植物研究雑誌 J. J. Jpn. Bo t. 66: 66: 134-146 (1 991)
Studies in the Systematics of Filmy Ferns VIII C 同phalomanes Presl subgen. Cephalomanes
Kunio IWATSUKI
Botanical Botanical Gardens ，Faculty of Science ，University of Tokyo ，
3-7-1 ，Hakusan ，Bunkyo ・ku ，Tokyo ， 112 JAPAN
コケシノブ類の分類に関する研究班 ソテツホラゴケ亜属
岩槻邦男
東京大学理学部附属植物園 112 東京都文京区白山3-7-1
(Received (Received on January 7， 1991)
A filmy fern subgenus ， Cephalomanes subgen. Cephalomanes ， was taxonomically revised. Taxo- nornic nornic characters were surveyed and re-evaluated from systematic viewpoints. Six species including infraspecific infraspecific taxa were recognized and keyed ou t. Enurneration of all taxa was given with notes on
distribution distribution and taxonomy. C句phaloman 回 crassum and C. madagascariense are restricted to particular
localities localities and are rare with a few collections. C 句phalomanes densinervium and C. singaporianum are also also restricted to narrow areas and are not variable ， but C. atrovirens and C. javanicum have wide areas areas of distribution and are variable with many infraspecific taxa. Rheophytic nature ， or adaptation to to the habitat along stream ， is discussed in relation to habitat and phenetic features. (Continued (Continued from Acta Phytotax. Geobot. 35: 165-179 ， 1984.)
Cephalomanes was described by Presl in 1843 following to Copeland in its position. Most of the on the basis of C. α trovirens and some ten species species have been ill-defined ，and a revision is have currently been referred to subgen. necessary for this complex group. Cephalomanes (Iwatsuki 1984) from the Old In the present part of this series ， subgen. World tropics: East Madagascar ， India to con- Cephalomanes will be treated in its strict sense ， tinental tinental Southeast Asia ，Hainan ，Taiwan ， southern though the generic classification was revised and Ryukyus ，throughout 恥1alesia to Polynesia ， south broadly circumscribed leaving Cephalomanes s. str. to to Queensl 加 d. Copeland (1 938) sugg 回 ted th 瓜 this as a subgenus (l watsuki 1984). Before going further was derived from the Vandenboschia radicans on each species ，it may be contributive to note a group and placed it next to M 白 opteris and few distinct features of subgen. Cephalomanes in Callistopteris. 恥10rton (1 968) ranked it in a section general to elucidate the systematic evaluation of of Trichomanes subgen. Pachychaetum 如 d placed their characteristics. it it together with sects. Nosopteris and Callistopteris ，
句、A 『 d June June 1991 Journal of Japanese Botany Vo l. 66 No. 3 135
The specimens of most larger herbaria were The pinnae are principally subentire to denticu- examined examined on the way of this study. Sincere thanks late ， and the dentation at margin is uninervate at are are due to the directors and curators of these least at its end. Th e uninervate dentation is in most
herbaria ，some of which are recorded in the tex t. cas 岱 very short and hardly waved at margin; in The author is grateful to Dr. 恥1asahiro Kato who some cases ，however ， the dentation elongates ， or read read the manuscript and gave it valuable is separated from at the base or near costae ，
comments. comments. A p 紅 t of this study was supported by forming very narrow or linear segments like those a Grant-in-Aid for Scientific Research no. of Macroglena in appearance. Lamina is nearly 02454017 from Ministry of Education ，Science and lacking on the basiscopic side ， where several hair- Culture. Culture. like segments ， or linear uninervate dentations ， occur. occur. The origin of this type of variation is
Taxonomic characters unknown ， but there 紅 e various intermediate forms Frond construction which suggest that this type of variation is under
The fronds 紅 e simply pinnate throughout ， the in f1 uence of environmental conditions. The
except except for C. crl α ssum in which free pinnae 訂 eonly hair-like segments are present throughout subgen. a few at the base of a frond and more than 20 Cephalomanes ，and there is no specific difference lateral lateral pinnae are adnate to the rachis ，giving an of this type of variation observed. The linear appearance appearance of pinnatifid construction. The adnate segments may suggest the relationship of this group b 出 e of pinnae is contiguous to that of neighboring to Macroglena ，and the other features ，including ones ，and the rachis is broadly winged throughou t. soral morphology and cell-walls ， are not incon- In In the other species ， the rachis is usually described sistent with the suggested relationship. as as terete ， but in fact it is very narrowly winged at The species of Cψ halomanes usually become least least in its upper half. The wing is more distinct soriferous at a very young stage ，and even a small in in smaller plants ，and young leaves usually have frond bears sori rather commonly. In the small distinct distinct wings on both sides of a rachis. plants in which there are little development of Lamina segments dentation at margin of lobules ， the sori appear In In sterile segments ， the margin is usually sub- comparatively large and the sinus is distinc t. entire entire to denticulate or variously lobulate. In fertile Venation segments ，sinus 紅 e often formed and sori are The venation pattern is rather difficult to trace placed placed at their bottom. The linear segments (see in pinnate fronds because of various modifications. the the next paragraph) 紅 e often separated from the In all the species of Cψ halomanes ， venation is laminae laminae next to the sinus. Even in the sterile anadromous and ， as the species with this venation segments ， the sinus is sometimes formed 泊 a place in general ， the sori are paratactic in 紅 rangement equivalent equivalent to the sinus of fertile segments. (cf. Prantl 1875 ，Morton 1968). In dwarfed However ， in portions without any sori ， as seen in species ，it is rather difficult to distinguish the vena-
the .the distal portion of C. singapori α， num ， there is no tion type ， but the general tendency Can be traced laceration laceration of pinnae or at most minute denticula- especially when the soral position is also taken into tion. tion. The denticulation is distinct in this section account.
