shaped forms.
The classification and nomenclature of the Haemosporidia are in a very unsettled condition. For an account of the various systems and modifications hitherto adopted, the article of Minchin (see underSporozoa:Bibliography) should be consulted.Classification.With the realization that the life-history in the case of the “Haemamoebae” and the Haemogregarines is fundamentally similar in type, the chief reason for grouping them as distinct suborders has disappeared. It is most convenient to regard them as separate, but closely allied families, thePlasmodidae(“Haemamoebidae”) and theHaemogregarinidae. ThePiroplasmata, on the other hand, constitute another family, which is better placed in a distinct section or sub-order. In addition there are, as already noted, two or three genera whose systematic position must be considered as quite uncertain. One is the well-knownHalteridiumof Labbé, parasitic in various birds; the type-species isH. danilewskyi(Gt. and Fel.). Another is the much-debated parasite of white blood-corpuscles (leucocytes), originally described in birds by Danilewsky under the name ofLeucocytozoon, a form of which has been recently observed in Mammals.
In conclusion, the chief members of the above-mentioned families may be enumerated.
Fam.Plasmodidae(“Haemamoebidae”).
GenusLaverania, Gr. and Fel. (syn.Haemamoenas, Ross), for L.malariae, Gr. and Fel. (synn. L. s.Plasmodium, s. “Haemamoeba,” &c.,praecoxs.immaculatum, &c.), the parasite of pernicious malaria. GenusPlasmodium, March. and Celli (syn. “Haemamoeba”) forP. vivaxandP. malariae, the tertian and quartan parasite, respectively. There is also a form known in apes,P. kochi. GenusHaemoproteus, Kruse (syn.Proteosoma), forH. danilewskyi(syn.Proteosoma grassi,Plasmodium praecox, &c.), parasitic in numerous birds. Recently, another form has been described, from reptiles, which Castellani and Willey have termedHaemocystidium simondi.
Remarks.—The distinguishing characters of the malarial parasites have been mentioned above. Some authorities would includeLaveraniain the genusPlasmodium, as differing only specifically from the other two forms. It has, moreover, been suggested by Sergent that all three are merely different phases of the same parasite, predominating at different seasons; this idea cannot be regarded, however, as in any way proved so far. From what is known of the morphology and mode of manifestation of these forms, the differences betweenLaveraniaand the two species ofPlasmodiumare considerably more pronounced than those betweenP. vivaxandP. malariae; if the latter are to be considered as distinct species, the first-named is probably generically distinct. Lühe, it may be noted, in his recent comprehensive account of the Haematozoa, also takes this view. Lastly, whatever be the correct solution of the above problem, there is certainly not sufficient justification for including the Avian genusHaemoproteus, as also only a species ofPlasmodium, which is done by some. Its different Vertebrate habitat, and also the fact that its Insectan definitive host is Culex and notAnopheles, differentiate it sharply fromLaveraniaandPlasmodium.
a, Young trophozoite in a blood-corpuscle,
bandc, Older trophozoite.
dande, Sporulation.
d, Precocious sporulation with few merozoites.
e, Sporulation of a full-grown schizont, with numerous merozoites.
f, Gametocyte.
N, Nucleus of blood-corpuscle.
n, Nucleus of parasite.
p, Pigment.
mz, Merozoites.
r.p, Residual protoplasm.
a, Blood-corpuscle with young trophozoite.
b, Older trophozoite.
c, Full-grown trophozoite, ready to leave the corpuscle.
dande, Trophozoites free in the blood-plasma, showing changes of form.
f-i, Trophozoites, still within the blood-corpuscle (not drawn), showing the structure of the nucleus, the coarse chromatoid granules in the protoplasm and the manner in which the parasite grows into theU-shaped Haemogregarine without increase of body-mass.
j, Commencement of sporulation; the nucleus has divided into eight nuclei, and the body of the parasite is beginning to divide up into as many merozoites within a blood-corpuscle.
N, Nucleus of the blood-corpuscle.
n, Nucleus of the parasite.
Fam.Haemogregarinidae.—The different genera are characterized chiefly by their size relative to the blood-corpuscles, and their disposition in the latter. Here, again, it has been suggested to unite the various types all in one genus,Haemogregarina, but this seems at least premature when it is remembered how little is known in most cases of the life-cycle, which may prove to exhibit important divergences.
GenusHaemogregarina, Danilewsky (syn.Danilewskya, Labbé).The body of the parasite exceeds the blood-corpuscle in length, when adult, and is bent upon itself, like aU. A very great number of species are known, mostly from reptiles and fishes; among them may be mentionedH. stepanovi(fig. 6), fromEmysandCistudo, whose sexual-cycle in a leech has been worked out by Siegel (see above),H. delagei, fromRaja,H. bigemina, from blennies, andH. simondi, from soles. Recently one or two Mammalian forms have been observed,H. gerbilli, from an Indian rat (Gerbillus), andH. jaculi, from the jerboa.
GenusLankesterella, Labbé (syn.Drepanidium, Lankester). The parasite is not more than three-quarters the length of the corpuscle.L. ranarumfromRanais the type-species; another, recently described by Fantham, isL. tritonis, from the newt.
