日本産ムカデノリ科の分類学的研究えの寄与

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(1)Title. 日本産ムカデノリ科の分類学的研究えの寄与. Author(s). 川畑, 清作. Citation. 北海道学芸大学紀要. 第二部. B, 生物学,地学,農学編, 13(1): 22-51. Issue Date. 1962-09. URL. http://s-ir.sap.hokkyodai.ac.jp/dspace/handle/123456789/5752. Rights. Hokkaido University of Education.

(2) Vol. 13 No. 1 Journal of llokkaido Gakugei UniverBity Dec., 1962. A CONTEIBUTION TO THE SYSTEMATIC. STUDY OF GEATELOUPIACEAE. FEOM JAPAN (1) Seisaku KAWABATA. Biolngical Laboratory, Iwamizawa Branch, Hokkaido Gakugei University, Hokkaido Japan. Contents. I. .. Introduction. .............................................................................................. l. ][. Historical survey .................................................•...............•••...••••.•••.••••••• 2. Ill. Description of the species wth special reference to the anatomy and tlie developmental morphology ..................................................................... 3. IV. Explanation of plates ................................................................................. 13. I. Introduction A number of systematic studies of Grateloupiaceae have hitherto been made by various. investigators since Pr. Schmitz (1889) established the present family. In those studies, however, the characteristic of each genus belonging to this family is chiefly based on the external and the internal structures of the thalli and on the distribution of the tetrasporangia. Therefore there arise many points of questions and indistinctness in characters among the genera as among the species. As regards the Grateloupiaceae from Japan, Okamura (1936) gave an account of 39 species and 9 genera in his Nippon Kaiso-shi, some of them are illustrated by him in Icones of Japanese Algae and Nippon Kaiso-shi. Among these speciese, such species as Grateliipia twuturu Yamada, Pach.ymeniopsis lcmceolata Yamada (Syn. Aeodes lanceolata Okamura), Pachymeniopsis yendoi Yamada (Syn. Pcichymenia canwsa Yendo, (non J. Ag.), Pachymeniopsis eilipfica Yamada (Syn. Grateloupia el/.ipti.ca Holmes), Pachymeniopsis pseudoeiliptica sp. nov. (in manuscript), Phyllymenia spcn'su (Okam.) Kylin, and some undetermined species have a considerable resemblance both in the external appearances and in the internal structures. The external and the internal structures of the thalli of these species were described by Holmes, Okamura, Yendo and Yamada in their works. On the other hand, the development of reproductive organs of these algae have been only partially known from the brief remarks made on the cystocarps given by above mantioned authors. Since 1953, the writer has been conducting the studies on the structure of the thalli - 22 -.

(3) Seisaku Kawabata. and the reproductive organs of the above mentioned species to find out a distinct differences among them, so as to contribute to the systematic study of Grateloupiaceae under the guidance of Prof. Yamada. In the present study a most careful observation was made upon the following items : 1) Whether the distinction between the outer and the inner cortex is clear or not. 2) Whether the cells of inner cortex are stellate or not. 3 ) The thickness of cortex. 4) Whether the disposition of cell rows in cortex is anticlinal or not. 5) Whether the transverse connections among the cortical cells are present or not. 6 ) Whether the cystocarps have a tendency of scattering or that of collecting. 7) Whether the cystocarps are enveloped with special enveloping filaments or not. 8) Whether the enveloping filaments of cystocarps are reticulated or not. 9) Whether the tetraspores are scattered or forming nemathecia. 10) The developmental stages of cystocarp. The materials used for the present study were collected in the following localities. Muroran, Hokkaido, Gruteloupia turutwu Shirikishinai, Hokkaido, Grateloupia ttirutu.ru. Koshigoe, Prov. Sagami, Gratelonpia turuturu Oma, Shimoldta peninsula, Prov. Mutsu, Pachymaniopsis yencloi Onahama, Prov. Iwaki, Phyllymenia sparsci Oarai, Prov. Hitachi, Pliyllymenia sparsa Kominato and Chikura, Prov. Awa, Pachymemopsis ellipticn p.. IcmceoUita. p.. elliptica. p.. lanceolata. p.. pseudoelliptica. Koshigoe and Enoshima, Prov. Sagami,. Yabitsu and Koza, Prov. Kii, Saga, Prov. Tosa,. Kominato, Prov. Awa, P. pseiuloelHptica The materials were preserved in formalin. Dry matierials were also used as supplement to formalinpreserved materials. Here the writer wishes to express his heartiest thanks to Prof. Dr. Y. Yamada under whose guidance this work was carried out. II. Historical survey. The family Grateloupiaceae was established by Fr. Schmitz in 1889 as comprising 11 genera. Afterward, Schmitz and Hauptfleisch (1897) revised the present family, and added to it 2 genera. De-Toni (1905) gave an account of this family enumerating 15 genera.. Recently 19 genera are enumerated in this family by Kylin (1954), The external ancl 23-.