創 nong the filmy ferns generally with univeined By simply pinnate construction of fronds with teeth. teeth. lanceolate outline ，Ceph α lomanes subgen. 136 136 植物研究雑誌第66 巻第3 号 平成3 年 6 月
Cephalomanesis Cephalomanesis comparable with the New World tics. Copeland (1933 ， p l. 53) illustrated the group group of filmy ferns referred to Trichomanes different distribution of sori on one pinna for five subgen. subgen. Trichomanes. However ， the latter is species. If we tre 剖 only typical extremes we can distinct distinct in having catadromous venation and distinguish different forms: 1) When sori 紅 eafew epitactic 紅 rangement of sor i. In some Am erican on a pinna ， they are at the distal end or on the species species with dimidiate basiscopic base of pinnae ， acroscopic side near distal end; on a pinna with the the venation is rather irregular and needs further several sori ， they are on the acroscopic side from observation observation especially on young plants. The distal end towards acroscopic base; and in case in difference difference in frond construction seems to indicate which there are more numbers of sori ，th 句T are also a systematic remoteness between Trichomanes on the basiscopic side. 2) Wh en sori 紅 ea few on subgen. subgen. Trichomanes and Ceph α lomanes subgen. a pinna ， they are on the acroscopic side usually at Cephalomanes ，and the former may better be midst between distal and basal ends; and even in separated separated from the latter and its allies at generic case there are more sori on a pinna ， they are rank rank (Iwatsuki 1984). restricted to the acroscopic side ， though the sori Distribution Distribution of sori are observed even at the distal end. This typical
The sori are distributed on a frond from the difference is observed in most distribution 紅 eas ， apical apical portion downwards ，except for a case in C. but my field observation does not admit this foersteri ，a problematic form which was described difference as to hold a specific difference. The
ぉ having sori only on the middle and lower pinnae. distribution of the sori on a frond is rather Sori Sori are sometimes borne only at the very apical variable ， and it is hardly concluded from the varia- portion portion of fronds forming spike-like structure ， tion of phenetic characters that the above tendency apparently apparently relating to plant size. There are several is genetically fixed. The two varieties 紅 e geo ・ species' ‘species' distinguished only by this particular graphically isolated in general but coexist in character ， in addition to the others ， such as T. Borneo ， where this feature is more variable. acrosorum ， T. boryanum var acranthum ， T. kingii ， In C. singaporianum the distribution of sori on and T. sumatranum. a pinna is different from that of the other species. Plants Plants with the sori gathering to the apical part The sori have a tendency toward being distributed of of fronds in most cas 白 have larger involucres and at the basal portion of a pinna: most sori are on long long stalks. It is unknown whether they are under the basal half of acroscopic side of pinnae ，and an an influence of the position of sori or not ， though even in fully fertile plants with sori on both the the general tendency is obvious in the species of acroscopic and basiscopic sides ， most sori distri- subgen. subgen. Cephalomanes. The combination of these buted in basal and middle portions of pinnae ， never characters ，larger sori with long stalks gathering on the distal end. This is 仕ue also in the case of at at apical portion of fronds ， forms a spike-like C. densinervium. In these species ， the distal part construction construction of soral region. of pinnae is subentire or at most denticulate at The distribution pattern of sori on a pinna is margin ， even though there are many linear another another character to identify the speci 回 ofsubgen. segmen 臼 in the basal and middle. parts on both Cephalomanes. Cephalomanes. Cephalomanes javanicum var. acroscopic and basiscopic sides. In C. asplenioides asplenioides was distinguished from v紅 .javanil ωm singaporianum the sori are in notches on pinna- by this feature in addition to the other characteris- margin and the mouth of involucre is usually June June 1991 Journal of Japanese Botany Vo l. 66 No. 3 137 situated situated at a level lower than the apex of lobes ， tion was taken from embarkment in damp shaded or or the sori never extruded beyond the pinna- ravine (no ecological data for type collection). 