GenusKaryolysus, Labbé. The parasite does not exceed the corpuscle in length; the forms included in this genus, moreover, although not actually intranuclear, have a marked karyolytic and disintegrating action upon the nucleus of the corpuscle. The type-species is the well-knownK. lacertarum, of lizards; another isK. (Haemogregarina) viperini, fromTropidonotus.
In the section of thePiroplasmatathere is only the genusPiroplasma, Patton (synn.Babesia, Starcovici,Pyrosoma, Smith and Kilborne), the principal species of which are as follows:P. bigeminum, the cause of Texas cattle-fever, tick-fever (Rinder-malaria) of South Africa, andP. bovis, causing haemoglobinuria of cattle in Southern Europe; there is some uncertainty as to whether these two are really distinct;P. canis,P. ovisandP. equiassociated, respectively, with those animals. Lately, a very small form,P. parvum, has been described by Theiler in Rhodesia, which causes East-African coast-fever; and another,P. muris, has been observed in white rats by Fantham.
Bibliography.—(The older literature is enumerated in most treatises on Sporozoa—see bibliography underSporozoa). P. Argutinsky, “Malariastudien,”Arch. mikr. Anat.59, p. 315, pls. 18-21 (1901), andop. cit.61, p. 331, pl. 18 (1902); A. Balfour, “Haemogregarine of Mammals,”J. Trop. Med.8, p. 241, 8 figs. (1905); C. A. Bentley, “Leucocytozoan of the Dog,”B.M.J.(1905), 1, pp. 988 and 1078; N. Berestneff, “Über einen neuen Blutparasiten der indischen Frösche,”Arch. Protistenk.2, p. 343, pl. 8 (1903); “Über das’Leucocytozoan’ danilewskyi,”op. cit.3, p. 376, pl. 15 (1904); A. Billet, “Contribution à l’étude du paludisme et de son hématozoaire en Algérie,”Ann. Inst. Pasteur, 16, p. 186 (1902); (Notes on various Haemogregarines).C. R. Soc. Biol.56, pp. 482, 484, 607 and 741 (1904); C. Börner, “Untersuchungen über Hämosporidien,”Zeitschr. wiss. Zool.69, p. 398, 1 pl. (1901); T. Bowhill, “Equine piroplasmosis,” &c.,J. Hyg.5, p. 7, pls. 1-3 (1905); Bowhill and C. le Doux, “Contribution to the Study of ’Piroplasmosis canis,’”op. cit.4, p. 217, pl. 11 (1904); E. Brumpt and C. Lebailly, “Description de quelques nouvelles espèces de trypanosomes et d’hémogrégarines,” &c., C. R. Ac. Sci. 139, p. 613 (1904); A. Castellani and A. Willey, “Observations on the Haematozoa of Vertebrates in Ceylon,”Spolia Zeylan. 2, p. 78, 1 pl. (1904), andQ. J. Micr. Sci.49, p. 383, pl. 24 (1905); S. R. Christophers, “Haemogregarina gerbilli,”Sci. Mem. India, 18, 15 pp., 1 pl. (1905); H. B. Fantham, “Lankesterella tritonis, n. sp.,” &c.,Zool. Anz.29, p. 257, 17 figs. (1905); “Piroplasma muris,” &c.,Q. J. Micr. Sci.50, p. 493, pl. 28 (1906); C. Graham-Smith, “A new Form of Parasite found in the Red Blood-Corpuscles of Moles,”J. Hyg.5, p. 453, pls. 13 and 14 (1905); R. Hintze, “Lebensweise und Entwickelung vonLankesterella minima,”Zool. Jahrb. Anat.15, p. 693, pl. 36 (1902); S. James, “On a Parasite found in the White Blood-Corpuscles of Dogs,”Sci. Mem. India, 14, 12 pp. 1 pl. (1905); R. Koch, “Vorläufige Mitteilungen über die Ergebnisse einer Forschungsreise nach Ostafrika,”Deutsch. med. Wochenschr., 1905, p. 1865, 24 figs.; A. Labbé, “Recherches sur les parasites endoglobulaires du sang des vertébrés,”Arch. zool. exp.(3) ii. p. 55, 10 pls. (1894); A. Laveran, “Sur quelques hémogrégarines des ophidiens,”C. R. Ac. Sci.135, p. 1036, 13 figs. (1902); “Sur uneHaemamoebad’une mésange (Parus major),”C. R. Soc. Biol.54, p. 1121, 10 figs. (1902); “Sur la piroplasmose bovine bacilliforme,”C. R. Ac. Sci.138, p. 648, 18 figs. (1903); “Contribution à l’étude deHaemamoeba ziemanni,”C. R. Soc. Biol.55, p. 620, 7 figs. (1903); “Sur une hémogrégarine des gerboises,”C. R. Ac. Sci.141, p. 295, 9 figs. (1905); (On different Haemogregarines)C. R. Soc. Biol.59, pp. 175, 176, with figs. (1905); “Haemocytozoa. Essai de classification,”Bull. Inst. Pasteur, 3, p. 809 (1905); Laveran and F. Mesnil, “Sur les hématozoaires des poissons marins,”C. R. Ac. Sci.135, p. 567 (1902); “Sur quelques protozoaires parasites d’une tortue d’Asie,”t.c.p. 609, 14 figs. (1902); Laveran and Nègre, “Sur un protozoaire parasite deHyalomma aegyptium,”C. R. Soc. Biol.58, p. 