Page 1 MORPHOLOGY of REBOULIACEAE III. Development

MORPHOLOGY OF REBOULIACEAE III. III. Development of sex organs ， sporogonium and interrelationships interrelationships of the various genera 1

By P. KACHR00 2

P . KACHROO : デソガサゴケ科の形態乍的研究(;I( =: ).生殖掠干?の発生， l胞子体及び何々の属聞の類縁

Introduction Mehra and K achroo (195 1) in their deta i! ed account of sporeling g erminati on stlldies stlldies in the family fOllnd the spore strllct ure built on a common plan and probab ・ ly ly affording a feature of phylogenetic value. Kachroo (19 54a) discussed the distribution tion of the family in India. He further (1954b) gave a morphological account of Mannia indica ， a few species of Asterella ，and Plagiochasma aρ'pendiculatum ，and described a number of morphologically interesting forms in Asterella. Since the present Inve s tigation gives an account of d 巴velopment of sex organs ， spo rogonIum and interrelationships of the various genera it Is thought desirable to g ive a concise historical acco unt of work done so far on these.

Roboulia :- Cavers (1911 ) repo rted the presence of 2 archego n ia in the same re- ceptacular ceptacular notch and a carpocep halum structure similar to Saute パα， formation of octants octants in the embryo (c .f Cavers ， 1904) and a we l1 developed Ii d which fa l1 s off in frangments on opening of the capsule. W oo dburn (1919 ) wrote a preliminary note note on the embryo logy and a comprehens i ve accou nt of the same was Iater given by Haupt (1 921 ) wh o aIso discussed the affinit i es of Retoulia to other Marchantia - ceae ceae on the basis of development of air chambers ，development of sex organs and the the sporophyte; reporting antheridia developing intermitten tI y without interrupting the the activi ty of the apical cel l. The latter is ，however ，i nvolved in the formation of the the archegon ial recepta cI e. The reduction in the numb er of archegonia in Marchan- tiaceae tiaceae reaches highest express ion in Ree O'U li α. The embryo is of the filamentous type type and the potentially sporogenous tissue is diverted to form e laters later than in

March α冗 tt α viz. both belong to the same cell generation. In this re sp ect the spo ・ rophyte rophyte of Reb 側 lia is primitive. Mannia:-Leitgeb Mannia:-Leitgeb (1881 ) stud ied structu re of the recepta cI e and the orig in of a rchegonia. Kienitz -Ge rI off (1874 ，18 75) found regular formation of octants in the

sporogoni um . Cavers regar d ed the ge n us as closely related to Re' 肘 d 句 except tha t the the u pper portion of the ca psule wall forms a well-deve loped lid and the capsule shows the pre se nce of an annulu s. Campbe lI (191 8) stated th at the deh i scence of the the capsu le is by a lid .

1 Part 1 appeared in the Bryolo gist ，57 : 159 -166 ，1954 ; and Part II in the J ourn. Hat- tor i Bot. L ab. 12 : 34 -52 ， 1954. The major portion of this work was completed at Panjab University ，India ， as a special Research Sc ho l ar du ri ng 1948 - 50; and so me p art at Gauhati Un iv ersity ，India ，where the author served as a Le ctur er. Thanks are due to the authorit i es at at both the places for laboratory faci ¥ities. ~ Malaria Re search Laboratory ，Damodar Valley Corporn .，Burdwan ，Indi a. 2 Journ. Hattori Bot. Lab . No. 19 1 958

Astere Ua: -Le itgeb (1881) gave an account of structure and origin of the recepta c¥ es and Cavers (1911) regarded the perianth as neare st ally of that of the highest highest type of Marchantiaceae and that each involucre ， in most of the species ，

contained contained more than one archegonium and that it resembles M α88a1o 党，go αexcept for the the absence of a perianth in the latter genus. Pierce (1902) investigated the discharge of spermatozoids in A_ calif m- nw α，which is explosive as in 白悌cephalum conicum. Campbe Il (19 18 ) studied the morphology of A. cαlif m-n w α，and reported a quadrant type of embryo. Peissel (1925) described the development of sex organs and sporogonium in A_ bl 叫 meana. She found the growing point of the thallus con- sisting sisting of a transverse row of 4 cells of which the apical cell seen in a median longitudinal longitudinal section is but one member and that the carpocephalum arises as a dorsal sal outgrowth of the tha Il us as in Plagiocha8m α， that the marginal spical meristem remains at the base of the young recepta cJ e and fina Il y becomes lost in the formation tion of the stalk_ Haupt (1929) reinvestigated A. califm 叫 cαand found the development of sex organs similar to other Marchantiaceae ，absence of mucilage hairs in

the the antheridial pits (ot herwise present in Rebo 叫 li α) ，2-3 archegonia in each group and neck canal ce Il s numbering 8; after formation of archegonia there is development of new growing points beneath the posterior region of each wing to continue the the growth of thallus but these were adventitious ，having no relation to original apical apical cell; further that in A. pa1m 6' ri the embryo was filamentous . Mahabale and Bhate (1945) in A. απ gust αreported the development of sex organs similar to those of of other Marchantiaceae ，a filamentous embryo and the presence of radical thickenings ings on the walls of ce Il s on the upper part of capsule. Crgptomitrium :-Abrams (1899) and Howe (1899 ) studied the life history of C.

te 憎 rum ， the former describing the compound nature of carpocephalum and an embryo similar to that of A. Cα1ifm 明白α(Campbell ，1918 ). Cavers found th e involucre contai ning upto 5 arc hegonia with centrifugal development and carpocephalum of

the the composite type. Haupt (1942) in a reinvestigation of C. t計 wrum found the development of sex organs typical of Marchantiaceae ，a 自lamentous embryo of 4 superimposed cells and remarked that early embryogeny resemles that of Reboulia ， A8terell αand Plag 必ch α8m αfurther that 3- 4 archegonia d

and development of sex organs as usual in Marchantiaceae. Plagiochasma: -Le it geb (188 1) and Goebel (1930 ) gave a general account of the genus; genus; the former as well as Cavers (1911) ， interpreted the archegoniophore as an adventitious adventitious branch arising on the dorsal surface of the thallus and whose apex is used used in development of carpocephalum as in Athal αmia. Cavers reported a definite lid lid breaking up into fragment s on dehiscence of the capsule. Meyer (1916 ， vide Campbe Il， 1918) described a filamentous embryo in the genus. Kashyap (1914) dis - cussed cussed the nature of the horse -s hoe shaped recepta cJ e in P. α付加u αtum and P. αp- pendicu1atum which is due to forking and ( according to him ) homologous with those of of higher Marchantiales ， representing a branch system. He further established a homology between the stalk of the female recepta cJ e in this genus ， as als o in Ath α- lam 似， with a similar sta lk of higher forms of Marchantiales and regarded the genus as a last stage in a series of reduced forms from the level of M αrch α ntia through through Reboul αStar (1916) in an investigation of a Mexican species reported that that the main apical cell of the tha Il us was not involved in formation of the archegonia l recepta cJ e or the antheridial recepta c¥ e; the latter may follow the formation of of the former; that the apical cell did not function while the antheridial group was active active but later continued grow th of the thallus and that the embryo was of the 昭和 33 年服部植物研究所報告第 19 号 3

自lamentous type. Kapoor (1931) wrote a short morphological account on P. α付必u ・ 1αt 叫明 Mahabale and Deshpande (1947) in a detailed study of the same species noted noted bisexual receptacles ，gametogenesis similar to other Marchantiales and an embryo embryo of the 五lamentous type.