1 m 町田n. have never seen C. densinervium actually growing Morphology of sori in the field ，and its labels often read on slopes or The sori of subgen. Cephalomanes are typically even at ridge. Apparent resemblance of general cup-shaped cup-shaped with very long extruded receptacles. habit of this species with C. singaporianum may The mouth is truncate to more or less dilated. The result from the habitat similarity. All the other degree degree of dilation at mouth has been a most con- species have pinnae more or less oblique to the venient venient key character to distinguish the species of rachis ，and pinnae are lanceolate with narrowly subgen. subgen. Cephalomanes. There are various degrees cuneate bases. C 句phalom αnes javanicum var. of of dilation ，and typological discrepancy can hardly sumatranum always grows on rocks in stream-beds be be available if we treat all the materials from or along streams and is an obligate rheophyte. various various areas ， although the difference in degree of The pinnae are placed obliquely to the rachis and dilation dilation is clear when the materials from particular are narrowly lanceolate. The segments are often distribution distribution area are taken into accoun t. deeply cut at margin and give an appearance of The size of sori is not very variable ，compared bipinnatifid fronds ， though the incision is quite to to the size of plants. The sori are generally larger irregular having either n訂 row or broad segments. when they are gathering to the apical portion of It is rather difficult to include this incision in fronds. fronds. C 同phalomanes jav αnicum var. sumat- rheophytic characters adapted to rapid water ranum has large sori ，and more apparent ones current ， because the s加 ne type of incision is not in in small plants ，compared to va r. javanicum. rarely observed also for geophilous C. sing α- The sori are free and not immersed below the porianum. The young leaves are usually subentire margin of pinnae and usually with more or less and the leaves seem to be lacerated in age ，and distinct distinct stalks. In some cases ，especially when the often the laminar portion is removed remaining larger larger sori gather to the apical portion of frond ， only the veins and at most a few rows of cells along the the stalks are long and distinct. Even in C. them. It is st i1l unknown whether the irregula r1 y singaporianum ， in which the mouth of sori is below lacerate segments of C. kingii are comparable with the the level of pinna-margin ， the sori are fr 白 and the those noted above or not. short short stalks are attached at the sinus-bottom in the Distribution notches. notches. Among the six species of Cephalomanes Ecology Ecology subgen. Cephalomanes credited here ， four species
The species of subgen. Cephalomanes usually are rather rare and endemic to restricted 訂 eas: C. grow on rocks or on clayey banks along streams crassum and C. madag α'scariense are endemic to or or in stream-beds in lowlands. Therefore ， most of the Philippines (by two collections) and to them are rheophytic ，obligately or facultatively Madagascar (and Bourbon) ，respectively ，and C. according according to the species. C 句phalomanes sing α， densinervium and C. singaporianum are known porianum is often found on earth-banks near from New Guinea to the Solomon Islands and streams ，and has more or less patent pinnae. 恥1alay Peninsula to Borneo ，respectively. The other C 問phalomanes crassum is another species with two species have several infraspecific taxa and are widely widely oblique pinnae ， though the Samar collec- rather widely distributed. The two subspecies of 138 138 植物研究雑誌第66 巻第3 号 平成 3 年 6月