964, 6 figs. (1905); (for various earlier papers by these authors, reference should be made to the C. R. Ac. Sci. and C R. Soc. Biol. for previous years); C. Lebailly (On Piscine Haemogregarines)C. R. Ac. Sci.139, p. 576 (1904), andC. R. Soc. Biol.59, p. 304 (1905); J. Lignières, “Sur la ‘Tristeza,’”Ann. Inst. Pasteur, 15, p. 121, pl. 6 (1901); “La Piroplasmose bovine; nouvelles recherches,” &c.,Arch. parasit.7, p. 398, pl. 4 (1903); M. Lühe, “Die im Blute schmarotzenden Protozoen,” in Mense’sHandbuch der Tropenkrankheiten(Leipzig, 1906), 3, 1; F. Marceau, “Note sur leKaryolysus lacertarum,”Arch. parasitol.4, p. 135, 46 figs. (1901); W. MacCallum, “On the Haematozoan Infection of Birds,”J. Exp. Med.3, p. 117, pl. 12 (1898); G. Mauser, “Die Malaria perniciosa,”Centrbl. Bakter.(1) 32, Orig. p. 695, 3 pls. (1902); C. Nicolle (On various Reptilian Haemogregarines),C. R. Soc. Biol.56, pp. 330, 608 and 912, with figs. (1904); Nicolle and C. Comte, “Sur le rôle ... deHyalomma... dans l’infection hémogrégarinienne,”op. cit.58, p. 1045 (1905); Norcard and Motas, “Contribution à l’étude de la piroplasmose canine,”Ann. Inst. Pasteur, 16, p. 256, pls. 5 and 6 (1902); G. Nuttall and G. Graham-Smith, “Canine piroplasmosis,”J. Hygiene, p. 237, pl. 9 (1905); F. Schaudinn, “Der Generationswechsel der Coccidien und Hämosporidien,”Zool. Centrbl.6, p. 675 (1899); “Studien über krankheitserregende Protozoen—II.Plasmodium vivax,”Arb. Kais. Gesundheitsamte, 19, p. 169, pls. 4-6 (1902); E. and E. Sergent (On different Haemogregarines),C. R. Soc. Biol.56, pp. 130, 132 (1904),op. cit.58, pp. 56, 57, 670 (1905); J. Siegel, “Die geschlechtliche Entwickelung vonHaemogregarina,” &c.,Arch. Protistenk.2, p. 339, 7 figs. (1903); P. L. Simond, “Contribution à l’étude des hématozoaires endoglobulaires des reptiles,”Ann. Inst. Pasteur, 15, p. 319, 1 pl. (1901); T. Smith and F. Kilborne, “Investigations into the Nature, Causation and Prevention of Texas Cattle Fever,”Rep. Bureau Animal Industry, U.S.A., 9 and 10, p. 177, pls. (1893); A. Theiler, “ThePiroplasma bigeminumof the Immune Ox,”J. Army Med. Corps, 3, pp. 469, 599, 1 pl. (1904); J. Vassal, “Sur une hématozoaire endoglobulaire nouveau d’un mammifère,”Ann. Inst. Pasteur, 19, p. 224, pl. 10 (1905); L. B. Wilson and W. Chowning, “Studies inPiroplasmosis hominis,”J. Infect. Diseases, 1, p. 31, 2 pls. (1904).
(H. M. Wo.)
1Compare, for example, the flagellated granules of certain Coccidia, which point unmistakably to a Flagellate ancestry.2A possible exception is a doubtful species ofHaemogregarina, which has been described from the walls of the blood-vessels of an Annelid.3For an interesting account of the biological relations between parasites and their hosts, and the penalty Man pays for his roving propensities, the reader should see Lankester’s article in theQuarterly Review, July 1904.4This does away with one of the principal reasons on account of which some authorities considerPiroplasma(Leishmania)donovanias quite distinct from otherPiroplasmata(seeTrypanosomes).5It must not be forgotten that one species ofHalteridium(H.[Trypanomorpha]noctuae) is said to have well-marked trypaniform phases in its life-cycle; these are preferably considered underTrypanosomes(q.v.), and therefore, to avoid repetition, are only thus alluded to here. WhetherH. danilewskyialso becomes trypaniform in certain phases, and how far it really agrees with the criteria of a Haemosporidian above postulated, are matters which are not yet definitely known.
1Compare, for example, the flagellated granules of certain Coccidia, which point unmistakably to a Flagellate ancestry.
2A possible exception is a doubtful species ofHaemogregarina, which has been described from the walls of the blood-vessels of an Annelid.
3For an interesting account of the biological relations between parasites and their hosts, and the penalty Man pays for his roving propensities, the reader should see Lankester’s article in theQuarterly Review, July 1904.
4This does away with one of the principal reasons on account of which some authorities considerPiroplasma(Leishmania)donovanias quite distinct from otherPiroplasmata(seeTrypanosomes).