Materials and methods The present investigation includes the following spec ies: Mannia indica ，As- lerella lerella blumeana ， A. reticulata ， A. mussuriensis ，A. sanguinea ，A. ρathankotensis ， A. A. sps. (probably identical with A. khasian α(Gri 伍th) ， referred to as A . “Mani- pur" in the following pages) and Plagiochasma appendiculatum. The life history has been studied in detail in all species except in A. “Manipur ". Mannia indica is is figured in detail and in others only those features are figured by which they differ differ from this species. The species were collected at Pathankot ，Hoshiarpur ，Delhousie ， Simla ，Hard- war ，Dehradun ，Mussurie ， Nainital (Western Himalayas); Gauhati ，Jowai ， Shillong and Manipur (Khasi Hills ，Assam) in spring ，summer and autumn during March 1948 1948 - August 1952. All were fixed in the field and the simple suction pump (Kachroo ， 1951) was employed to sink the material in the 白xative. F.A.A. ，CRAF and Lewitsky's fixatives were used but CRAF proved best for cytological and morphological phological studies. Microtome sections were cut 8-12μthick; safranin and light green ，and crystal -vio let and erythrosin staining schedules were followed.

Nomen c1 ature of the cited species 3 Il A Il combinations proposed here as new under Asterella have actually been ad- opted opted by Kachroo (1954) and Pande et al (1954) but are there technically not valid under the current International Code as the authors unfortunately omitted the basionyms. basionyms. The genera as we l1 the species under them are arranged alphabetical- ly ly in the following list.

Asterella Asterella angusta (St.) comb. nov. ，basionym Fimbriaria angusta St. Spec. Hepa t. 1: 1: 104 ， 1899. A. blumeana ( Nees) comb. nov. ，basionym Fimbriaria blumeana Nees ， in Gottsche et et al. ， Syn. Hep. p. 564 ， 1846. A. A. bolanderi (Aus t.) Underw. ， Bot. Gaz. 20: 61 ，1895 . A. lindenberghiana (Corda) Lindberg ，Musc. Scand ，1 ， 1879. A. ludwigii (Schwaegr.) Underw. ， Bot. Gaz . 20: 61 ， 1895. A. multijiora (St. ) comb. nov. ，basionym Fimbriaria multijioaa St.， Spec. Hepat. 1: 1: 124 ，1899 . A. A. mussuriensis (Kashyap) comb. nov. ，basionym Fimbriaria mussuriensis Kashyap ， J. Bombay Nat . Hist. Soc. 24: 345 ， 1916 A. A. nepalensis (Taylor ) comb. nov. ，basionym Fi mbriaria ne ρalensis Taylor ， Trans. Linn. Linn. Soc. 17: 387 ， 1837. A. ρ almeri (AusU Underw. ， Bot. Gaz . 20: 63 ， 1895.

~ Thanks are due to M r. V. Raghavan (Gauhati Univ.) for his assistance i n collecting some some of this information. 4 Journ. Hattori Bo t. Lab. No. 19 19 5 8

A. ρaρ ulosa (St.) comb. nov. ，basionym Fimbriari αραρ ulosa St. ， Spec. Hepat. 6: 16 ， 1917. A. ρ arviPora (S t.) comb. nov. ，basionym Fimbriaria ρα rvi ρ ora St.， Spec. Hepat. 1: 116 ，1899 . A . ρathankotensis (Kashyap ) comb. nov .，basionym Fimbriaria tathankot ensis Ka- shyap ， J. B ombay Nat. His t. Soc. 24: 344 ， 1916. A. reticulata (Kashyap) comb. nov. ，basionym Fimbriaria reticulata Ka shyap ，J . Bombay Na t. Hist. So c. 25: 279 ， 1917. A. saccata (Wah l.) Evans ， Contrib. U.S. Nat. Herb. 20: 276 ， 1920. A. sanguinea ( L. et L. ) comb. nov. ，basionym Fimbriaria sanguinea Lehm. et Lin - denb. ， in Lehm. ， Pug_ 4: 5， 1832. A. tenella (L.) Pa l. de Beauv .， in Cuvier ed. ，Dic t. Sci. Nat . 3: 258 ， 1805. Cryptomitrium Cryptomitrium himalayense Kashyap ，New Phyto l. 14: 2， 1915. C. C. tenerum ( Hook.) Aust. ex Underw. ，Bul l. 111 . State Lab . Nat . Hist . 2: 36 ， 1884. Mannia californica (Gottsche ) Wheeler ，The Bryol ogist 37: 88 ， 1934. M. fragrans (Balbis) Frye & Clark ， Univ. Washington Pub l. Bio l. 6: 62 ， 1937. M. indica (St ，) comb. nov. ，basionym Grimaldia indica St .， Spec. Hept. 6: 10 ， 1917. M. ρilosa (Hornem.) Frye & Clark ，Univ . Washington Pub l. Bio l. 6: 67 ，1937 . M. ru ρestris ( Nees) Frye & Clark ，Univ . Washingt on Pub l. Bio l. 6: 64 ，19 37 . Plagiochasma Plagiochasma a ρ'pendiculatum Lehm. et Li ndenb. ， Pug. 4: 14 ，1832 . P. P. articulafum Kashyap ，New Phyto l. 13: 320 ，1914 . P. P. cuneatum Evans ，Amer. J. Bo t. 19: 627 ， 1932. P. P. intermedium Lindenb. et G .， in Gottsche et al. ，Syn . Hep. p. 513 ，1846 . P_ P_ ru ρestre (Forst.) St.， Spec. Hepa t. 1: 80 ， 1898. P. P. simlense Kashyap ，J . Bombay Na t. Hi st. Soc. 25 : 279 ， 1917. Reboulia Reboulia hemisthaeric α( L.) Raddi ， Opusc. Scient di Bolo gna 2: 357 ， 1818.

Receptacles Receptacles and the development of sex organs In In M archantia and .other genera Marchantiaceae the growth of the thallus ceases ceases with the development of archegoniophore while in Plagiochasma and Atha- lamia lamia the apex of the thallus con tinue s to grow in length after the differentiation of of the archegoniophore. All All the species under consideration here are strictly protandrous in development and by the time the female receptacle becomes prominent most of the mature antheridia have released their spermatozoids. In Asterella the antheridia are are aggregated into circular or oval cushions which project but slig htl y above the surface surface of the tha l1 us; except occasiona l1 y in A. sanguinea where they are much raised -up forming false receptacles (c .f. Kachroo ，) 1954b ， or in A. “Manipur" where they are in the form of receptacles - either more or less horse-shoe sha ped or or crescentic (Figs. 85 ，90 ). In A. blumeana such cushions are borne just behind the the stalk of the female receptacle; in A. reticulata on later al sh ∞ ts borne along the the anterior or the posterior region of the thallus; in A. pathankotensis and A. sanguinea sanguinea terminal on the main thallus and in A. mussuriensis on the lateral shoots . In M annia indica they are linear in form ，and situated either terminally or or dorsally (as occasiona l1 y in A. ρathankotensis) on the main thallus. Plagiochas- ma appendiculatum is characteristic in having sessile male recepta c1 es dorsal in 昭和 33 "1'- 服部 lîll 物 (v[Jb 河機 ~'i.~; 19 J，} d" position position and variable in fo rm; hor se.sho e shaped to ovaL The fo rmati on of an theridi a is acropetal and usually accompa nies the apical grow th of the tha l1 us or lateral shoots . But in A. californica (Haupt ，1929 ) they