C. C. atrovirens are isolated in their distribution: ssp. as several Bornean species are not distributed in atrovirens atrovirens in the S. Ryukyus ，Philippines to Malaya but reach only this small island (Holttum Queensland Queensland and to the Solomon Islands and ssp. 1955). Another reophilous variety of C.javanil ωm ， boryanum in Micronesia ，and New Hebrides to var. sum αtranum ， has its range in Hainan ， Polynesia Polynesia (Fiji ， Tonga ， Samoa ， and New Vietnam ，Sumatra ，Java ，and Borneo. Caledonia). Caledonia). There is only one collection of C. No information is available as yet on the atrovirens atrovirens in the southern tip of the Ryukyus features of phytochemistry and gametophytes for (Iriomote (Iriomote Isl ょin which there are several Philippine the species of this subgenus. species species which are not found in T 剖wan. It is ， Chromosomes therefore ， not very particular to have only one Trichomanes jαvanicum was first observed collection collection of the Philippine element ， in addition to cytologically by Manton (1955) and was reported the the common occurren ∞ of other Malesian species. to have n= ca. 32 chromosomes. Braithwaite Cephalomanes javanicum has the area covering (1 969 ， 1975) counted chromosome numbers for entire entire Malesia. Vars. javanicum and asplenioides four species ，and Mitui (1 976)reported n=32 for are are allopatric except in Borneo and New Guinea; C. oblongifolium in the Ryukyus. The informa- var. var. javanicum is in Thailand ，Malaya ，Sumatra ， tion available is summarized in Table 1 according Java ，Borneo ， east to New Guinea ，and va r. to the nomenclature in this paper. This number ，
asplenioides asplenioides is in the Philippines ，Ryukyus ， n= 32 ，is constant in 0 町 subgenus and found in Taiwan ，and Pulau Tioman off Malay Peninsula Trichomanes subgen. Trichomanes ， but no other
in in addition to the 訂 eas from Borneo to New subgenera of Cephalomanes. Guinea. Guinea. In East Kalimantan ， var. javanicum is always always in lowlands ，less than 4∞m from sea-level ， Taxonomy and var. asplenioides in higher elevations ， higher Ceph α lomanes Presl in the sense of Copeland than than 500 m altitude. The of occurrence var. (1938 ， 1947) and others (e.g. ，Morton 1968 ，ぉ a asplenioides asplenioides on Pulau Tioman ， against var. section) is a natural and definable group of the
javani ω， m in Malay Peninsula ，is not very peculiar ， following six species and is placed here as a distinct
Table Table 1. Chromosome numbers counted for the species of Cψ halomanes subgen. Cephalomanes
Taxa Chromosome number Origin References
C. C. atrovirens ssp. ssp. atrovirens n=32 Solomon Is. ，New Hebrides Braithwaite (1969 ， 1975) ssp. ssp. bory αnum n=ca.64 New Hebrides ，Fiji Braithwaite (1 975) C. C. densinervium n=32 Solomon Is. Braithwaite (1969) C.jl αvα nicum var. var. Javamcum n=32 恥1alaya Manton (1955) var. var. asplenioides n=32 Solomon Is. Braithwaite (1969) n=32 Ryukyu Is. 恥1i tui (1976) June June 1991 Journal of Japanese Botany Vo l. 66 No. 3 139 subgenus subgenus of the genus Cephalomanes in broader filmy ferns. They are distributed throughout sense sense (Iwatsuki 1984). A key to the species and palaeotropics: Madagascar ， Bourbon ， India ， infraspecific infraspecific taxa is given and an enumeration of Burma ， Thailand and Indo-China ， Hainan ， all all the taxa belonging to Cephalomanes subgen. Taiwan ， S. Ryukyu ，Malay Peninsula to Malesia
Cψ halomanes is prep 紅 ed. throughout ， Pacific islands (恥1i cronesia ， Genus Cephalomanes Presl emend. K. Iwats. Melanesia ，Polynesia) ， south to Queensland. Acta Phytotax. Geobo t. 35:176. 1984. Terrestrial Terrestrial or on rocks ， of mediocre size or Key to the species and infraspecific taxa larger larger among the family; rhizome erect ， or when 1. Fronds monomorphic ，pinnate ， with more than creeping creeping short and thick; fronds fasciculate or close 10 pairs of free pinnae. together ， simply pinnate to pinnately decompound; 2. Fronds linear lanceolate ， pinnae narrowly venation venation anadromous; sori paratactic; involucres oblong to lanceolate ， the breadth less than tubular ，truncate at apex or with dilated mouth. one-third of the length ， the basiscopic base There There are seven subgenera recognized in this often dimidiate. genus genus (Iwatsuki 1984). 3. 恥10uth of involucre dilated. Subgenus Subgenus Cephalomanes; K. Iwats. Acta 4. Stipe usually up to 5 cm long ，less than