(4) A CONTRIBUTION TO THE SYSTEMATIC STUDY OF GRATELOUPIACEAE FROM JAPAN internal structures of vegetative thallus of the family were described by various authors, while the reproductive organs of the family were quite imperfectly known before Daines. (1913). Berthold (1884) was the first to describe the general structure of the present family giving a brief account of the developmental morphology of genus Grateloupia. Schmitz. and Hauptfleisch (1897) gave a brief account of Grafelou.pia filicina (Wulf.) Ag. The first detailed description of the developmental morphology of the genus Prionitis was made by. Daines (1913). Sjostedt (1926) made a detailed investigation on the Cryptonemia borecilis Kylin and the Prionitis lcmceolata Harvey, and he found that Dairies' observations were. incorrect in some points. Kylin (1930) made an intensive study on Grateloupia filicina (Wulf.) Ag. and, afterwards described in detail the developmental morphology of Grateloupiaceae in 1937.. Balakrishnan (1949) made an investigation on the developmental morphology and cytology of Grateloupia Hthophiia Boergs.. In America, Smith (1944) and Taylor (1928, 1960) made investigations on the Grateloupiaceae and some species belonging to it. On the species from Japan here examined by the writher, the investigations have hitherto been made by the following authors. E. M. Holmes, 1895, described a red alga from Enorhima Prov. Sagami as a new species under the name of Grateloiipia elliptica Holmes.. K. Yendo, 1911, gave a detailed account of Grateloupia elliptica Holmes ; 1914, identified a red alga from Mutsu Bay with Pachymenia canwsa J. Ag. and gave a detailed account of it. K. Okamura, 1916, 1936, gave an account of G. elliptica Holmes ; 1934, described a red alga from middle Honshu, Japan as a new species under the name of Aeocles lcmseolata Ok., and a red alga from Onahama Prov. Iwaki undr the name of Cyrtymenia sparsa Ok., ; 1936, gave a detailed account of Pachymenia carnosa Yendo (non J. Ag.). K. Inagaki, 1933, gave an account of Pachymenia carnosa Yendo (non J. Ag.). S. Kawabata, 1954-1958 gave a detailed accounts of the structure of fronds and the reprodutive organs of Grateloupia turuturu Yamada, Pachymeniopsis lcmceolcifa Yamada, Pachymeniopsis yendoi Yamada, Pachymemopsis elliptica Yamada, Cyrtymenia sparsa Okamura and a red alga belonging to the Grateloupiaceae. Y. Yamada, 1941, described a red alga from Muroran, Hokkaido, as a new species under the name of Grateloupia tu.ritturu Yamada. In 1952, gave a lecture at the 17 th Annual Meeting of the Bot. Soc. of Japan on the Grateloupia elliptica and a group of red algae resembling it. In this lecture he proposed to establish a new genus Pcichymeniopsis. The marked characters of the genus Pachymeniopsis according to Prof. Yamada's manuscript, may be tabulated as follows. 1. The cortex is constructed of small cells, which are disposed in anticlinal rows, dichotomously branched, progressively smaller toward the thallus surface,. -24-.

(5) Seisaku Kawabata. 2. The transverse connections among the cortical cells are frequently present, though in some species they are present only in the lower part of cortex, while in other species they are found in upper part. 3. The comparatively large cells in the lower part of the cell rows in cortex are becoming larger downward, and the contents ere connected with adjacent cells, showing stellate forms. 4. The medullary layer is composed of filamentous cells which are more or less densely interlaced. 5. The carpogonial branch arises from the middle part between the outer and the middle cortex. 6. The cystocarp contains several gonimoblast, and a net-work of filaments enveloping the cystocarp as seen in Aeodes mtidissima J. Ag. is not developed. 7. The mature cystocarps are immersed near the surface of the thallus in some spacies, and more or less deeply immersed in other species, but not so deeply. 8. The cystocarp is scattered on both surfaces with a carpostoma, and in some species has a tendency ef gathering irregularly as in Pcichymeniopsis lanceolata Yarn.. 9. The tetrasporangia are evenly scattered on both surfaces, arise from the outer cortex as the branches of the cortical filaments, and are divided into cruciate. Prof. Yamada offered an opinion that those species, Grateloupia elli.ptical Holmes, Pachymenia carnosa Yendo and Aeodes lanceolata Okam. should be removed to the new genus Pachymeniopsis. A limited observation on the development of the spermatangia of the Gratelopiaceae, such as Grateloupia ftn'uturn Yarn. G. el.iiotica Holmes, Aeodes lcmceohita Okam. and Pachymenia carnosa Yendo was made by N. Tazawa in 1960.. The reports regarding the habitats of the present algae have been published by the following authors. Hasegawa, Y., 1949 ; Hh-ohashi, T., 1937 ; Hirose. H., 1958 ; Kawabata, S., 1939, 1960 ; Kawashima, S., 1955 ; Okamura, K., 1927, 1930 : Saito, Y., 1956 ; Segawa, S., 1935 ; Takamatsu, T., 1938, 1939 ; Toldda, J., 1954, 1959. III. Description of the species with special reference to the anatomy and the developmental morphology 1. Crateloupia tururu Yamada Pl. 1-2 Yamada, Notes on Some Jap. Alg. IX, 1941, p. 205, Pl XLVI ; Syn. Halymenia twutw'u Okamura in Herb. ; Kawabata, Sorui (Bull. Jap. Soc. Phycol.), 1954, Vol. 2, No. 2, pp. 29-32 ; Tazawa, Sorui Vol. 8, No. 3, p. 112. Tokida, The Mar. A\g. of South. Saghalien 1954., p. 163. Japanese name : Tiiruturu Material The material was collected by the writer from Muroran, Hokkaido, the type locality, in.