1 2 3 4 5

6 7 8 9 10

14 14 15 16

Figs. Figs. 1 -16. Developmen t of anther idi um. 1-6，1 0，11 ，Mαn 叫4α4 匁dic α7-9 ， Aste1 ・ell αblu me αnα12 -15 ，A. reticul at α 16 ，Plagioch ωm αα .ppendicul αtum (m-mucilag 邑 hair ). Figs. 1-10 ，12-15 ，x5 1O; 11 ，16 ， x390. 6 ]ourn. HatlorI Ro t. I. ab. No . 19 |リ !i ~ sometimes have an intermittent development resu 1t ing in formation of sterile patches patches in between. Usua l1 y they are in 1-2 rows. Each is sepa rated from one another another by septa which persist even after the antheridia di scharge sperma tozoids. Towards the dorsal surface air chambers in many layers are quite distin ct in the space space between the passages through which the antheridial chambers open to outside. side. It is interesting that mucilage hairs only arise within the antheridial cavities in in Plagiochasma ， (as also in Reboulia and Cry ρtomitri 切 n) and not in Mannia and Asterella. Asterella. The antheridial cavities open to outside through a pore which resembles the the air pore in development except that the inner cells do not become so small and pointed. pointed. The archegonia are borne on stalked recepta c1 es which arise singly in the apical apical notch ，making their appearance as knobs. 1n the development of receptacles the the apical cell of the thallus undergoes two successive dichotomie s forming 4 cells arranged arranged in a cross-wise fasion ，which ultimately form 4 archegonia. The dome- shaped recepta c1 e is the resu 1t of the intercalary growth of the cells between these these vegetative points. 1n this tissue later air chambers appear schizogenously. A vertical section of the receptacle at a later stage shows air chambers in more than than one layer ， the dorsal ones opening to outside through compound pores. Mean- time ，a short stalk develops with an anterior rhizoidal groove. The receptacle bears bears ventrally rhizoids and scales ， the latter often showing degenerating chloro- plasts. plasts. The stalks ，when young ， is composed of cells more or less the same size and shape but later the cells towards the base become more elongated and those towards towards the apex remain sma l1 er ， densely granular and with prominent nuclei. 1n Mannia and most species of Asterella the stalk is long ， but in A. “Manipur" the stalk stalk is always sub-sessile and rarely the carpocephalum is sessile (Figs. 8&， 89) as as also in A. sanguinea (Fig. 92). The former species al so exhibit s abnormalities in in the nature of the receptacle (Figs. 86-88). 1n Plagiochasma appendiculatum the stalk stalk is often subsessile and always without rhizoidal groove. Usually Usually 4 archegonia are formed ，one in each involucre; in contrast to usual 8 in in Marchantia (Andersen ， 1929; McNaught ，1929 ). 1n these species each of the original original growing points undergoes dichotomy. 1n Plagiochasma aρρ endiculatum occasionally casionally 9 invo 昭和 33 :r 服部陥物倒究所線引第 19 号 7

ln ln A. caltfornica Haupt (1929 ) reported that the receptacle may grow out from the OOttom OOttom of the dichotomy ，an ordinary branch appearing on each side. “ This would seem to indicate dorsal origin ， the apical region of the branch continuing it growth but but undergoing dichotomy at once." The two lateral branches ，thus ，formed grow out out and the receptacle then appears to grow out from the bottom of the dicho ・ tomy . Each or both of these branches may bear receptacles. In In Mannia and Asterella it is common to observe the development of apical ， subapical subapical or lateral ventral shoots ， either arising anteriorly or on either side at the base base of the receptacle; or from the ventral surface of the thallus. They usually arise from the surface of the midrib or from the region within the notch. A detailed study study of the development of such shoots is in progress. At the moment the idea of of dorsal nature of female receptacle is improbable as the apical cell is utilized in its its formation. (l n Reboulia hemisPharica var.ρ 'angiensis Kashyap ，and often in Asterella “Manipur" and rarely in Mannia indica and Plagiochasma aρρ endicula. tum the thallus forms apical innovations ， as in Preissia ， to facilitate continued elongation elongation of thallus after formation of carpocephalum or antheridial cushion. Such ventral ventral sh ∞ ts grow out from beneath the receptacle and finally give the thallus a jointed jointed appearance). Antheridium: The antheridial initial is observed as a conspicuous cell (Fig. 12a) 12a) just behind the apex of the thallus. Later each becomes sunken in the antheridial theridial pits formed by the outgrowth of the surrounding tissue. A transverse wall wall divides it into 2 cells-the primary antheridial cell and a lower stalk cell (Fig. (Fig. 1). Transverse walls are formed in the former resulting in formation of a short short filament of 3-4 cells (Figs. 2，3 ， 12b) as in Reboulia (Haupt ，1921) ，As terella californica californica (Haupt ，1929 )，A. angusta (Mahabale and Bhate ，1945) ， Plagiochasma articulatum articulatum (Mahabale and Deshpande ，1947) ，Cry ρ tomitrium tenerum (Haupt ， 1942) and C. himalayense (Chopra and Khosla ，unpub l. ) and other Marchantiaceae (see Durand ， 1908; Andersen ，1929 ，McNaught ， 1929; Ahmad ， 1938; Mehra and Mehra ， 1939 ). In Asterella bolanderi (Campbell ， 1918) a row of 5 cells is produced before the the first vertical walls are formed (although his figure shows only 3 transverse walls walls in the superficial portion of the antheridum ) and the same is the case occasionally casionally in Reboulia (Dupler ，1 8 Journ . Hattori Bo t. Lab . No . 19 1 9 5 g

Similar Similar conditions restrict the size of the sporogonium. By this time the antheridium is sunk within the antheridial chamber which opens opens to out side by a simple pore. The antheridial wall grows by further formation tion of radial walls and is usually hyaline in adult condition (Fig. 11) . In P . aρ- ρ endiculatum the antheridium is associated with unicellular hairs which arise near its its base (Fig. 16) as in C. tenerum and Reboulia. They secrete mucilage in the antheridial antheridial chambe r. Mahabale and Bhate (1 945 ) de s cribed that in Asterella angusta exist occasional. ly ly a triangular apical cell at the apex of the antheridium adding segments on 3 sides sides (though not clear from their figure in the text ). Such a cell is not discernible in in the present species. Spermatogene sis follows the normal course described for Marchantiales Marchantiales (Champbell ， 1939; also Woodburn ， 1922). The primary spermatoge- nous nous cell is with conspicuous nucleus and nucleolus. It divides diagonally in the usual usual fashion. 9 chromosomes are present in Mannia indica ，Asterella mussuriensis ， A. A. pathankotensis and Plagiochasma ap ρendiculatum; and 18 in A. blumeana and A. A. reti culata. 4 Irregular spermatogenesis is observed in P . aρρ endiculatum and P. articulatum. articulatum. Each spermatozoid is biciliate and with a more or less twi st ed body . They were observed to be dischar ged explosively in Mannia indica and Asterella μ thankotensis as in certain other Marchantiaceae (see Peirce ， 1902; Caver s， 1903; Haupt ，1921 ). Campbell (1918 ) fo und the top of the antheridium in A. californica prolonged into into a beak or tubular neck . Such a condition is not found in any of the specie s under consideration nor could Haupt (1929) locate it in the same speci es. Archegonium: The archegoni a l initials are also superficial in origin and they arise arise in early stages of development of the carpocephalum but do n ot involve the growing points. However ， the activity of the latte r is checked by the growi ng archegonia. archegonia. The development of the archegonium is of the usual type in Mar- chantiales chantiales (Le itgeb ， 1881; Goebel ，190 5; St ar， 1916; Campbell ，1918 ; Haupt ，1921 ， 1926 ，1929 ， 1942; Mahabale and Bhate ， 1945; Mahabale and Deshpande ， 1947; Mehra and Mehr a，1939 ). The initial (Fig. 24) either divides by a transverse wall forming ing an upper archegonial initial and a lower basal cell or directly behaves as an archegonial archegonial initial (Fig. 18). It is cut off by 3 inter sect ing walls into an axial cell and 3 periph