5It must not be forgotten that one species ofHalteridium(H.[Trypanomorpha]noctuae) is said to have well-marked trypaniform phases in its life-cycle; these are preferably considered underTrypanosomes(q.v.), and therefore, to avoid repetition, are only thus alluded to here. WhetherH. danilewskyialso becomes trypaniform in certain phases, and how far it really agrees with the criteria of a Haemosporidian above postulated, are matters which are not yet definitely known.
HAETZER,orHetzer,LUDWIG(d. 1529), Swiss divine, was born in Switzerland, at Bischofszell, in Thurgau. He studied at Freiburg-im-Breisgau, and began his career in a chaplaincy at Wadenswil, on the Lake of Zürich. At this time his attachment to the old faith was tempered by a mystical turn, and by a devotion to the prophetical writings of the Old Testament, which he studied in the original. By 1523 we find him in Zürich, where he published, at first anonymously and in Latin (Judicium Dei), later with his name and in German (Sept. 24, 1523), a small tract against the religious use of images, and bearing the motto attached to all his subsequent works, “O Got erlösz die (or dein) Gefangnen” (“O God, set the prisoners free”). An attempt to give effect to the teaching of this (frequently reprinted) tract was followed by a public religious disputation, of which Haetzer drew up the official account. In 1524 he brought out a tract on the conversion of the Jews, and published a German version of Johann Bugenhagen’s brief exposition of the epistles of St Paul (Ephesians to Hebrews); in the dedication (dated Zürich, June 29, 1524) he undertakes to translate Bugenhagen’s comment on the Psalter. He then went to Augsburg, bearing Zwingli’s introduction to Johann Frosch. Here he came for a time under the influence of Urbanus Regius, and was for a short time the guest of Georg Regel. Returning to Zürich, he was in intercourse with leading Anabaptists (though his own position was simply the disuse of infant baptism) till their expulsion in January 1525. Again resorting to Augsburg, and resuming work as corrector of the press for his printer Silvan Ottmar, he pushed his views to the extreme of rejecting all sacraments, reaching something like the mystical standpoint of the early Quakers. He was expelled from Augsburg in the autumn of 1525, and made his way through Constance to Basel, where Oecolampadius received him kindly. He translated into German the first treatise of Oecolampadius on the Lord’s Supper (in which the words of institution are taken figuratively), and proceeding to Zürich in November, publishedhis version there in February 1526, with a preface disclaiming connexion with the Anabaptists. His relations with Zwingli were difficult; returning to Basel he published (July 18, 1526) his translation of Malachi, with Oecolampadius’s exposition, and with a preface reflecting on Zwingli. This he followed by a version of Isaiah xxxvi.-xxxvii. He next went to Strassburg, and was received by Wolfgang Capito. At Strassburg in the late autumn of 1526 he fell in with Hans Dengk or Denck, who collaborated with him in the production of hisopus magnum, the translation of the Hebrew Prophets,Alle Propheten nach hebraischer Sprach vertuetscht. The preface is dated Worms, 3 April 1527; and there are editions, Worms, 13 April 1527, folio; Augsburg, 22 June 1527, folio; Worms, 7 Sept. 1527, 16º; and Augsburg, 1528, folio. It was the first Protestant version of the prophets in German, preceding Luther’s by five years, and highly spoken of by him. Haetzer and Denck now entered on a propagandist mission from place to place, with some success, but of short duration. Denck died at Basel in November 1527. Haetzer was arrested at Constance in the summer of 1528. After long imprisonment and many examinations he was condemned on the 3rd of February 1529 to die by the sword, and the sentence was executed on the following day. His demeanour on the scaffold impressed impartial witnesses, Hans Zwick and Thomas Blaurer, who speak warmly of his fervour and courage. The Dutch Baptist Martyrology describes him as “a servant of Jesus Christ.” The Moravian Chronicle says “he was condemned for the sake of divine truth.” His papers included an unpublished treatise against the essential deity of Christ, which was suppressed by Zwingli; the only extant evidence of his anti-trinitarian views being contained in eight quaint lines of German verse preserved in Sebastian Frank’sChronica. The discovery of his heterodox Christology (which has led modern Unitarians to regard him as their proto-martyr) was followed by charges of loose living, never heard of in his lifetime, and destitute of evidence or probability.
See Breitinger, “Anecdota quaedam de L. H.” inMuseum Helveticum(1746), parts 21 and 23; Wallace,Antitrinitarian Biography(1850);Dutch Martyrology(Hanserd Knollys Society) (1856); Th. Keim, in Hauck’sRealencyklopädie(1899).
See Breitinger, “Anecdota quaedam de L. H.” inMuseum Helveticum(1746), parts 21 and 23; Wallace,Antitrinitarian Biography(1850);Dutch Martyrology(Hanserd Knollys Society) (1856); Th. Keim, in Hauck’sRealencyklopädie(1899).