Phytotax. Phytotax. Geobo t. 35:177. 1984. -C 句phalomanes one-third of the length of fronds; pinnae Presl ，Hymen. 17. pl. 5.1843; Cope l. Phi l. J. Sci. more or less cut to lacerate " 67:66. 67:66. 1938; Genera Filicum 40. 1947. - Lacostea 1. C. atrovirens v.d.B. v.d.B. sec t. Cephalomanes (Presl) Prantl ，Unters. 5. 恥10uth of involucre slightly dilated ， the Morph. Gefaesskryp t. 1:50. 1875. - Trichomanes dilated lips up to 0.3 mm broad; plants L. L. subgen. Cephalomanes (p resl) C. Ch r. Ind. Fi l. in S. Ryukyu ，Philippines ，New Guinea xiv. xiv. 1906. - Trichomanes L. sec t. Eutrichomanes to Solomon Islands and Queensland ser. ser. Cephalomanes (Presl) v.A.v.R. Ma l. Ferns 84. ssp. atrovirens 1908. 1908. - Trichomanes L. subgen. Pachychaetum 6. Sori on normal pinnae and not confined Presl Presl sec t. Cephalomanes (Presl) Morton ， Contr. to the apical part of fronds. U.S. U.S. Na t. Herb. 38:189. 1968. Type: Trichomanes 7. Rachis terete and wingless at least in atrovirens atrovirens Pres l. the lower portion; sori not or shortly Trichomanes Trichomanes L.‘ subgen.' Schizophlebium stalked ...... f.α trovirens v.d.B. v.d.B. P l. Jungh. 1:551. 1856. Type: Trichomanes 7. Rachis narrowly winged throughout; ja νanicum Blume. sori distinctly stalked; endemic to N ew Terrestrial Terrestrial or epipetric ， often in rheophytic Guinea ...... f. kingii habitat; habitat; fronds simply pinnate ， pinnae dark green ， 6. Sori borne in a terminal spike or blackish blackish in dried specimens ，unistratose except on restricted to apex of fronds; endemic to the the veins ，subentire or variously cut at margin; New Guinea .. . .. f. acrosorum veins veins once or twice forked ， their ends usually 5. Mouth of involucre conspicuously forming forming teeth at margin of pinnae; sori paratactic ， dilated ， the dilated lips usually 0.5 mm receptacles receptacles long extruded. broad; plants in 恥1i cronesia ，恥 1elanesia ，
Six Six speci 邸 are c1 0sely related to each other and and Polynesia ...... ssp. boryanum are are more or less isolated from the other groups of 4. Stipe more than 10 cm long ，more than a 140 140 植物研究雑誌第66 巻第3 号 平成 3 年 6 月
half half of the length of frond; pinnae only F l. Phi l. 1:77. 1958; Tindale ，Cont r. N.S.W. Na t. shallowly shallowly dissected; endemic to New Herb. Fl. Ser. 201:45. 1963; Croxall ，Aust r. J. Bo t. Guinea and Solomon Islands . 23: 540. 1975. - Trichomanes atrovirens (Presl) 2. 2. C. densinervium Kunze ，Bo t. Zeit. 5: 37 1. 1847; Cope l. Phi l. J. Sci. 3. 3. Involucre truncate or the mouth hardly 51: 25 1. pl. 52. f. 3， p l. 55. f. 2. 1933. - dilated. dilated. Trichomanes javanicum var. atrovirens (Presl) 8. 8. Sori salient ， not immersed in notches ， the C. Ch r. Ind. Fi l. 168. 1905 ， 635. 1906. Type: mouth of involucre usually higher than the Philippines ，Cuming 169 (Praha ，dup l. in B GH level level of lobe-apex .... 3. C. jav αnicum K L). 9. 9. Sori not very large ，commonly 1. 7-2.5 Trichomanes boryanum Kunze ， Farnkr. 1: 237. mm long ，on normal pinnae and not con- pl. 97. 1847 ， based on Trichomanes alatum Bory ， fined fined to the apical part of fronds. Dup. Voy. Bo t. 1: 282. t. 38. f. 2. 1828 ，non Swartz 10. 10. Sori on acroscopic margin of pinnae ， 1801 ， nec Schkuhr 1809 ， nec Hooker 1821; Cope 1. not not on basiscopic margin .. Phi 1. J. Sci. 51: 254. pl. 52. f. 4. 1933. -
var. var. Jαvα nzcum Cephaloman 釘 alatum (Bory) Presl ，Abh. Boehm. 10. 10. Sori gathering to the distal portion of Ges. Wiss. V. 5: 334. 1848. - Cephalomanes pinnae and distributed towards boryanum (K unze) v.d.B. Ned. K r. Arch. 4: 351. acroscopic acroscopic margin ， or on basiscopic 1859. - L αcoste α bory αnα(Kunze) Prantl ， margm m some cases. Hymen. 50. 1875. - Trichomanes javanicum va r. . var. asplenioides α latum C. Ch r. Ind. Fi 1. 168. 1905. Type: 9. 9. Sori larger ，commonly more than 3 mm Carolines. long ， borne in a terminal spike or panicle Cephalomanes curvatum v.d.B. Ned. Kr. Arch.
var. var. sumatr α， num 4: 35 1. 1859. - Trichomanes curv αtum J. Smith ， 8. 8. Sori in notches on margin ， the mouth of J. Bo t. 3: 417. 1841 ，nom. nud. - Trichomanes
involucre involucre usually lower than the apex of jανα ni ωmv， 紅 . curvatum (v.d.B.) C. Ch r. Ind. Fil. lobes lobes ...... 4. C. singaporianum 638 ， 642. 1906. Type: Malay Arch. ，Cuming 184 2. 2. Fronds linear ， pinnae oblong to linear- (K; dup 1. B).