4 A detailed account of cytology of some North W est Himalayan Marchantiales will appear appear jointly with Prof. P. N. Mehra . H11 府133 '1'- Ilji 泊¥hU 物研究所被-:'i U'; 19 号 9 tally tally or is nearly penden t. There are six vertical rows of neck cells in all species ， w i th each row having 8 cells in A. reticulata ，A. blumeana and A. mussuriensis. In In A. ca lifornica Campbell (1918) reported 7 and Haupt (1929) 8; while in Rebouli α Haupt (192 1) reported 18-20 in each row. In In Asterella just before the fertilization there appears a perianth around the base base of the archegonium (Figs. 28 ， 29) and after an archegonium is fertilized it grows over it as a conspicuous conical sheath projecting from the carpocephalum. マh 18 18 17 17 19 20

25 24 26

....--.-.+-ー佳話事

32 31 30

Figs. Figs. 17 -32. Development of archegonium. 17 ，九 f. i時d化α18 -23 ，A . blu -

制閥均α 24 -28 ，A. 何 tic 叫 lα ta. 29 ，A. su 冗.gu 'i畑 α 30- 32 ，A. 叩 U88 叫 riensi8. Figs. Figs. 17-29 ， x 425; 30-32 ， x 325. (p - perianth initial ，yp- young perianth ， both in V.8.). V.8.). 10 10 J ourn. Hatlori 臼ot. Lab. No. 19 19 58

Fertilization: Fertilization: Fertilization take s place during the first shower of ra i ns and it it is later that the stalk increases in length. Just before it occurs the neck opens along along the lid cells due to hygroscopic pressure inside ，which forces them apart to let let in the spermatozoids. The latter wriggle their way down the neck till they reach reach the egg membrane . One fertilizes the egg. The zygote enlarges in size till it it nearly fills the cavity of the venter (Figs. 23 ， 29). The la t ter is one layered before before fertilization but afterwards its cells enlarge and divide to form a jacket of seve ral layers around the young embryo. This is the calyptra ，and it persists even after after the spores lie freely within the capsule.

Spo rog oniu m Meyer (1931 ) described filamentous and quadrant types of embryos in Mar - chantiales chantiales and regarded the former as p rimitive and thelatter as secondary. He included included Plagiochasma Mannia ，Reboulia ， Asterella and Fegatella (and probably Cyathodium Cyathodium and Cr y' ρ tomirium as well) in the filamentous type and Marchantia ， Preissia ，Astroporae ， Corsiniacea and Riccia in the latter type. The embryos of the the species investigated here are of the filamentous type ，no exception having been observed . The development of the embryo starts immediately after fertilization. The first first division in the zygote is transverse dividing it more or less equally in Aste- rella rella blumeana and A. sanguinea (Figs. 47 ，64 ，65 ) or unequally in other species Fig. (Fig. 61) ， thus forming 2 cells-an epibasal ce Il and a hypobasal cel l. In A. mus - suriensis suriensis there is an indication that rarely the division may be oblique (Fig. 62). In In Asterella ， Plagiochasma and occasiona Il y in Mannia indica next a transverse division division appears in the epibasal cell dividing the embryo in a3 celled filament (Figs. (Figs. 33 ，48 ，53 ，63 ，66 ，71 ，77) ， in which the lower forms the foot ， the middle seta seta and the upper capsule . A 自lamentous embryo is a!so described for Reboulia ， Asterella ρalmeri ，Cr y' ρ tomitrium tenerum (Haupt ，1921 ，1929 ，1942) ， A . blumeana (Peissel ，1925 )， A. angusta (Mahabale & Bhate ，1945) ， Plagiochasma articulatum Mahaba!e & Deshpande ， 1947 )，Mannia (Meyer ， 1916) and Cry. ρ tomitrum hima- layensis layensis (Chopra & Khosla ，unpub l.). Previously Campbell (1918) described a quadrant quadrant embryo for A. californica and Abrams (1899 ) described the same for C. tenerum . Usually in Mannia indica a4 celled embryo is organi sed (Fig s. 34-36) as in A. californica and C . tenerU1 η(Haupt. 1929 ). Though the early divisions in this filamentous embryo are more or les s variable in in different species they broadly follow the plan described for other Rebouliaceae. After After the first division of the embryo one can easily detect the further divisions in in the 3 ( or 4 in case of Mannia indica ) regions of the embryo (Figs . 34-36 ，66-68 ， 71-73 ). Vertical wa lI s first appear in the capsular region (Figs . 36 ，37 ，50 ，63 ， 71) in in contrast to Reboulia (Haupt ，1921 ) whe re they first appear in the lower tiers; or lI alI tiers may undergo simultane ous divisions (Fig. 67 )， or often the terminal cell of

Figs. Figs. 33-52. Development of sporo gon i um. 33 -46 ，M . i叫dic α 47 -52 ， A. blume α冗α. 42a ，archesporial cell; b，c ，d ivi sion of the same into spore mother cell and elater cel l. 43a ， b，44 ，stages in spiralization in elate r. Fi gs. 33 -40 ，43 ，47 -52 ， x 325; 41 ，x 却 0; 42 ，44 ， 45 ， x 425; 46 ， x 125. t9 月 II U{3，f 113J 年服部他物研究所線 ~1;.ZI\

33 34 36 37 39

45 b 43 α

49 "'w 50 12 12 Juurn. Hattu ri Bu t. Lab . No. 1!:J 1 !:J ，'S ti

5 ラ 56

品川

68 76 66 67 昭和 33 年服部随物研究所報告第 19 号 13 the the embryo is the 1ast to undergo a vertica1 division (Figs. 35 ，49 ，54). Later more vertical vertical walls are laid in nearly al1 the tiers (Figs. 38 ，68 ，72 ， 78). The ce l1 s of the the seta are meristematic for some time and result in the formation of a short seta. seta. In the mature sporogonium usua l1 y it remains short but in Asterella sanguinea sanguinea it is somewhat elongated (Fig. 69) recalling to some extent the seta of M archantia. The basal ce l1 of the embryo does not divide simultaneously with the capsular and the seta initials so that in Asterella blumeana and A. reticulata when the embryo is elongated and many ce l1 s long it is still a single cell ， though enlarged and prominent (Figs. 49 ，50 ，55 ，56). The later development of foot is quick and it it soon assumes a bulbous shape with the periphera1 ce l1 s highly granular and the central central ones less so in the mature state (Figs. 59 ，70). This character is persistent even after the spores are shed. It is believed to be haustoria1 in function though no particular processes are developed for this purpose. By this time the sporogonium is elongated and now the ce l1 s in the capsular region region divide in all directions (Figs. 39 ，39 ，51 ，57 ，74 ，78-80). Later this mass of ce l1 s is di 任erentiated into an outer one cell thick layer and a centra1 archesporium Figs. (Figs. 40 ，52 ，58 ，75 ， 81) and the capsule becomes globular in shape (Figs. 41 ，69 ， 82). 82). The parietal layer is persistent but loses its contents and becomes enlarged by the time the spores are shed (F igs. 46 ，84) ， retaining the nu c1 ei up to the last stage. stage. The cells of the archesporium stain deeper due to their dense cytoplasmic contents contents and have prominent nu c1 eoli. The sporogenous mass is formed from about 2/3 of this tissue and about 1/3 forms the belt separating the archesporium from the the seta. There is no definite arrangement of sterile and fertile cells but they are readily readily distinguishable by their form. The elater mother cells and spore mother cells cells are sister cells (Fig. 42a-c) as in Reboulia and Cryptomitrium tenerum (Haupt ， 1921 ，1942) . The sterile cells are more or less elongated and the fertile ones nearly rounded. rounded. The sporogenesis is as usual for Marchantiaceae. The spore mother cells round round 0 任 and form a tetrad after reduction division. When the spore mature the outer outer cell walls thicken and are later differentiated into 3 layers-the exine develop. ing ing characteristic sculpturing and the perisporium is in form of a sac. The spore wall wall is already 3