(A. Go.*)
HĀFIZ.Shams-ud-din Mahommed, better known by histakhallusornom de plumeof Hāfiz, was one of the most celebrated writers of Persian lyrical poetry. He was born at Shiraz, the capital of Fars, in the early part of the 8th century of the Mahommedan era, that is to say, in the 14th of our own. The exact date of his birth is uncertain, but he attained a ripe old age and died in 791A.H.(A.D.1388). This is the date given in the chronogram which is engraved on his tomb, although several Persian biographers give a different year. Very little is actually known about his life, which appears to have been passed in retirement in Shiraz, of which he always speaks in terms of affectionate admiration. He was a subject of the Muzaffar princes, who ruled in Shiraz, Yazd, Kirman and Ispahan, until the dynasty was overthrown by Timur (Tamerlane). Of these princes his especial patrons were Shah Shujā’ and Shah Mansūr. He early devoted himself to the study of poetry and theology, and also became learned in mystic philosophy, which he studied under Shaik Mahmūd ‘Aṭṭār, chief of an order of dervishes. Hāfiz afterwards enrolled himself in the same order and became a professor of Koranic exegesis in a college which his friend and patron Haji Kiwam-ud-din, the vizier, specially founded for him. This was probably the reason of his adopting the sobriquet of Hāfiz (“one who remembers”), which is technically applied to any person who has learned the Koran by heart. The restraints of an ascetic life seem to have been very little to Hāfiz’s taste, and his loose conduct and wine-bibbing propensities drew upon him the severe censure of his monastic colleagues. In revenge he satirizes them unmercifully in his verses, and seldom loses an opportunity of alluding to their hypocrisy. Hāfiz’s fame as a poet was soon rapidly spread throughout the Mahommedan world, and several powerful monarchs sent him presents and pressing invitations to visit them. Amongst others he was invited by Mahmūd Shah Bahmani, who reigned in the south of India. After crossing the Indus and passing through Lahore he reached Hurmuz, and embarked on board a vessel sent for him by the Indian prince. He seems, however, to have been a bad sailor, and, having invented an excuse for being put ashore, made the best of his way back to Shiraz. Some biographies narrate a story of an interview between Hāfiz and the invader Timur. The latter sent for him and asked angrily, “Art thou he who was so bold as to offer my two great cities Samarkand and Bokhara for the black mole on thy mistress’s cheek?” alluding to a well-known verse in one of his odes. “Yes, sire,” replied Hāfiz, “and it is by such acts of generosity that I have brought myself to such a state of destitution that I have now to solicit your bounty.” Timur was so pleased at his ready wit that he dismissed the poet with a handsome present. Unfortunately for the truth of this story Timur did not capture Shiraz tillA.D.1393, while the latest date that can be assigned to Hāfiz’s death is 1391. Of his private life little or nothing is known. One of his poems is said to record the death of his wife, another that of a favourite unmarried son, and several others speak of his love for a girl calledShākh i Nabat, “Sugar-cane branch,” and this is almost all of his personal history that can be gathered from his writings. He was, like most Persians, a Shi‘ite by religion, believing in the transmission of the office of Imām (head of the Moslem Church) in the family of Ali, cousin of the prophet, and rejecting theHadith(traditional sayings) of Mahomet, which form the Sunna or supplementary code of Mahommedan ceremonial law. One of his odes which contains a verse in praise of Ali is engraved on the poet’s tomb, but is omitted by Sudi, the Turkish editor and commentator, who was himself a rigid Sunnite. Hāfiz’s heretical opinions and dissipated life caused difficulties to be raised by the ecclesiastical authorities on his death as to his interment in consecrated ground. The question was at length settled by Hāfiz’s own works, which had then already begun to be used, as they are now throughout the East, for the purposes of divination, in the same manner as Virgil was employed in the middle ages for the divination calledSortes Virgilianae. Opening the book at random after pronouncing the customary formula asking for inspiration, the objectors hit upon the following verse—“Turn not away thy foot from the bier of Hāfiz, for though immersed in sin, he will be admitted into Paradise.” He was accordingly buried in the centre of a small cemetery at Shiraz, now included in an enclosure called the Hāfiziyeh.
His principal work is theDīwān, that is, a collection of short odes or sonnets calledghazals, and consisting of from five to sixteenbaitsor couplets each, all the couplets in each ode having the same rhyme in the last hemistich, and the last couplet always introducing the poet’s ownnom de plume. The whole of these are arranged in alphabetical order, an arrangement which certainly facilitates reference but makes it absolutely impossible to ascertain their chronological order, and therefore detracts from their value as a means of throwing light upon the growth and development of his genius or the incidents of his career. They are often held together by a very slender thread of continuous thought, and few editions agree exactly in the order of the couplets. Still, a careful study of them, especially from the point of view indicated by the Sufiistic system of philosophy, will always show that a single idea does run throughout the whole. The nature of these poems has been the subject of much discussion in the West, some scholars seeing in their anacreontic utterances nothing but sensuality and materialism, while others, following the Oriental school, maintain that they are wholly and entirely mystic and philosophic. Something between the two would probably be nearer the truth. It must be remembered that Hāfiz was a professed dervish and Sūfi, and that hisghazalswere in all probability published from atakia, and arranged with at least a view to Sufiistic interpretation. At the same time it is ridiculous to suppose that the glowing imagery, the gorgeous and often tender descriptions of natural beauties, the fervent love passages, and the roystering drinking songs were composed in cool blood or with deliberate ascetic purpose. The beauty of Hāfiz’s poetry is that it is natural. It is the outcomeof a fervent soul and a lofty genius delighting in nature and enjoying life; and it is the poet’s misfortune that he lived in an age and amongst a people where rigid conventionality demanded that his free and spontaneous thoughts should be recast in an artificial mould.