(6) A CONTRIBUTION TO THE SYSTEMATIC STUDY OF GRATELOUPIACEAE FROM JAPAN Oct. 1953. It grows on rocks in lower littoral zone. Structure of the Thallus This alga has a linear-lanceolate thallus arising from a small disc-shaped holdfast with a short stipe. The blade is suddenly spreading from the stipe, sometimes provided with porliferations, almost always undulate (Pl. 2, c-d). The thallus is about 400-500 /-< thick, usually reaches a length of about 30 cm or more, has a width of about 5-10 cm. The cortex is commonly 5-6 layers of cells, showing individual variations in thickness. In the basal portion of tha thallus, the cortex is composed of about 20 layers of cells. The outermost cortical cells are oblong, while those of the second and the third layer are almost spherical. The cells of inner cortical layer are irregularly shaped, rarely taking stellate forms. Among the inner cortical cells, transverse connections are frequently found. The medullary layer is composed of lax filamentous cells with a length several times the breadth. The medullary filaments are 5-6 /.< in diameter, often dichotomously branched, loosly interwoven, some of them tending to be disposed perpendicularly to the surface of the cortex. The distinction between the outer cortex, the middle one and the inner one is not evident. The cells of the tetrasporic frond are somewhat larger than those of cystocarpic frond, the former being more laxly disposed than the latter (Pl. 1, a. Pl. 2, a, b).. Tetrasporang-ia. The tetrasporangia arise as secondarily developed simple-celled branches from the cells of the inner cortical filaments. The tetrasporangia are usually homologous with the secondary cells from the apex of cortical filaments, divided cruciately, and scattered equally on both surfaces of the thallus (Pl. j, k).. Development of the Cystocarp The carpogone and the auxiliary cell are formed in separate bushy filaments known as ampullae, ampullary cell clusters or filamentous baskets. The carpogonial branch is two-celled, the upper cell of which is the carpogone, and the lower one is the hypogynous cell. The hypogynous cell is supported by the supporting cell in the bottom of smpulla. The ampullar filaments surrounding the carpogonial branch start as an accessory formation from a cell in the inner cortical layer. The carpogone produces a long trichogyne which is conspicuously swollen near the carpogone. The swollen part of the trichogyne has an appearance of being a cell, but practically there is only spirally coiled protopoasmic substance present in that part. The trichogyne projects above the surface of the thallus when fully developed (Pl. 1, b, c). After fertilization the carpogone fuses with the hypogynous cell (Pl. 1, d-f), but the writer was not able to ascertain vyhether other neighbouring cells fuse with them or not. From the fused cell, several connecting filaments are sent out, each of which may grow. -26-.

(7) Seisaku Kawabata. to an auxiliary cell. After fusion with a connective filament, the auxiliary cell cuts off a primary gonimoblast, from which the secondary gonimoblast is produced. This secondary gonimoblast gives off another cell which grows successively and splits off a number of gonimoblast cells of higher order (Pl. 1, g-i). It seems that, in mature cystocarp, the auxiliary cell is fused with neighbouring cells and forms a large fusion cell. The nucleus is invested with enveloping filaments which are originated from exhausted ampullar filaments. Besides this enveloping filaments there are another ones which originated from medullay tissue. The mature cystocarps are scattered all over the frond except the basal portion, showing a tendency of gathering in small groups (Pl. 1,1). Spermatang-ia Researches on the spermatangia have been conducted by N. Tazawa (1960). The spermatangial mother cells are formed in the surface layer of male frond. They are born from the apical cells of each cortical filaments, homologize with the surface cells of the tetrasporic and cystocarpic frauds. The mature spermatangium is spherical or ellipsoidal, 4-5 1.1. in diameter. The sperm escapes outside through the breaking at the top of the spermatangium.. To the Garteloupin fiiicina (Wulfen) J. Ag. the type species of genus Grateloupia (Agardh. 1822) (Pl. 25, a), the present species has a considerable resembrance in inner structure of the thallus as well as in feature of cystocarp. There is no question raised in placing the present species in the Grateloitpia. 2. Pachymeniopsis lanceolata Yamada Pl. 3-6 Yamada, Manuscript of the lecture at the 17 th Annual Meeting of the Bot. Soc. of Jap. 1952 ; Syn. Aeocles lanceolata Okam., Icon. Jap. Alg. Vol. 7, 1934, p. 42, Pl. 322 ; Aeodes J. Ag., Epicr. 1876, p. 678 ; Kawabata, Sorui, 1954, Vol. 2, No. 3, pp. 67-71 ; Tazawa, Sorui, 1960, Vol. 8, p. 115. Japanese name : Fudaraku. Material The material was collected by Prof. Yamada from Shichirigahama, Sagami-bay on April 6, 1954., and by the writer from Koshigoe, Sagami-bay on April 15, 1956. The male frond was collected by N. Tazawa from Mera, Prov. Awa, in April 1960. Structure of the Thallus The thallus is membranaceous, dull rose-reddish in colour, with one or more blades, growing from a dis-shaped holdfast, usually with a short stipe, often sessile. The blades are rapidly expanding from the basal portions, upper half of which are at times longitudianlly incised into several segments. In the examined specmens the mature female thalli. are forming an irregular patches, while the tetraspric thalli are rugose as in the thalli of -27 -.