Figs. Figs. 53-76. Development of sporogonium (contd.) 53~0. A. 何 ticulat α， 61-63 ， A. mUS8U パ側 8is. 64 -70 ，A. 8 側仰ti-憎 α 71-76 ，A. Pα訪問kote π8i8. 59 ， t.8. foot. 60 ， 76 ，apices of capsules in l.8 . 70 ， t.8. foot in V.8. Figs. 53-68 ，70 ，75 ， x 325; 69 ， x 125; 71 -74 ，76 ， x 425. 14 14 Journ. Hattori Bo t. Lab. No. 19 1 958

The apical region of the capsule-wall in vertical section shows 2 layers of cells in in Asterella blumeana ， A. reticulata (Fi g. 60 ) and A. mussuriensis ， while in A. athankotensis ρathankotensis (Fig. 76) ， A. sanguinea ，Mannia indica and Plagiochasma ap ρendi- culatum culatum (Fig. 83) it is very prominent and 3 cells in thickness. An operculum is present in all the species. In Mannia indica ， Asterella ρathankotensis ， A. sanguinea and Plagiochasma appendiculatum it is detached as a circular circular lid from the capsule. In other species it is detached in fragments. In either either case the dehisced caps u le is left as an urn-shaped structure. The cells of the the operculum show radial thickenings which are more prominent where the lid detaches detaches as a disc. Cavers (1911) described an annulus-like structure in capsules of Mannia ， such a structure is not detected in M. indica. The spores are liberated either singly ，few at a time or in a mass ，depending on the velocity of the currents of air or movements of insects. The elaters help in in dispersal in rains or after the spores fall on moist ground.

Discussion Discussion

IndividuaUty of Rebouliaceae: ー The various genera included within the family family justify their separation in a group on basis of their morphological characters: ters: carpocephalum always stalked ，terminal ，with a single rhizoidal furrow; male receptacle receptacle or cushion truly dorsa l. In Plagiochasma both are dorsal ，and rhizoidal furrow is absent. ( In A . “Manipur" the carpocephalum stalk has air-spaces as well). well). The thallus has usually air chambers in many layers. In all species there is is uniformly a 白lamentous embryo (e xcept for a report of quadrant embryo in Asterella Asterella californica and previous report of a similar embryo in Reboulia and Cry ρ tomitrium tenerum ). Asterella is unique in having a perianth . lt is interest . ing ing to note that all genera have reticulate ，w in ged spores with slight variation in Asterella Asterella where they are occasionally ridged. This is in contrast to the coarsely papillate papillate spores of Sauteriaceae; to some extent resembbling those of Targionia (compare e.g. spores of A. blumeana with those of T. hypophylla ). In the family Marchantiaceae the spores range from smooth to lamellate. Cavers (1911 ) also while discussing their relationships with other genera of Marchantiaceae concluded that the "group may be regarded as forming an independent series showing some striking parallelisms with Astroporae on one hand and the Compositae on the otne r." He (l .c. p. 195 ) used the family name Ayto- niaceae niaceae since he regarded Plagiochasma (=Ay tonia ) as the most primitive member of of this family. Evans (1923 ，5 p. 9; 1939 ) chose Reb ouliaceae. 6 Plagiochasma ， though retaining the well developed carpocephalum ， is specialized due to the dorsal nature nature of the reproductive branches and the consequent loss of the rhizoidal furrow.

The correct citation for the name Rebouliaceae should be : Rebouliaceae (Reichenbach ) Evans ，1923.-Proskauer (in litt .，1957). 6 While the name Aytoniaceae Cavers ，1911 ， p. 195 ，is the correct name and has priority ority over Rebouliaceae Evans ，1923 ， in North American Flora 14 : pt. 1， p. 9， the latter is here here employed. The name Aytoniaceae is based on a synonym rejected as a homonym and the the case has been drawn to the attention of a memb er of the International Committee on Nomenclature Nomenclature of Bryophytes for f ¥，l rther action. 昭和 33 i' 手服部他物研究所報告げ~ 19 J，} 15

The only character that might be regarded primitive in Reboulia is the aggrega. tion tion of antheridia in recepta c1 es ， instead of a cushion ，and compound pores on the same! Asterella might with justification be regarded as primitive since in a few species species it has retained air chambers in one layer with assimilatory filaments ， the perianth perianth is present in all its species. Thus ， with regards to primitiveness of any one of these genera one has to be cautious since the various genera show striking parallelisms parallelisms with each other and the ‘position of primitive genera should be based upon a combination of character rather than upon a single character (E vans ， 1939)¥ Position Position of sex organs: - The position of male and female recepta c1 es jcushions on the thallus within the various genera and occasionally within the same species is is variable as shown below. (The species marked with an asterisk were studied by the author for this feature) .

Se x distribution Position of sex organs Species

Dioecious Dioecious e le dorsal; 室 stalked …terminal. | A.mgustα ホ il A … a*. A “Manip 日 A Monoecious Monoecious ♀ stalk 叫 terminal; アづ 222fr 守おう込 35422 ヤgf 合 dorsal ，just behind it. í;~eìí α ， A. palmeri ; M. rupe8t 付 8; c. l teneru 肌 c. himalaye 倒♂.

|♀ s坑御州凶ta a討l附 t匂町町e釘erm …m A. r・eticul αt ♂ M. pilosα R. hemis. } 会 on ventral shoots. pha 町化計.

A. pαth α叫 kotensis ホ， A. S. αnguine α* "♀ stalked ， on ventral A. pannpora ，A. muUifiora; M. i四- shoots ，合 aplcaιdorsa l. dica ヘM. cαlifor ・洞化 a. ， ♀ stalked; ♀ and 会 ter. minal minal on ventral shoots. 。A. mU8su 付en8is* ，A. bol α骨 d げぜ. ♀ stalked ，terminal on one M. fragra 畑 R. hemisphae γica* ; P. lobe; 合 dorsal.terminal on " another lobe. cune αtum ，P . simle 何 8e.

会 and ♀ dorsal on main I P. αppeπdiculatu骨~*， P .町 ticulatum ‘， ~ thallus. thallus. P. rupestre ，P. 加 te 門司edium.

A-Asterella ，M-Mannia ，C-Cryptomitrium ，R - Rebouli α，P - Pl αgiochasm α.

In In A. mussuriensis where the sex organs are restricted to the ventral shoots occasionally occasionally the stalked carpocephalum is borne apica l1 y on the apical continuation of of the main thallus. A. sanguinea also exhibits this reversal in position of male and female recepta c1 es. Here the carpocephalum is borne on the ventral sh ∞ ts and the male cushion (occasionally a faIse recepta c1 e) is apicaI on the main tha l1 us. But there are cases where the male is borne on a Iateral elongation of the apex and the apical.ce l1 .region is continued as the stalk of the carpocephalum ，which thus thus occupies a terminal position. It yet another case the carpocephalum was terminal terminal and the male was borne on a small ventral shoot arising from the ventral s ide just near the region from where the carpocephalum arose. This interesting inter. 16 16 Journ. Hattori Bo t. Lab. No. 19 1 958 shifting shifting of sex organs from their normal position points to the fact that probably the the ancestoral feature was that both the sex organs were terminal on the thallus but but later in the course of evolution they shifted to the ventral shoots. Thus ，on this this feature the species with both sex organs on the main thallus are primitive and those those with either of these on the lateral sh ∞ ts derived; those with 加 th on the