Besides theDīwān, Hāfiz wrote a number of other poems; the Leipzig edition of his works contains 573ghazals(forming theDīwān), 42kit‘asor fragments, 69ruba‘iyātor tetrastics, 6masnaviyātor poems in rhyming couplets, 2kasāïd, idylls or panegyrics, and 1mukhammesor poem in five-line strophes. Other editions contain severaltarji‘-bandor poems with a refrain. The wholeDīwānwas translated into English prose by H. Wilberforce Clarke in 1891, with introduction and exhaustive commentary and bibliography; a few rhyming versions of single poems by Sir William Jones, J. Nott, J. Hindley, Falconer, &c., are to be found scattered through the pages of theOriental Miscellanyand other periodicals, and a fine edition containing a verse rendering of the principal poems by H. Bicknell appeared in 1875. Other selections by S. Robinson (1875), A. Rogers (1889), J. H. M‘Carthy (1893), and Gertrude L. Bell (1897). The principal German versions are by von Hammer Purgstall (1812), which gave the first impulse to Goethe’sWestöstlicher Diwan; a rhyming and rhythmical translation of a large portion of Hāfiz’s works by Vincenz von Rosenzweig of Vienna (Vienna, 1858), which contains also the Persian text and notes;Der Diwan des Schemseddīn Muhammed Hāfis, by G. H. F. Nesselmann (Berlin, 1865), in which the rhyming system of the original is imitated. Besides these, the reader may consult d’Herbelot,Bibliothèque orientale, article “Hafiz”; Sir William Ouseley’sOriental Collections(1797-1798);A Specimen of Persian Poetry, or Odes of Hafiz, by John Richardson (London, 1802);Biographical Notices of Persian Poets, by Sir Gore Ouseley (Oriental Translation Fund, 1846); and an excellent article by Professor E. B. Cowell inMacmillan’s Magazine(No. 177, July 1874); J. A. Vullers,Vitae poëtarum Persicorum(1839, translated from Daulatshah); S. Robinson,Persian Poetry for English Readers(1883). The best edition of the text is perhaps that edited by Hermann Brockhaus of Leipzig (1854-1856). which is based on the recension of the Turkish editor Sudi, and contains his commentary in Turkish on the first eightyghazals. See also H. Ethé inGrundriss der iranischen Philologie, ii. (Strassburg, 1896); P. Horn,Geschichte der persischen Literatur(Leipzig, 1901).
Besides theDīwān, Hāfiz wrote a number of other poems; the Leipzig edition of his works contains 573ghazals(forming theDīwān), 42kit‘asor fragments, 69ruba‘iyātor tetrastics, 6masnaviyātor poems in rhyming couplets, 2kasāïd, idylls or panegyrics, and 1mukhammesor poem in five-line strophes. Other editions contain severaltarji‘-bandor poems with a refrain. The wholeDīwānwas translated into English prose by H. Wilberforce Clarke in 1891, with introduction and exhaustive commentary and bibliography; a few rhyming versions of single poems by Sir William Jones, J. Nott, J. Hindley, Falconer, &c., are to be found scattered through the pages of theOriental Miscellanyand other periodicals, and a fine edition containing a verse rendering of the principal poems by H. Bicknell appeared in 1875. Other selections by S. Robinson (1875), A. Rogers (1889), J. H. M‘Carthy (1893), and Gertrude L. Bell (1897). The principal German versions are by von Hammer Purgstall (1812), which gave the first impulse to Goethe’sWestöstlicher Diwan; a rhyming and rhythmical translation of a large portion of Hāfiz’s works by Vincenz von Rosenzweig of Vienna (Vienna, 1858), which contains also the Persian text and notes;Der Diwan des Schemseddīn Muhammed Hāfis, by G. H. F. Nesselmann (Berlin, 1865), in which the rhyming system of the original is imitated. Besides these, the reader may consult d’Herbelot,Bibliothèque orientale, article “Hafiz”; Sir William Ouseley’sOriental Collections(1797-1798);A Specimen of Persian Poetry, or Odes of Hafiz, by John Richardson (London, 1802);Biographical Notices of Persian Poets, by Sir Gore Ouseley (Oriental Translation Fund, 1846); and an excellent article by Professor E. B. Cowell inMacmillan’s Magazine(No. 177, July 1874); J. A. Vullers,Vitae poëtarum Persicorum(1839, translated from Daulatshah); S. Robinson,Persian Poetry for English Readers(1883). The best edition of the text is perhaps that edited by Hermann Brockhaus of Leipzig (1854-1856). which is based on the recension of the Turkish editor Sudi, and contains his commentary in Turkish on the first eightyghazals. See also H. Ethé inGrundriss der iranischen Philologie, ii. (Strassburg, 1896); P. Horn,Geschichte der persischen Literatur(Leipzig, 1901).
(E. H. P.)