subdeltoid ， the breadth about two 田 fifths of the Cephalom α nes australicum v.d.B. Ned. K r. length ， auricled at acroscopic base ， the Arch. 5(2): 139. 1861 ，J. Bot. Neer 1. 1: 35 1. 186 1. basiscopic basiscopic base broadly cuneate ， not - Trichomanes javanicum var. australicum dimidiate; dimidiate; endemic to Madagascar and (v.d.B.) C. Chr. Ind. Fil. 168. 1905. Syntypes: New Bourbon ...... 5. C. madagascariense Caledonia ，Cuming 8; Fりi，Seeman 769 (K)' Milne 1. 1. Fronds subdimorphic ， pinnate only at base ， or 181 (K); New Hebrides ，An eityum Is 1.， Kajewski only only a few basal pinnae free and more than 883 (P BO SING). twenty twenty pinnae adnate to rachis; endemic to the Cephalom α nes wilkesii v.d.B. Ned. K r. Arch. Philippines Philippines ...... 6. C. crassum 5(2): 140. 1961 ， J. Bo t. Neer 1. 1: 345. 186 1. -
Trichomanes Trichomanes jα， vanicum var. wilkesii (v.d.B.) C. Enumeration Ch r. Ind. Fi l. 169. 1905. Syntypes: Fiji (Ovalao) ， 1. 1. Cephalomanes atrovirens Presl ，Hymen. 18. Wilkes; Samoa ， Wilkes 24 (GH); Samoa or l. pl. 5. 1843; Copel Phi l. J. Sci. 67:67. 1938; Fern Navigator's Is 1.， Wilkes 24 (US). June June 1991 Journal of Japanese Botany Vo l. 66 No. 3 141
Trichomanes Trichomanes kingii Cope l. Phi l. J. Sci. 6: 72. dilated; stipes usually less than one-third of the 1911 ， 51: 253. pl. 53. f. 2. 1933. - Cephalomanes length of fronds. kingii kingii (Cope l.) Cope l. Phi l. J. Sci. 67: 68. 1938. C.javani ω， m: involucres truncate or the mouth Type: Type: New Guinea ，Lakekamu ， C. King S 13 hardly d i! ated; stipes usually more than one-third (MICH). of the length of fronds. Trichomanes Trichomanes acrosorum Cope l. Phi l. J. Sci. 6: Tindale (1 963) described that the stipes of C. 72. 72. 1911 ， 51: 254. pl. 53. f. 3. 1933. - atrovirens 1-4.5 cm and of C. javanicum 8-12 Cephalomanes acrosorum (Cope l.) Cope l. Phi l. J. cm. Croxall (1 975) correctly measured that the Sci. Sci. 67: 68. 1938. - Trichomanes acrosorum var. stipes of C. javanicum were proportionately alatum alatum v.A.v.R. Bull. Bui t. 11 11: 23. 1913. Type: longer. New Guinea ，Lakekamu ， C. King 352 (MICH). 1. ssp. atrovirens Trichom α nes ledermannii Brause ，Bo t. Jahrb. Cephalomanes atrovirens Presl ， l. c. 56: 56: 35.1920. Type: NE New Guinea ，Ledermann Trichom α nes curvatum J. Sm. l. c. - Trichomanes 8638 ，9622 (B). kingii Cope l. l. c. - Trichomanes acrosorum Trichom α nes maluense Brause ，Bo t. J ahrb. 56: Cope l. l. c. - Trichomanes lederm αnnii Brause l. c. 36. 36. 1920. - Type: NE New Guinea ，Ledermann Trichomanes maluense Brause ， l. c. 6343 6343 (B). Trichomanes acranthum H. 110 ，l. c. Trichomanes Trichomanes acranthum H. 110 in Nakai ，Ic. Pl. 1) f. atrovirens As. As. O r. 2: 109. t. 45. 1937. - Cephalomanes Distr. S. Ryukyu (l riomote Is よ Philippines ， acranthum (H. It o) Tagawa ，Acta Phytotax. Celebes ，Moluccas ，New Guinea ，Solomon Islands ， Geobo t. 14: 45. 1950; Serizawa ，Sci. Rep. Takao New Hebrides ，and Queensland. Mus. Na t. His t. 7: 14 (1975). Type: Ryukyu ， As noted in the section of distribution ，one lriomote lriomote Is. ， S. Sonohara & H. Ito s.n. (TI). collection was made in R 抑止 yu ， at upper course Cephalomanes densinervium (Cope l.) Cope l. of the river Nakara ，and no additional collection sensu sensu H. It o，Bo t. Mag. Tokyo 67: 215. 1954. has been made. The tendency of gathering the sori Trichomanes Trichomanes rhomboideum J. Smith in Hook. at apex is not particular in this plant ，and is seen J. J. Bo t. 3: 417. 1841 ， n.n. polytopically in various places. There There is no problem to reduce T. maluense to 2) f. kingii (Cope l.) sta t. nov. - Trichomanes this this species ， though the former is distinguished by kingii Cope l. Phi l. J. Sci. 6C: 72. 191 1. the the following features: the whole plants are smaller Distr. N ew Guinea. and delicate ， fronds linear lanceolate ，up to 8 cm 3) f. acrosorum (Cope l.) sta t. nov. - long ， 1.5 cm wide ， the hairs on axes more or less Trichomanes α crosorum Cope l. Phi l. J. Sc i. 6C: persistent ， pinnae 10-15 pairs ， oblong to sub- 72. 191 1. orbicular ， proportionately deeply lobed at Dist r. New Guinea. acroscopic acroscopic m 訂 gin ，thinner in texture ，sori mostly Several collections were made from New on acroscopic side of pinnae ，and often on Guinea ，and the habitat is always ， if given on basiscopic basiscopic side as wel l. labels ，on stones in and along creeks in deep shade Cephalomanes atrovirens is distinguished from in the primary forests. This form with the sori C.j αvanicum ， as follows: gathering to the apical part of fronds seems to be C. C. atrovirens: mouth of involucre more or less derived in parallel in various species of this genus 142 142 植物研究雑誌第66 巻第3 号 平成 3 年 6 月
in in such habitats. from Palau and Yap. These materials in TI 紅 e Trichomanes Trichomanes acranthum was described from the different from New Guinean type in the short stipes Ryukyus ，and the distribution of sori is quite and serrate to lobed margin of pinnae. Although similar similar to this New Guinean form ， though the the dilated lips of involucre are narrower ， general general habit is so different that the Ryukyu plant commonly about 1.3 mm in breadth ，these plants is is referred to f.α trovirens ， together with some have no distinct difference from C.α trovirens ssp. West Malesian plants having the similar distribu- bory αnum. tion tion of sori on the frond. 2. 2. ssp. boryanum (Kunze) sta t. nov. - 3. Cephalomanes javanicum (B 1.) v.d.B.