77 77 78

曲、品、- 1k

a 、久 o ，" CQJ ， λ^~Qコ o~ ~&? ~ - 0 -

Figs. Figs. 77 -84 . Development of sporogonium (contd.) P . αppendiculatum. 83 ， apex of capsu le in l.8. Figs. 77-80 ，84 ，x 390; 81 -83 ， x 240. RH 平11 33 i!，. 服部情物研究所報告ー節目号 17 leteral leteral sh ∞ ts still more removed from the ancestoral type. In the latter group this this specialization may have resu It ed in great morphological advantage over the other other species ， since the apical cell is free to undergo unlimited vegetative growth and may go on producing side by s ide numerous sexual shoots as wel l. It It is interesting to note that in the family Sauteriaceae (of Evans ， 1939) Sau- teria teria has similarity with Aslerella blumeana in position of sex organs on the thallus ， air air chambers in many Iayers ， stalk without green tissue and with a single rhizoidal groove . In Peltole ρis the male and female recepta c1 es are terminal on adjoining lobes lobes and air chambers in many layers as in Mannia fragrans; but the stalk though without green tissue ，has occasionally two rhizoid furrows. In Athalamia ， though the female recepta c1 e is dorsal without a gr ∞ve in the stalk ， as in Plagio- chasma ， the male recepta c1 e is reduced ，has become indistinct ， irregular and is without without surrounding bracteoles. These facts bring forth striking parallelisms be - tween the two families rather than showing a merging of one into another. R eceptacles: - The carpocephalum is conservative in having retained its stalked nature ，compound pores ，and probably in some species of Asterella the presence of of air chambers in one layer with assim i¥ atory filaments. It has undergone reduction tion in nature of air chambers in most of the species ， reduction to one rhizoidal groove groove in Mannia ，Asterella ， Cryptomilrium and Reboulia; and to its total absence in in Plagiochasma. In In A. "Manipur" (Figs. 88 ， 90) and A. blumeana (Pande et a! ， 1953) the stalk of of the carpocephalum is occasionally branched into two ，each branch bearing apically ly a female recepta c1 e in the usual fashion. Often in most of the Rebouliaceae the the number of involucres may be as great as 9， but the stalk remains always with a single furrow. However ， A. blumeana (Pande & Srivastava ， 1953) shows a double furrow in the stalk of the carpocephalum ，which is a feature of Compositae; further further a reduction from 2 to one furrow in the same stalk and in one case ac- companied by a dichotomy of the stalk at its apex! A. “Manipur" also shows the various various forms of this reduction from 2-1 furrow ， but in different plants and without out any correlation with the bifurcation of the stalk. Furthermore ， this species shows the presence of air spaces (usua lly 3) towards the posterior region of the stalk stalk (Fig. 91) ， thes

7 While th is pa 伊 r was in Mss. form Pande et al (1953b ) published an excellent account on on some abnormal female receptacles of Asterella kh αsian α， which makes it un.necessary for for me to append a similar account 1 wrote on the basis of abnormal female recepta cJ es of A.

“Manipur" (F igs. 86 -90 ) and A . 何 ngui 四 α(Fig. 92 ). Reference may also be made here of the the fruitful discussion on the Reduktionen in den Geschlechtsst 詰nden in Marchantiales (Burge 仔， 1943 ， pp. 244 -2.'>3)

8 A. Cαlifo ゲnic α (Campbell ，1918 )，RelJOul ω (Cavers ， 1904) and C. te 冗伺-u m (A brams ， (1899 ) were previously described with quadrant embryos; but on reinvestigation Relouli αand C. C. tenerum were found to have 自lamentous embryos. 18 18 Journ. Hattori Bot . Lab . No . 19 1 95 8

86 87 85

91 91 α

‘ .一・‘ム. . ‘' 6866bO 昭和 33 1f~ 服部槌物研究所報告第 19 号 19

In In Corsinia ，Sauteria ，Exormothec α，Ste ρhensoniella ， Preissia and Marchantia the embryo is of ∞ tant-type. Meyer (193 1) regarded the filamentous embryo as primi - tive tive and the 促 tant type advanced. (l t appears that linear phylogeny of Marchan ・ tiales ， ascending or descending ，no longer deserves serious consideration). However ， it it may be noted that Corsiniaceae and Sauteriaceae have an octant embryo; Tar- gionaceae gionaceae and Rebouliaceae a filamentous type; and Marchantiaceae mostly an octant octant type and occasiona l1 y a filamentous one . Thus ，a filamentous type can not be visualized to have been sandwiched between an octant and octant type. It would be best to visualize that in each group this development is independent of other other features. Incidenta l1 y it may be noted here that in Ricciocarpus and Riccia both both type s are met with. Concluding remarks: -Ac cording to Cavers (1 911 )，Kashyap (1919) and Frye and Clark (1937) Reboulia and Plagio chasma are derived one from the othe r. The apparent apparent relationship between the two is very great but it is to be noted that they differ differ a great deal in morphology. Plagiochasm αis apt to be regarded as a derived genus or at least as 'later evolved' as shown by its thallus and position of sex organs. organs. But the two might have descended from a common ancestor and followed parallel parallel lines of evolution. Asterella in the presence of a perianth resembles Mar - chantia chantia and Preissia and on this basis most clearly approaches the ancestoral form from which the Marchantiaceae have descended (Goebel ， 1930)_ This suggests that Asterella Asterella is a comparatively primitive genus and could not have been derived from Plagiochasma through either Massalongoa or Mannia and Cryptomitrium. It may have ancestors similar to those of Reb oulia- Plagiochasma group ，even though it di 任ers from them in spore structure ， nature of germ disc and in many species in the the structure of the thallus. Cavers (1911) derived Mannia aIso from Plagiochasma through through Reboulia while Frye and Clark (1937) did so directly from Plagiochasma. This is unsound. It could not have evolved from a genus with dorsal sex organs and the usual absence of ventral shoots. It resembles Asterella in its plan of spore ling germination and in the range of vegetative structure as revealed by the various various speci es of each genus. Resemblance s between some species are so great that that before a perianth appears it is difficult to differentiate one from the other (c f. A. ρα tha

Figs . 85-92. A8terell α “ Manipur ". 85 ， plant showing horse -s hoe shaped male re. ceptacle ceptacle an d continuation of apex into a heart-shaped lobe bearing carpocepha lum . 86 ， same ， note subsessile carpocephalum and dorsal involucre (di); the latter in V.8. in 87 (dia- grammatic). grammatic). 88 ， showing sessile carpocephalum on left side and a dorsal involucre in the one one on the right side; the former in 1人8. in 89 (diagram. ). 90 ，s howing concentric ma¥e receptacle receptacle and bifurcation of stalk (a lso in 88). 91 ， t.8. stalk show ing air spaces (α) and rhizoidal rhizoidal groove ， both double in b. 92 ， A. 8αngui7 恒久 a sma l1 tha l1 us with sessile carpoce- halum. halum. Figs.85 ，86 ，88 ，90 ，92 ， x4; 87 ，89 ，x lO; 91 ， x125. ~

Dioecious ， air chambers in one layer ， assimilatory filaments present ; sex organs terminal ，合 sessile receptacle ，♀ stalked; 1 rhizoidal furrow ; perianth present.