(8) A CONTRIBUTION TO THE SYSTEMATIC STUDY OF GRATELOUPIACEAE FROM JAPAN Phyllymenia sparsa (Okam.) Kylin (Pl. 5, a-c). The thallus is about 1000 /-< thick. The cortical layer is commonly about 8 layers of cells, often 10 or more layers of cells. The outer cortical layers are composed of compacted, dichotomously branched filaments standing perpendicular to the surface, with no transverse connection. The outermost cortical cells in the yunger part of the thallus bear partly a more or less dense growth of elongated hairs (Pl. 6, f). The middle cortical layers are composed of about 3 layers of large elliptical cells, frequently with transverse connections. The inner cortical layers consist of large cells, which are up to 40 /.( in diameter, irregularly shaped, commonly spherical or stellate, with transverse connections, often contain granular substance, and the innermost cells are connected with medullary filaments. The medullary filaments are compactly interlaced, tending to lie parallel to the surface of the thallus. The filamentous cells have a length several times the breadth, frequently stellate, often contain granular substance (Pl. 6, c, e). Tetrasporangia The tetrasporangia arise from the cells of middle cortical layer, scattered on both surfaces of the thallus, oblong in side view, cruciately divided. The surface of mature tetrasporophyte are rugose, the rugosities are running longitudinally (Pl. 4, e). Spennatangia The spermatangia are formed all over the surface of the male thallus except the basal portion. The cortical filaments producing the spermatangia are dichotomously branched and consist of small spherical cells, being easily distinguished from steril cortical filaments which are composed of large, long cells (Pl. 6, d). In the male frond there is noticed neither the characteristic patches as of the carposporophyte nor the rugosity as of the tetrasporophyte. Development of the Cystocarp The carpogonial branch and the auxiliary cell are formed in separate ampullae. The carpogonial branch consists of a carpogone and a hypogynous cell is arising from the bottom of carpogonial ampulla. The supporting cell corresponds to the 3 rd-7 th cell from the stellate cell in middle cortical layer. The trichogyne is conspicuourly swollen near the carpogone and projects appreciably above the surface of the thallus (Pl. 3, b). The auxiliary cells also are formed in the bottom of ampulla, very large in shape and proto-. plasmic (Pl. 4, b). The ampullar branch which produces the auxiliary cell is divided from the innr cortical stellate cell. The cells of carpogonial ampulla are somewhat smaller than those of auxiliary ampulla, some of the cells of the former being usually cup-shaped,. while those of the latter are usually spherical or elliptical (Pl. 3, b). After fertilization the carpogone fuses with the hypogynous cell, and then several connective projects arise from the fusion cell towards adjacent inner ceotical cells and the medullary cells. The fusion cell increases its contents and becomes larger, presumably having fused with more neighbouring cells. From this large irregulary shaped fusion 28 -.

(9) Seisaku Kawabata. cell is given off non septum connecting filaments (Pl. 4, a). The connecting filaments seek out the auxiliary cells and fuse with them (Pl. 3, e, g). After fusion with the connecting filament, the auxiliary cell cuts off primary gonimoblast, and successively secondary gonimoblast. From the secondary goniblast, higher order gonimoblasts are cut off (Pl. 3, e-g). During the formation of nucleus, the ampullar filaments divide a number of branchlet from them. The terminal of these branchlet are connected with neighbouring medullar filaments and inner cortical cells (Pl. 3, f). The ampullar filaments are evidently nutritve in function and are used up in the course of development of the gonimoblast and the carpospores. The mature cystocarps are invested with a few enveloping filaments, but the filaments form no net-work. The mature cystocarp with an ostiole is half immersed into the medullary layer, and never protects out of the surface of the thallus (Pl. 4, c, d) In maturity the cystocarps have a tendency to be collected in an irregular patches. In some old thalli the borders between the patches disappear and the cystocarps are evenly scattered. The present species was first described by Dr. Okamura in 1934 under the name of Aeodes lanceolafa Okamura in his "Icones of Japanese Algae Vol. 7, p. 42.", but he noticed in his description that "Plant seems to de closely resembling Aeodes nitidissima J. Ag. of which, I have no reliable specimen to compare with". Prof Yamada examined the present species, and judging from the structure of the thallus and the characters of cystocarp, he proposed in 195.2 to remove this species from Aeodes to Pachymeniopsis. Originallry, the genus Aeodes is confined to Southern Hemisphere, and the type species is Aeodes nitidissima J. Ag.. The writer was able to make observation on the structure of the thallus and the female reproductive organ of Aeodes nitidissima J. Ag. using the material from Auckland which was offered him by Prof. Yamada. The microscopic photograph is shown in Pl. 23, 24. In this observation, the cystocarp of the present species is found to be quite different from Aeodes nitidissima J. Ag. in respect of the enveloping filaments of cystocarp. Therefore, the present alga is to de removed from Aeodes to Pacliymeiv'opsts.. 3. Pachmeniopsis yendoi Yamada Pl. 7-10 Yamada, 1952, 1. c. ; Syn. Pachymenia carnosa Yendo (non J. Ag.), Notes on Alg., New to Jap., II, 1914, p. 279 ; Pcichymenia J. Ag., Epicri. 1876, p. 143 ; Inagaki, Mar. red Alg. of Oshoro Bay, 1933, p. 26 ; Kawabata, Sorui, 1957, Vol. 5, No. 1, pp. 8-13 ; Okamura, Nippon Kaiso-shi, p. 545 ; Tazawa, 1960, 1. c. Japanese : Akahacia Material The matreial was collected by the writer from Oma, Shimokita Peninsula Prov. Mutsu in April 1954. The male frond was collected by S. Kawashima from the same locality. on Aug. 5th 1956.. -29-.