Trichomanes Trichomanes bOl ァanum Kunze ，F 紅 nkr. 1: 237. p1. Hymen. Jav. 30. pl. 22. 1861 ， Ned. K r. Arch. 4:
97. 97. 凶184 仰7. 一 C ψ句ph αalom α仰n白 αω 似ωt仏LS ωS吋tかra α'(l lic αum η1 v.d.B. μ.1. 心C 350. 1861; Cope l. Phi l. J. Sci. 67: 67. 1938. 一 Ceph α10m αnes wilkesii v.d.B. 1. c. Trichomanes javanicum B 1. Enum. P 1. Jav. 224.
Cephaloman 白 densinervium (Cope l.) Cope 1. sensu 1828; Cope 1. Phi 1. J. Sci. 51: 246. pl. 52. f. 1. 1933; H. It o， l. c. Tard. & C. Chr. in Lecomte ， F 1. Gen. Indo-Chine Distr. Distr. Micronesia ，New Hebrides ，New 7(2): 68. 1939; Holt t. Rev. Fl. Malaya 2: 102. f. Caledonia ，Fiji ，Samoa ，and Pacific islands. 38. 1955. - Lacostea jανα nica (B 1.) Prantl ， Hymen. 50. 1875. Type: Java ， Blume
2. 2. Cephalomanes densinervium (Cope 1.) Cope 1. (L-908 ・325 ・638). Phi l. J. Sci. 67: 67. 1938. - Trichomanes Trichomanes asplenioides Presl ，Hymen. 129. densinervium densinervium Cope 1. Phi 1. J. Sci. 6C: 71. 1911; 51: 1843; Kunze ，F 紅 nk r. 218. p1. 89. 1847; Cope l. Phi 1. 253. 253. p1. 53. f. 1. 1933. Type: New Guinea ， C. King J. Sci. 51: 249. pl. 52. f. 2， p l. 55. f. 1. 1933 ，non
150 (MICH). Sw. 1788. - Cψ halomanes a伊 lenioides Presl ， Trichomanes Trichomanes infundibulare v.A.v.R. Nova Abh. Boehm. Ges. Wiss. V. 5: 334. 1848. - Guinea Guinea 14: 55. 1924. Type: New Guinea ，Lam 439 Trichomanes javanicum var. asplenioides (non
(BO). (BO). Presl) C. Chr. In d. Fil. 635. 1906. - Trichoman 回 Cephalomanes densinervium is similar to C. preslii Morton ， Contr. U.S. Na t. Herb. 38: 190. atrovirens atrovirens in a variety of important features but 1968. Cephalomanes javanicum va r. different different from it in more coriaceous and less in- asplenioides (C. Chr.) K. Iwats. J. Fac. Sci. Univ. cised cised pinnae ，proportionately longer stipes ，and the Tokyo 111. 13: 549. 1985. Type: Philippines ， sori sori aggregated towards the acroscopic side of Cuming 184. pinnae. pinnae. The combination of these features is Trichom α nes laciniatum Roxb. Calc. J. Na t. similar similar to that of C. singaporianum ，and it is His t. 4: 518. 1844; Morton ，Cont r. U.S. Na t. suggested suggested here that C. densinervium may have Herb. 38: 382. 1974. Lectotype: Molucca Is l. evolved evolved from C.α trovirens in parallel to C. (Amboina ?)， C. Smith s.n. in Herb. Roxburgh singaporianum singaporianum which have evolved from a mother (Brussels ， not seen). stock stock like C. javanicum. Both C. densinervium and Trichomanes oblongifolium Presl ，Epim. Bo t. C. C. singaporianum grow in less rheophytic habitat 19. pl. 10. 1849; Cope l. Phi l. J. Sci. 67: 67.1938 ， among species of Cephalomanes. 73: 467. 1940 ， Fern Fl. Phi l. 1: 78. 1958. Type: Cephalomanes densinervium is recorded from Philippines ，Cuming 169 pt. (Praha ，dup 1. in K). Micronesia Micronesia by H. It o based on collections several C 句phalomanes zollingeri v.d.B. Hymen. Jav. June June 1991 Journal of Japanese Botany Vo l. 66 No. 3 143
1. 31. pl. 23. 1861 ，Ned. K r. Ar ch. 4: 35 1. 186 1. 1. var. javanicum. - Cξ phalomanes jανα， nicum Lectotype: Lectotype: Java ，Zollinger 1464 (L) designated (B l.) v.d.B. l. c. - Cψ halomanes zollingeri v.d.B.