Tendency towards bisexu 直 lity ， loss of assimilatory fila. ments and subsequent increase in layers of air chambers.

oC】『ロ・出血丹件。ユ切 Estab 1i shment of bisexuality ，development of ventral ventral sh ∞ts ， restriction of either sex to iド片(ω如制凶釘出向…i均ian 叩n t一糊叫tra …oぽr absent ，…bisexua l. apex of thallus ，会 cushion; perianth present. on - - とー yZ0 ♀ or 会 1 1 0 'a~~: 前向 1 11 ♀ terminal ，会 sex organs Reduction Reduction of dorsal ， loss tMminalm ll zdmal 勺 1j ・合 to cushion Asterell α 1 1 ，5? _，，"- v: _oa. ， 1 1 thallus or on of rhizoid 司l 同但 (A. α匁伊t8 ta ). ventωshoots l| (qvt 帆ι11 ;~.~t凶油∞ ts groove (到α- (Mannia ). 1 1 …… J・ 11 (Re 加山.α ). gioch ω ma). f、ご¥ ♀ terminal ，日♀ terminal ， S;h~ 1 1 ♀ 刷会 11 合 terminal ，会 dorsal ， 11 合 dorsal v entral 1 1 terminal on 1 1 ♀ on stalk stalk with air 11 behind ♀ shοots adjoining 11 v entral spaces spaces (A . 11 re 四 ptacle (A. (A. mua . 1 1 lobes (M. 1 1 s hoots “Man ipur ") .11 blu me ωω ). S urie 畑 is ). I I jlT 吋 rans ). II (M. 伽 dica ). 戸埠印∞ 昭和 33 ;1'. JI~~ilfttI物倒究所報 ~ ' i'~; 19 J，j 今 21 occasional occasional presence of air chambers in one layer with assimilatory filaments in Asterella. Asterella. The removal of Cry ρtomitrium from Rebouliaceae to Marchantiaceae (Verdoorn ， 1932) is also uncalled for ， since this genus is so much like Rebouliaceae that that such a doubt should not have arisen; e.g. in its filamentous embryo ，a definite lid lid in the capsule ，structure of male cushions and female receptacles and thallus morphology. In con c¥ usion the accompanying table is presented embodying these results and speculations. 9

Summary

Life Life history is studied in M αnni αt刊dic α，A8terell α blu η1.e απ a ，A. reticulata ，A. mU8 ・

8urien8 匂，A.8 側 .gui 刑 α，A. P αtha 叫 kote 叫8i8 and Pl αgioch α8m ααppendicul α.tum. Develop- ment of sex organs is of the general type in Marchantiales. The embryo is filamentous: 4・ celled celled in M. indic αand 3.celled in other species. The various genera on basis of thallus morphology ， nature of reproductive organs and spore character justify their inclusion under a single family. Asterell αand M. αn '11. iαshow striking parallelisms in position of sex organs ; and in the former genus variation and interchange in position of sex organs (receptacles) on the the thallus ， as shown by various species ， suggest that in ancestoral form both were terminal on the main thallus. A. sa '11. guine αand A. “Manipur" have rarely sessile carpocephalum ， the latter showing occasionally occasionally a branched stalk and usually abnormalities in the number of involucres. The inter-relationships inter-relationships of the various genera are discussed . 1 am indebted to Pro f. P. N. Mehra for having suggested this problem and for his keen interest interest in me in general and in this problem in particular; to Prof. J. Proskauer for his criticism criticism and for suggesting a few modifications incorporated in the text; to Profs. W. C. Steere ，A. J. Sharp and Dr. T. Herzog for encouragement; to Mr. T. N . Khoshoo for help- ing ing me in a number of ways; and to my wife for her appreciation ， cooperation and encouragement. ment.

Literature Literature cited

Abrams ， L. R. 1899 . The structure and development of 0ヤyptomitrium ten 肘・um. Bot. Gaz. Gaz. 28 : 110-12 1. Ahmad ， Sultan. 1938. A study of Aiichisoniell αhim αlα Ye l'l sis. Proc. lndian Acad. Sc i. 7: 206 -224. . 1940 . Morphological study of Exormotheca tuberifer αBot. Gaz. 101 : 948-954.

Andersen ， E. N. 1929. Morphology of sporophyte of M 即 ch α'11. tia dom 伽 g側劉s. Ibid. 88: 15 0- 166. Burgeff ，H . 1943 . Genetische studien an M αγ ch αnti αJena. Campbell ， D. H . 1918. Structure and Development of Mosses and Ferns. New York. . 1939. Embryophyta. Stanford.

9 Burge 任 (1943 ) in his Genetical Studies in M αrch α'11. ti αpresented a Schema der Mar. chantiales chantiales (p. 241 ， f. 213 ・reproduced here ) in which he suggested a polyphyletic origin of most of of the genera of the Marchantiales (Fig . 93 ). Reboul 必，Asterell αand M. α'11. n仰 are derived di stantly from Plagi ∞hα S1n αthe latter genus shares it s ancestory with Clet'e αand Athal α- mω. The genera Pel ωle pi8， & 山 ter 似，Exor 悦 othec α，Pl αがoch αsmα ，Cleve αand Athalami α， form a distinct ‘ line' in his scheme. 22 22 Journ. Hattori Bo t. Lab. No. 19 1 95 l'l

Cavers ， F. 1903 . Explosive discharge of antherozoids in Feg αtell αconic αAnn. Bot. 17 : 270-274. 270-274. . 1904. Contributions to the biology of the Hepaticae. Leeds. . 191 1. The inter.relationships of Bryophyta. New Phyto l. reprint no. 4.

Chopra ，R . S. and S. Khosla. Unpub l. Embryology of Cryptomit1 ・ium hi 例 αlayense Kash. (M . Sc. Thesis ，Panjab Univ. 1949 ). Dupler ， A. W. 1922. Ea r1 y embryogeny of Re 初旬li αhemisph αerica. Bo t. Gaz. 74: 143- 157. 157.

. 1922. The male receptacle and antheridium of Rebouli α he 悦 .isph αerica. Ame r. J. J. Bo t. 9: 285-295 Durand ， E. J. 1908. The development of sex organs and sporogenesis of l'if arch αnti α poly-

骨ωγ ph αBul l. Torrey Bot. Clb. 35: 321-335. Evans ，A. W. 1939. The c1 assification of Hepaticae . Bot. Rev. 5 ・48-96. Frye ，T. C. and 1.. Clark. 1937. Hepaticae of North America north of Mexico. Univ. Washington Pub l. Bio l. 6: 42 - 102. Goebel ，K. 1905. Organography of Plants (E ng. ed.). Oxford. 1930. 1930. Organographic der Pflanzen. Zweiter Tei l. Bryophyten-Pterodophyten Dritte Auflage. Auflage. 643-1363 . Haupt ， A. W. 192 1. Development of gametophyte and sex organs in Reboul ω hemisphae- ric α Bot. Gaz. 71 : 61 - 74. 一一一一一 . 192 1. Embryology and sporoge nesi s in Rebo uZ ωhemisphae γ化αIbid. 71: 446- 453. 453. . 1926 . Morphology of Preissi αqu αdr αtαIbid. 82: 30-54 . 一一一. 1929. Studies in Californian Hepaticae. 1. Asterell αcalifo 門前αIb id. 87: 302- 318. 318. . 1942 . Studies in Californian Hepaticae. 11. Cγ yp ωmitrium te 叫酔-u m. Ibid. 104 : 264-272. 264-272. Howe ，W. A. 1899. Hepaticae and Anthocerotes of Californica. Mem. Torry Bo t. Clb. 7: 7: 1-208. Kachroo ， P. 195 1. A s imple suction pump . Curr . Sc i. (l ndia) 19 : 113. . 1954. a. Distributi on of Rebouliaceae in Indi a. The Bryologist ， 57: 159 - 166.

. 1954. b. Morphoolgy of Rebouliaceae . II. On some species of Mannia ，Aste 何 lla and Pl α:;ioch αsm αJourn. Hattori Bo t. Lab. 12: 34 - 52. Kapoor ， A. S. 193 1. A study of Pl α.qioch ωmααγ ticul αtU 7n. Proc. Indian Sci. Cong. 28: 2臼. Kashyap ， S. R. 1914 . The androecium of Pl αgioch αsm αα ppendicul αtum 1.. et. L. and P.