(10) A CONTRIBUTION TO THE SYSTEMATIC STUDY OF GRATELOUPIACEAE FROM JAPAN Structure of the Thallus The thallus is attaining about 30 cm in height, 2-5 cm in width, tapering toward the apex, about 1000 p. in thickness. At first sight the thallus has some resemblance in extermal appearance to Pachymeniopsi.s eliiptica Yamada, especially so in the shape of the. holdfast which is produced from undersurface of the thallus. The thallus is split into several blades. In some thalli the basal portions are becoming very narrow, and consequently seem to have a stipe. It is a remarkable character of the present species that,. the thllus has angulated edges as if cut with a knife at right angle to the surface of the thallus (Pl. 8, c, d). The outer cortex is about 7 layers of cells. The cortical filaments are divided into dichotomously, which are composed of obovate or oblong cells, with some transverse connections. The inner cortical layers are constructed of large stellate cells, which are vertically and horizontally connected with each other (Pl. 10, c). The form of cells in. cortical layer much differs with the individuals and with their stages of maturity. The medullary layer occupies about two fifths in thickness of the thallus in the view of cross-section, and is composed of compactly interlaced filaments and stellate cells with long processes (Pl. 22, d). Most of the medullary filaments lie parallel to the long axis of the thallus, containing rich granular substance (Pl. 9, a). It may be mentioned. incidentally that the writer has found during the examination of the thallus, a quantity of parasitic algae resemblng Chaetosiphon sp. in the medullary as well as in the cortical layer.. Tetra sp oran g'ia. The tetrasporangia are formed in the middle cortical layer as the branches of the cortical filaments, cruciately divided, scattered on both surfaces (Pl. 8, b ; Pl. 10, a). They are scarcely discernible with naked eye neither in fresh specimens nor in dried ones.. Development of fhe Cystocarp The carpogone and the auxiliary cell are formed in separate ampullae. The ampullar. filament is originated from the longitudinal division of the cell in middle cortical layer. The cell divided from cortical cell cuts off in succession one or two cells, remaing in the common gelationus membrane with the initial cells then the new filament breaks the common gerationus membrane out, continues to grow and develops to ampullar filaments (Pl. 7, a-c). Though the well grown ampulla seems as if it was divided from the terminal cell of medullary filament, it is only secondarily connected with the terminal cell of medullary filament. The carpogonial branch is two-celled, develops from the supporting cell in the bottom of ampulla. The upper one of the two cells is carpogone, and the lower one is hypogynous cell. The trichogyne is usually very swollen up close to the carpogone, in whose swollen part the protoplasma is spirally coiled. It seems that,. the large nutritive stellate cell beneath the ampulla supplies nutrious substance to the -30-.

(11) Seisaku Kawabata. ampulla (Pl. 7, f). Compared with the carpogonial one, the auxiliary ampulla is more ramified, and the shape of the cell is somewhat larger than the latter (Pl. 7, d, f). At the upper portion of the auxiliary ampulla there is already occurred a depression before the auxiliary cell fuses with connecting filament. The auxiliary cell is protopoasmic, being much larger than the other cells of ampullar filaments, produced intercalary in the bottom of the ampulla (Pl. 7, d, e). In the present study, the writer was not able to observe the fertilized carpogone, but judging from the fact that some of the auxiliary cells are furnished with connecting filament, it is presumed that the fertilized carpogone might have sent out connecting filaments from it. The ramification of ampullar filament in the present species is not so remarkable as in those of Grateloupia turuturu Yarn., Pacliymeniopsis lanceolata Yarn. and Phyllymenia sparsa (Okam.) Kylin. The ampullar. filaments joint with the cells of middle or inner cortical layer by plasmic threads (Pl. 7, e). That is the reason, perhaps, why the enveloping filaments of cystocarps of the present species are poor. At the basal portion of the cystocarp, ampullar filaments remain as plasmic filaments. The auxiliary cell, after fusion with connecting filament, cuts off transversely primary gonimoblast, and the second division subsequently takes place in primary gonimoblast, then further successive transverse or diagonal division takes place (Pl. 7, h, i ; Pl. 8, a'). The mature cystocarp is invested with eveloping filaments only in the basal portion, scattered on both surfaces of the thallus, immersed to the cortical layer, not to the medullary layer so deeply. The rudimentary nucleus is not so evident as in other species of Pachymeniopsis. Spermatangia N. Tazawa (1960) has made an investigation on the spermatangia of the present species using the material from the same locality as the writer's, the description and illustration of which is given in Sorui Vol. 8, No. 3, p. 115. The present species was identified by Dr. Yendo (1914) with Pachymenia carnosa J. Ag.. The genus Pachymenia is originally confined to the Southern Hemisphere, and the type locality is Cape of Good Hope. The Pachymeniu carnosa J. Ag., according to the original description, has a cystocarp with network of filaments, while the cystocarp of the present species lacks it. Accordingly, as Prof. Yamada (1952) has proposed, the present species is to be removed from Pachymeni.a to Pachymeniopsis. 4. Pachymemopsis elliptica yamada Pl. 11-14 Yamada, 1952, 1. c. ; Syn. Gratelou.pia elli.pticf/ Holmes, New Mar. Alg. from Jap., J. of Un. Soc. Bot. Vol. 3, 1895, p. 253 ; Yendo, Kaisan Shokubutsugaku, 1911. p. 736 ; Kawabata, Sorui, 1958, Vol. 6, No. 1, pp. 16-23 ; Okamura, Nippon Sorui Mei-i, 1916, p. 109 ; Id., Nippon Kaiso-shi, 1936, p. 543 ; Tazawa, 1. c., pp. 113-115.. -31.