HAG.(1) (Probably a shortened form of the O. Eng.hægtesse,hegtes, cognate with Ger.Hexe, witch, Dutchhecse), a word common during the 16th and 17th centuries for a female demon or evil spirit, and so particularly applied to such supernatural beings as the harpies and fairies of classical mythology, and also to witches. In modern usage the word is generally used of a hideous old woman whose repulsive exterior is accompanied by malice or wickedness. The name is also used of an eel-like parasitic fish,Myxine glutinosa, allied to the lamprey.
(2) A word common in Scottish and northern English dialects for an enclosed piece of wood, a copse. This is the same word as “hedge” (seeHedges) and “haw.” “Hag” also means “to cut,” and is used in Scotland of an extent of woodland marked out for felling, and of a quantity of felled wood. This word is also used of a cutting in the peat of a “moss” or “bog,” and hence applied to the small plots of firm ground or heather in a bog; it is common in the form “moss-hags.”
HAGEDORN, FRIEDRICH VON(1708-1754), German poet, was born on the 23rd of April 1708 at Hamburg, where his father, a man of scientific and literary taste, was Danish minister. He was educated at the gymnasium of Hamburg, and later (1726) became a student of law at Jena. Returning to Hamburg in 1729, he obtained the appointment of unpaid private secretary to the Danish ambassador in London, where he lived till 1731. Hagedorn’s return to Hamburg was followed by a period of great poverty and hardship, but in 1733 he was appointed secretary to the so-called “English Court” (Englischer Hof) in Hamburg, a trading company founded in the 13th century. He shortly afterwards married, and from this time had sufficient leisure to pursue his literary occupations till his death on the 28th of October 1754. Hagedorn is the first German poet who bears unmistakable testimony to the nation’s recovery from the devastation wrought by the Thirty Years’ War. He is eminently a social poet. His light and graceful love-songs and anacreontics, with their undisguisedjoie de vivre, introduced a new note into the German lyric; his fables and tales in verse are hardly inferior in form and in delicate persiflage to those of his master La Fontaine, and his moralizing poetry re-echoes the philosophy of Horace. He exerted a dominant influence on the German lyric until late in the 18th century.
The first collection of Hagedorn’s poems was published at Hamburg shortly after his return from Jena in 1729, under the titleVersuch einiger Gedichte(reprinted by A. Sauer, Heilbronn, 1883). In 1738 appearedVersuch in poetischen Fabeln und Erzählungen; in 1742 a collection of his lyric poems, under the titleSammlung neuer Oden und Lieder; and hisMoralische Gedichtein 1750. A collection of his entire works was published at Hamburg after his death in 1757. The best is J. J. Eschenburg’s edition (5 vols., Hamburg, 1800). Selections of his poetry with an excellent introduction in F. Muncker’sAnakreontiker und preussisch-patriotische Lyriker(Stuttgart, 1894). See also H. Schuster,F. von Hagedorn und seine Bedeutung für die deutsche Literatur(Leipzig, 1882); W. Eigenbrodt,Hagedorn und die Erzählung in Reimversen(Berlin, 1884).
The first collection of Hagedorn’s poems was published at Hamburg shortly after his return from Jena in 1729, under the titleVersuch einiger Gedichte(reprinted by A. Sauer, Heilbronn, 1883). In 1738 appearedVersuch in poetischen Fabeln und Erzählungen; in 1742 a collection of his lyric poems, under the titleSammlung neuer Oden und Lieder; and hisMoralische Gedichtein 1750. A collection of his entire works was published at Hamburg after his death in 1757. The best is J. J. Eschenburg’s edition (5 vols., Hamburg, 1800). Selections of his poetry with an excellent introduction in F. Muncker’sAnakreontiker und preussisch-patriotische Lyriker(Stuttgart, 1894). See also H. Schuster,F. von Hagedorn und seine Bedeutung für die deutsche Literatur(Leipzig, 1882); W. Eigenbrodt,Hagedorn und die Erzählung in Reimversen(Berlin, 1884).
HAGEN, FRIEDRICH HEINRICH VON DER(1780-1856), German philologist, chiefly distinguished for his researches in Old German literature, was born at Schmiedeberg In Brandenburg on the 19th of February 1780. After studying law at the university of Halle, he obtained a legal appointment in the state service at Berlin, but in 1806 resigned this office in order to devote himself exclusively to letters. In 1810 he was appointedprofessor extraordinariusof German literature in the university of Berlin; in the following year he was transferred in a similar capacity to Breslau, and in 1821 returned to Berlin asprofessor ordinarius. He died at Berlin on the 11th of June 1856. Although von der Hagen’s critical work is now entirely out of date, the chief merit of awakening an interest in old German poetry belongs to him.
His principal publications are theNibelungenlied, of which he issued four editions, the first in 1810 and the last in 1842; theMinnesinger(Leipzig, 1838-1856, 4 vols, in 5 parts);Lieder der ältern Edda(Berlin, 1812);Gottfried von Strassburg(Berlin, 1823); a collection of Old German tales under the titleGesamtabenteuer(Stuttgart, 1850, 3 vols.) andDas Heldenbuch(Leipzig, 1855). He also publishedÜber die ältesten Darstellungen der Faustsage(Berlin, 1844); and from 1835 he editedDas neue Jahrbuch der Berlinischen Gesellschaft für deutsche Sprache und Altertumskunde. His correspondence with C. G. Heyne and G. F. Benecke was published by K. Dziatzko (Leipzig, 1893).