αγ ticul αt 叫 m Kashyap. New Phyto l. 18 : 235 - 238. . 1919. The inter .relationships of liverworts especially in the light of som 邑 recen t1 y discovered discovered Himalayan forms. Proc. Asiatic Soc. (Be ngal) 15 : c1 ii. c1 xv i. Kienitz.Ge r1 o仔， F. 1874. Vergleichende Untersuchungen uber die Entwick lungsgeschichte des

Lebermoos-Sporogoniums. Bo t. Zeit. 32: 161 ー172 ，193 -204 ，209 -217 ，225- 幻 5. 事 . 1875. Neue Bεitr 品ge zur Entw icklun g des Lebermoos.Sporogoniums. Bo t. Zei t. 33: 33: 777-782 ， 793-799.* Leitgeb ，H. 188 1. Untersuchungen uber die Lebermoose. V I. Die Marchantieen. Graz .* Mahabale ，T. S. and P. D. Bhate. 1945. The structure and life history of Fi 7n briarin angust αS t. Journ. Univ . Bombay 13 :5 - 15.

一一一一一 and S. R. Deshpande. 1947. Life history of Plagioch αS 悦 ααγ ticulatum Kash. Ibid. Ibid. 15: 23-37. McNaught ，H. 1.. 1929. De velopment of sporophyte of March α叫的 chenopod αBo t. Gaz. 88: 88: 400 -4 16. 昭和 33 (1 111 [1; 111111 物併究所W ;' i' zn19 号 23

Mehra ， P. N. and H . L. Mehra. 1939 . Life history of Stephensrmiella uret ヤedunculat 品 Kash. Kash. Proc. Indian Acad. Sci. 9: 287 - 315. and P. Kachroo. 195 1. Sporeling germination studies in Marchantiales . I. Reboulia- ceae . The Bryologist ， 54: 1- 16. Meyer ，K. I. 1916. A study of sporophyte in the liverworts of the group Marchantiales (in Russian ); vide Campbell ， 1918.* 193 1. Die Sporophytenentwicklung und die Phylogenie bei den Marchantiales.

Planta Planta 13 : 210 -220 .ホ Pande ， S. K. and K. P. Srivastava. 1953. Two more abnormal female receptacles ef A8terel- la la bl 初旬eα .n αNees. Journ. lndian Bot. Soc. 32: 137 -14 1. and S. A . Khan. 1953a. On some abnormal female receptacles of A8terel- la la blume αnαNees. Proc. lndian Acad. Sc i. 38: 21-26. 一一，一一一 and 一一 . 1953b . On some anomalous receptacles of Asterella kh α8i - α .na. The Bryologist ，56 : 229 -24 1. 一- and 一一一一一 . 1954. On some species of Asterell αBe auv. Journ. Hat- tori tori Bo t. Lab . 11: 1- 10. Peirce ， G. J. 1902. Forcible discharge of antherozoids in Asterell αωlifornic αBu ll . Torrey Bot. Clb. 29: 374 - 382. Peissel ， Ruth. 1925 . Bau und Entwicklungsgescichte von Fim~イαパα blumeanαGottsche. Bo tan Arch. 10: 434 -4 76 .* Star ， A. M. 1916. A Mexican Aytoni α Bo t. Gaz . 61 : 48 - 58. Verdoorn ，F . 1932. The classification of Hepaticae ， in Manual of Bryology ，The Hague. Woodburn ，W. L. 1919. Preliminary notes on the embrology of Reboulia hem 俗ph αeric α. Bull. Bull. Torrey Bo t. Clb . 46: 461 -4 64.

. 1922 . Spermatogenesis in Asterella hemisphaerica. Ann. Bot 目 36 : 535- 539.

APPENDIX

1. 1. Notes 00 Aytooiaceae.

During my previous investigations 011 the morphology of this family 1 chose the family name Rebouliaceae. However ，when the last of this series was sent to Dr. J. Proskauer for his kind criticism ，he drew my attention to the impropriety of such a selection ，since Aytoniaceae has priority over Rebouli aceae ，as a family name ，and ought to have been employed. This correspondence resulted in the following compilation ，( mainly from Proskauer's letters to the writer) ，which gives the various names used for this group in the pas t. Rebouilliariae Rebouilliariae H. G . L. Reichenbach ， 1837.

Grimaldieae Grimaldieae Reichenbach in Rabenhorstschen Kryptogamen ・Flora ， ed. 1， 1848; ( the original date date in 1841-Proskauer 初 lit. ) Operculatae Operculatae Leitgeb in Untersuchengen uber die Lebermoose ， V I. Die Marchantieen. Graz ， 1881 . Aytoniaceae Aytoniaceae Cavers in New Phyto l. rep r. no. 4， p. 195 ，191 1. Rebouliaceae Rebouliaceae Evans in North American F lo ra 15 (1):9 ， 1923 ( the correct citation should be Rebouliaceae Rebouliaceae ( Reichenbach) Evans 1923 - Proskauer 伽 lit. ). Asterelleae Asterelleae Jorgens en in Norges Levermoser ，Be rgens Mus. Skrifter Nr . 16 : 1-343 ， ηtit 25 Tα f. Be rgen ， 1934. Grimaldi a ceae (Reichenbach ) Muller in Rabenhor st Kryptogamen-Flor a 6: Erg anzungsband : 220 ， 1940. * Not seen in origina l. 24 24 jりU rl1 . llallori Bo t. Lab. No. 19 J !) !i 8

2. On Aswrella ev α nsii Kachroo.

1n 1n 1917 Kashyap described Asterella (= Fimbriari α ) reticulala with the fo Jl ow- ing ing characters: the thallus monoecious with air chambers in many layers and with simple simple pores; the ventral scales with single appendages; the antheridial cushions on very short lateral shoots arising on both sides of the rnidrib; the stalked car- pocephalurn pocephalurn terrninal on the main thallus ， the stalk with a single rhizoidal groove; and the spores brown ，reticulate-larnellate: reticulations obscure ，and with narrowly finely finely punctate wing . However ，Kachroo (1954 ) described the rnale cushion as often often found on lateral shoots that are OOrne apically and the exine of the spore decidedly decidedly with larger fewer reticulations inclosing rnany smaller reticulation. The species species is closely related to A. blumeana (Nees) Kachroo with which it agrees in its its spore structure ， thallus morhology and chromosorne number (n = 18 in OOth ， Mehra and Kachroo in press). Further it is not uncornmon to find specirnens intermediate intermediate between them ，and it is possible that A. reticulata is a derivative of A. blumeana. Evans (1920 )，presumably unaware of A. reticulata Kashyap also described a new species of Asterella which he also narned A. reticulata. Evans' “ A. reticulata" is is very close to A. elegans in the general appearance ， in the thallus structure and the the rnale inflorescence; the only difference between the two being spore structure ， which in case of A. elegans has a regular reticulum and in case of “A. reticulata " larger larger regular reticulations including smaller irregu l ar rneshes (c./. Evans ，l.c. ). Since Since A. reticulata (Kashyap ) Kachroo is described eariier ， it has priority over “A. reticulata" Evans. Therefore ， A. ev α nsii sp. n. is proposed to accommodate the the latter species. (Narned after Pro f. Alexander W. Evans). However ，a study of of the four species: A. blumeana ，A. reticulata ， A. evansii and A. elegans might be interesting from cytological and distributiona l points of view. Evans ，A. W . 1920. Contrib. U. S. Nat . Herb . 20: 302 -303 . Kachroo ， P. 1954. Jour. Hattori Bot . Lab. 12 : 38 -39 ，fig .3 ，a -1. Kashyap ， S. E. 1917. Jour . Bombay Na t. Hist. Soc. 25: 279; 1929- Li verwort s of the Western Himalayas Himalayas and Panjab Plain ，1: 65 ， p l. 14 ，figs. 1 -3. Mehra ，P . N. and P. Kachroo. Cytology of West Himalayan Marchantiales (in press ).