(12) A CONTRIBUTION TO THE SYSTEMATIC STUDY OF GRATELOUPIACEAE FROM JAPAN Japanese name : Tanbanori. The present species was first described as a new species by Holmes (1895) basing on the material from Enoshima Prov. Sagami under the name of Grateloupia elliptica Holmes. in his "New Marine Algae from Japan". Yendo (1911) and Okamura (1916, 1936) have each given a detailed account of this alga, but these descriptions are chiefly made in view of outer characters of the thallus. Prof. Yamada gave a lecture on this species (1952, 1. c.) and proposed to transfer this species to Pachymeniopsis. Material The present study is based on the materials collected from Koshigoe in March 1954,. April 1955, and from Chikura Prov. Awa in April 1956, 1957 by the writer. Structure of the Thallus Young thallus of the present species has no stipe, and attaches to substratum with discoidal holdfast, which is produced froiTi the central part of undersurface. As the thallus grows up, it splits into several blades. In mature thallus each blade is about 2030 cm high, 5-15 cm wide and about 1000 /-( or more thick (Pl. 12, d). The anatomical structure of the thallus shows various appearances in proportion as the thallus grows older, and in different parts of the same thallus. In mature thallus, the cortex is about 20 layers thick ; the outer cortex is about 10 layers thick, and the middle and inner ones are about 4 layers thick respectively. The outer cortex is composed of oblong assimilating cells disposed in dense, anticlinal rows. No transverse connections are found among the cells of outer cortex. The middle cortex consists of about 4 layers of cells, which are composed of large, oblong cells with transverse connections. The inner cortex consists of about 4 layers of cells, constructed of large, 3-5 armed, stellate cells (Pl. 11, e, f). The medullary layer, in mature thallus, occupying about one third of the thickness of the thallus (Pl. 22, e ; Pl. 13, c) is densely composed of interlaced, multi-ramous filaments. The cells of medullary filaments are usually about 10 /-< in diameter, with a length several times the breadth, frequently stellate or spherical. In young thallus the medulla occupies about two-thirds or more in thickness, and its filaments are loosely interwoven (Pl. 11, f ; Pl. 13, a, b). Tetrasporangia The tetrasporangia are formed in the middle cortical layer as the unicellular branches of the cortical filaments, scattered on both surfaces of the thallus, which are usually divided into cruciate, while in some thallus divided intp zonnate (Pl. 12, b, c). In some specimens tetrasporamgia and cystocarps occured mixed together. Development of the Cystocarp Both the carpogonial and the auxiliary ampullae develop in the middle cortex. The developmental stages of the ampullae are in the same way as in those of Paclt.ymeniopsis. -32-.