His principal publications are theNibelungenlied, of which he issued four editions, the first in 1810 and the last in 1842; theMinnesinger(Leipzig, 1838-1856, 4 vols, in 5 parts);Lieder der ältern Edda(Berlin, 1812);Gottfried von Strassburg(Berlin, 1823); a collection of Old German tales under the titleGesamtabenteuer(Stuttgart, 1850, 3 vols.) andDas Heldenbuch(Leipzig, 1855). He also publishedÜber die ältesten Darstellungen der Faustsage(Berlin, 1844); and from 1835 he editedDas neue Jahrbuch der Berlinischen Gesellschaft für deutsche Sprache und Altertumskunde. His correspondence with C. G. Heyne and G. F. Benecke was published by K. Dziatzko (Leipzig, 1893).
HAGEN,a town of Germany, In the Prussian province of Westphalia. Pop. (1905), 77,498. It lies amid well-wooded hills at the confluence of the Ennepe with the Volme, 15 m. N.E. of Elberfeld, on the main line to Brunswick and Berlin, and at the junction of important lines of railway, connecting it with the principal towns of the Westphalian iron district. It has five Evangelical churches, a Roman Catholic church, an Old Catholic church, a synagogue, a gymnasium, realgyrnnasium, and a technical school with special classes for machine-building. There are also a museum, a theatre, and a prettily arranged municipal park. Hagen is one of the most flourishing commercial towns in Westphalia, and possesses extensive iron and steel works, large cotton print works, woollen and cotton factories, manufactures of leather, paper, tobacco, and iron and steel wares, breweries and distilleries. There are large limestone quarries in the vicinity and also an alabaster quarry.
HAGENAU,a town of Germany, in the imperial province of Alsace-Lorraine, situated in the middle of the Hagenau Forest, on the Moder, and on the railway from Strassburg to Weissenburg, 10 m. N.N.E. of the former city. Pop. (1905), 18,500. It has two Evangelical and two ancient Catholic churches (one dating from the 12th, the other from the 13th century), a gymnasium, a public library, a hospital, and a theatre. The principal industries are wool and cotton spinning, and the manufacture of porcelain, earthenware, boots, soap, oil, sparkling wines and beer. There is also considerable trade in hops and vegetables. Hagenau is an important military centre and has a large garrison, including three artillery battalions.
Hagenau dates from the beginning of the 12th century, and owes its origin to the erection of a hunting lodge by the dukes of Swabia. The emperor Frederick I. surrounded it with walls and gave it town rights in 1154. On the site of the hunting lodge he founded an imperial palace, in which were preserved the jewelled imperial crown, sceptre, imperial globe, and sword of Charlemagne. Subsequently it became the seat of theLandvogtof Hagenau, the imperialadvocatusin Lower Alsace. Richard of Cornwall, king of the Romans, made it an imperial city in 1257. In 1648 it came into the possession of France, and in 1673 Louis XIV. caused the fortifications to be razed. In 1675 it was captured by imperial troops, but in 1677 it was retaken by the French and nearly all destroyed by fire. In 1871 it fell, with the rest of Alsace-Lorraine, into the possession of Germany.
HAGENBACH, KARL RUDOLF(1801-1874). German church historian, was born on the 4th of March 1801 at Basel, where his father was a practising physician. His preliminary education was received at a Pestalozzian school, and afterwards at the gymnasium, whence in due course he passed to the newly reorganized local university. He early devoted himself to theological studies and the service of the church, while at the same time cherishing and developing broad “humanistic” tendencies which found expression in many ways and especially in an enthusiastic admiration for the writings of Herder. The years 1820-1823 were spent first at Bonn, where G. C. F. Lücke (1791-1855) exerted a powerful influence on his thought, and afterwards at Berlin, where Schleiermacher and Neander became his masters. Returning in 1823 to Basel, where W. M. L. de Wette had recently been appointed to a theological chair, he distinguished himself greatly by his trial-dissertation,Observationes historico-hermeneuticae circa Origenis methodum interpretendae sacrae Scripturae; in 1824 he became professor extraordinarius, and in 1829 professor ordinarius of theology. Apart from his academic labours in connexion with the history of dogma and of the church, he lived a life of great and varied usefulness as a theologian, a preacher and a citizen; and at his “jubilee” in 1873, not only the university and town of Basel but also the various churches of Switzerland united to do him honour. He died at Basel on the 7th of June 1874.
Hagenbach was a voluminous author in many departments, but he is specially distinguished as a writer on church history. Though neither so learned and condensed as the contributions of Gieseler, nor so original and profound as those of Neander, his lectures are clear, attractive and free from narrow sectarian prejudice. In dogmatics, while avowedly a champion of the “mediation theology” (Vermittelungstheologie), based upon the fundamental conceptions of Herder and Schleiermacher, he was much less revolutionary than were many others of his school. He sought to maintain the old confessional documents, and to make the objective prevail over the purely subjective manner of viewing theological questions. But he himself was aware that in the endeavour to do so he was not always successful, and that his delineations of Christian dogma often betrayed a vacillating and uncertain hand.