(13) Seisaku Kawabata. yendoi Yamada (Pl. 11, a ; Pl. 13, d). The well grown ampulla is connected with middle and inner cortical cells in places, and it seems that the nutrition is supplied from the surrounding cells to the ampulla (Pl. 11, b). The carpogonial branch is composed of carpogone and a hypogynous cell.. The supporting cell is bearing the hypogynous cell at the bottom of the ampulla. The trichogyno is somewhat swollen up close to the carpogone (Pl. 11, b ; Pl. 14, b). The auxiliary ampullar cells are larger than those of carpogonial ampulla, and the cells are. mostly spherical (Pl. 11 g ; Pl. 14, a). Though in the present study the writer was not able to observe the connecting filaments issuing from the fertilized carpogone. judging from the fact that the auxiliary cells are fused with conecting filaments, it may be presumed that the fertilized carpogone issues out some connecting filaments from it. After fusion with connecting filaments, the auxiliary cell cuts off a primary gonimoblast, which grows successively and splits off a number of gonimoblast cells (Pl. 11, c, d). The mature cystocarp has no specific filamentous cluster, except in the basal portion, where it is surrounded by the exhausted ampullar filaments. The sizes of carpospores are various in different cases, the largest ones are 30 /-< in diameter, while the rudimental ones are less than 10 /-( in diameter. The ripe carpospore is covered with gelationus wall. In the examined materials, the germinations of carpospores frequently take place in the cystocarps, and branched threads are growing up from original carpospores. The carpostoma is furnished with incurved assimilating filaments, showing specific appearance, peariform in side view. The cystocarp is immersed in the cortical layer, not deeply into the medullary layer (Pl. 12, a, a' ; Pl. 14, d). In the examined material from Enoshima, the cystocarps were found only in the part of the first year, not in the part of the second year, while in the young materials collected from Chikura in April of the same year, young carpogonial and auxiliary ampullae were observed. In view of this fact, the good materials suitable for observing the developments of reproductive organs should be collected during summer and autumn in Kanto-district. Spermatangia The spormatangia was not examined by the writer. Mr. Tazawa has made an investigation of the male frond of this species and has given an account of it in Sorui Vol. 8, No. 3, pp. 112-115, in 1960. The present species also, as Prof. Yamada proposed, can not be placed in the genus Grateloupia in consideration of its thick cortex and the characteristic feature of cystocap.. This alga is the most typical one of the genus Pachymeriiopsis. (To be continued). - 33 -.

(14) A CONTRIBUTION TO THE SYSTEMATIC STUDY OP GkATELOUPIACEAL: FROM JAPAN. Explanation of Plates Abbreviation. A: AF : c: CP: PC: c:. H. auxiliary cell. HA MF. ampullar filament carpogone. NC. connecting fillament. s. fusion cell. SP. gonimoblast. hypogynous cell hair medullary filament nutritive cell supporting cell spermatium. IG:. primary gonimoblast (gonimoblast SM. spermatangial mother cell. mother cell). trichogyne. IG :. secondary gonimoblast. T. Plate 1 Gmteloupia turu.tu.ru Yamada a. Part of a sterile frond in trasverse view. X 400. b. Carpogonial ampulla with a carpogonial branch. X 400. c. Carpogonial branch with a sterile lateral branchlet, showing the spiral s tructure of the trichogyne. X 400. d-f. Carpogonial ampullae showing the fusions of the carpogones with the hypogynous cells, and the connecting filaments growing out from the fusion cells (PC). X 400. g. Young ampulla with a young auxiliary cell (A). X 400. h. Auxiliary ampulla with a auxiliary cell (A). X 400. i. Young cystocarp showing well developed gonimoblast. X 400. j, k. Parts of a tetrasporic frnds in transverse view. X 400. 1. Ripe cystocarp showing the fully developed carpospores and the rudimental ones.. Note the enveloping filaments which originate from the ampullar filaments (AF) and the medullary filaments (MF). X 200. Plate 2 Grateloiipia t.wutwu Yamada a. Transverse section through a segment of tetrasporic frond. X 200. b. Transverse section through a segment of cystocarpic frond. X 200. c. Habit of a mature tetrasporic frond. 1/3. d. Habit of a mature cystocarpic frond. 1/3.. Plate 3 Pachymeniopsis lanceolata Yamada a-g. X 370 a. Transverse section of a female frond showing the young ampullar filament.. - 34 -.

(15) Seisaku Kawabata. b. Carpogonial ampulla with a carpogonial branch. c. Young carpogonial ampulla with young carpogonial branch.. d. Carpogonial ampulla showing the fusion cell (PC) and the connection with the neighbouring cells. e-g. Development of gonimoblasts.. Plate 4 Pcichymeniopsis lanceoluta Yamada a. Fusion cell (FC) formed in a carpogonial ampulla after fercilization, and the connecting filaments (CF) growing out of ic.. X 370.. b. Auxiliary ampulla showing the auxiliary cell (A). X 370. c. Cross section of a thallus showing the young cystocarp. X 130. d. Ripe cystocarp. X130. e. Part of a tetrasporic f rod in transverse view. X 370.. Plate 5 Ptichymeniopsis lanceolatu Yamada a. Habit of a tetrasporic frond. 1/5. b. Habit of a male frond. 1/3. c. Habit of a cystocarpic frond. 1/5.. Plate 6 Pcicliymeniopsis lanceolata Yamada a. Ripe cystocarp in transverse view. X 100. b. Part of a female frond showing the young auxiliary ampulla. X 100. c. Part of a sterile frond. Note the cortical layers and the madullary layer. X 100. d. Part of a male frond in transverse view. X 800. e. Longitudinal section of a tetrasporic frond. X 100. f. Hairs growing out from the outer cortical cells (HA). X 100.. Plate 7 Pachymeni.opsis yendoi Yamada a-c. Developments of ampullar filaments. X 370.. d. Auxiliary ampulla with an auxiliary cell (A). X 370. e. Auxiliary ampulla producing out connective cell (CC) towards neighbouring cells.. X 370. f. Carpogonial ampulla with large nutritive cell (NC). X 370. g. Ripe cystocarp in transverse view. X 250. h, i. Developments of gonimoblasts. X 370.. - 35 ~.

(16) A CONTRIBUTION TO THE SYSTEMATIC STUDY OF GRATELOUPIACEAE FROM JAPAN Plate 8 Pachymeniopsis yendoi Yamada. a. Auxiliary ampulla with an auxiliary cell (A). X 370. a'. Auxiliary cell with a connecting filament (CF), cutting off a gonimoblast. X 370. b. Part of a tetrasporic frond in transverse view. X 370. c. Habit of a male frond. 1/3. d. Habit of a cystocarpic frond. 1/5.. Plate 9 Puchymeniopsis yendoi Yamada a. Part of a female frond showing the carpogonial ampulla, and the medullary cells containg granular substance. X 400. b. Part of a female frond showing the auxiliary ampulla and the auxiliary cell.. X 400. c. Transverse section of a young thallus showing the young ampulla. X 100. d. Part of a young thallus in transverse view. X 100.. Plate 10 Pcichymeniopsis yendoi Yamada a. Part of a tetrasporic frond in transverse view. X 400. b. Mature cystocarp in transverse view. X 100. c. Part of a cortical layer in transverse view, showing the transverse connectiosn among cells of the middle and inner cortical layers. X 400. d. Mature cystocarp with no rudimental carpospores. X 100.. Plate 11 Pachymeniopsis ellipti.cu Yamada a-d. X 400 e, f. X 250, g. X 400 a. Developmental stages of young amdullar filament. b. Carpogonial ampulla with a carpogonial branch. c. Auxiliary ampulla with an auxiliary cell having a connecting filament, cutting off the gonimoblasts. d. Auxiliary ampulla showing the development of gonimoblast. e. Part of a mature frond in transverse view. f. Part of a young frond in transverse view. g. Auxiliary ampulla with an auxiliary cell (A).. Plate 12 Pachymeniopsis elliptica Yamada a. Mature cystocarp in transverse view. X 250. a'. Germination of carpospores in cystocarp. X 400. - 36 -.

(17) Seisaku Kawabata. b. Part of a tetrasporic frond in side view, showing the developmental stages of tetrasporangia. X 400. c. Part of a tetrasporic frond, showing the tetrasporangia dividing zonately. X 400. d. Habit of a cystocarpic frond. 1/4.. Plate 13 Pachymeniopsis ellipica Yamada a. Part of a cortex of a young frond, showing the dichotomously branching cortical filaments and the stellate cells of the inner cortical layers. X 400. b. Transverse section through a young frond. X 100. c. Transverse section through a mature frond. X 100. d. Part of a female frond in transverse view, showing the young ampullar filament.. X 400. Plate 14 Pacliymeniopsis ellipt.icu Yamada a. Part of a female frond in transverse view, showing the auxiliary ampulla. X 400. b. Part of a female frond in transverse view, showing the carpogoninl amplla. X 400. c. Part of a female frond in transverse view, showing the thick cortex. X 100. d. Mature cystocarp in transverse view. X 100.. 37 -.

(18) A CONTRIBUTION TO THE SYSTEMATIC STUDY OF GRATELOUPIACEAE FROM JAPAN. d. E. - 38 -.

(19) Seisaku Kawabata. Plate 2. - 39 —.

(20) A CONTRIBUTION TO THE SYSTEMATIC S'l'UUY OF GRA't'ELOUPlACEAE FROM JAPAN. K. E. 03. - 40 -.

(21) Seisaku Kawabata. 1 fc. w. 41 -.

(22) A CONTRIBUTION TO THE SYSTEMATIC STUDY OF GRATELOUPIACEAE FROM JAPAN. IL iiil^ SiSSHiaS. teSSBRi '%. <t&ti::l. 01. t9. &<~. - 42 -.

(23) Seisaku Kawabata. ?SiyM M^Sffii'^ ^';'^^%ii ^. îa:^ ^SM^ y 'Aff'"s/f^-• "L y,. 'a^h ^"". îtSi. ^^? ^*r!^CS. '^WM i;^,-;--^,. - s.' ^ '>y^. •UK ^^ 'ViX. ^9.''.. -.". ^;t::. ww^..'.-. ^ '•' .•<» "^^.-^ b €••'. & A. ^.<t. <'^'' ^ ^. ^. ^! '' ^\'i. '.M/^'. :^'^^. •fy^yf^ 'î€^. ^.i_4^. •'»' :3-:^H. %<fS^,^ A ^ SVk. ''« 1" '. •,.' .•-'"< *•.. '^ (•,'-.,). 1\ ^ I.' ^ t • (I-. h ,. -43.

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(25) A CONTRIBUTION TO THE SYSTEMATIC STUDY OF GRATELOUPIACEAE FROM JAPAN Plate 7. - 45 -.

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(27) Seisaku Kawabata. Plate 10. ~; 'A1'- .V^'i «>**;,.-^r- • •-^.•-i%r „,<« ,„. • !'l!lliei^.'. -47-.

(28) A CONTRIBUTION TO THE SYSTEMATIC STUDY OF GRATELOUPIACEAE FROM JAPAN. g +^. c^j. _ 48 ~.

(29) Seisaku Kawabata. Plate 12. d. -49-.

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(32)