here. here. l. c. - Cephalom α nes rhomboideum v.d.B. 1心 Cephalomanes rhomboideum v.d.B. Hymen. Dist r. N. India ，Tha i1 and ，Vietnam ，Malaya ， Jav. Jav. 33. pl. 24.1861 ， Ned. K r. Arch. 4: 350.1861 ， Sumatra ，Borneo ，and Java. non T. rhomboideum J. Sm. n.n. Lectotype: 2. var. asplenioides (Presl) K. Iwats. J. Fac. Sci. Philippines ，Cuming 169 pt. (K) ，designated here. U. Tokyo 11 1. 13: 549. 1985. - Trichomanes Trichom αnessum α tranum v.A.v.R. B叫1. Dep t. asplenioides Presl ， Hymen. 129. 1843. - Ag r. Ind. Neer l. 18: 4. 1908; Cope l. Phi l. J. Sci. Trichomanes laciniatum Roxb. l. c. 51: 51: 248. pl. 53. f. 4. 1933; Tard. & C. Chr. in Trichomanes oblongifolium Presl ， l. c. Lecomte ， F l. Gen. Indo-Chine 7(2): 68. 1939. - Trichomanes su. 万ト utex v.A.v.R. l. c. Cψ halomanes sum α trana (v.A.v.R.) Cope l. Phi l. Distr. Ryukyu ，Taiwan ， Malaya (Pulau J. J. Sci. 67: 67. 1938. Type: Sumatra ，Burck 87 Tioman) ，Borneo ，Philippines ，Moluccas ，New (BO). (BO). Guinea ，and Solomon Is. ? Trichomanes foersteri Ros. Rep. Sp. Nov. 13: Brass 28244 (New Guinea) is particular in 213. 213. 1914; Cope l. Phi 1. J. Sci. 51: 255. 1933. Type: having long-stalked sori on the basiscopic margin Sumatra ，W. Grash o. グ 43. of pinnae on some fronds. The distribution of such Trichom α nes suffrutex v.A.v.R. Nova Guinea particular sori seems to be abnorma l. The normal
14: 14: 56. 1924. Type: New Guinea ，L αm 1197 (BO; sori are on acroscopic side ，usually in sinus betw 田 n K SING US). lobes short ，and not distinctly stalked. There is no In In every characteristic feature ， the type and the peculiarity in vegetative features ， except for some other other collections referred to T. su. グト utex are not thinner texture and dark green pagina. particular particular and this is identified as C. jlα vanicum 3. va r. sumatranum (v.A.v.R.) K. Iwats. J. Fac. var. var. asplenioides. Sc i. U. Tokyo 11 1. 13: 549. 1985. - Trichomanes 1 have seen no authentic specimens of T. sumatranum v.A.v.R. Bull. Dep t. Ag r. Ind. Neer 1. foersteri. foersteri. (Copeland did not see anY specimens.) 18: 4. 1908. There There are two distinct features described for this Dist r. Annam ，Hainan ，Sumatra ，Borneo ，and species: species: the involucres bearing crown-like Java. appendages appendages similar to those of T. superbum and Compared with two preceding varieties ， var. the the fertile pinnae restricted to the middle and lower sumatranum is distinct ， in addition to the soral parts parts of the frond. 1 have not seen any materials characters ， in such vegetative features as; the size having having these features among Cephalomanes of fronds is smaller ，commonly up to 20 cm long ， species. species. Cephalomanes species become fertile even 4.5 cm wide; pinnae are proportionately narrower ， in in very young stage ，and the soriferous pinnae are often lacerate with uninervate linear segments ， often often in middle and/or lower part of the frond in placed with acute angles ，35-45 0 ， to rachis. They these these plants. The crown-like structure of the in- are the so-called rheophytic features ，and it seems volucres volucres as described suggests that this is Nesopteris that this variety is adapted to this habitat most species ， although it should be investigated on the conspicuously in this subgenus which itself is type type materials. There is no Sumatran specimen rheophytic as a whole. The fronds are dark green examined referable to T. foersteri. in the field usually in deep shade or under gaps 144 144 植物研究雑誌第66 巻第3 号 平成 3 年 6 月 of of crowns along the streams. The fronds become Boivin 35 in Herb. de Fraqueville (P). black black after drying up ，and this is conspicuous in Distr. Madagascar; a Bourbon collection (P) is var. var. sumatranum even among the members of this also referable to this. subgenus. subgenus. Both Christensen (1 932) and Copeland (1 933 ，
1938) 1938) doubted the occurren ∞ofCψ haloman 白 sp. 4. 4. Cephalomanes singaporianum v.d. Bosch ， in Madagascar ， but Tardieu-Blot (1951) drew a Ned. K r. Arch. 4: 35 1. 1859 ， based on typical form of Cephalomanes on the basis of
Trichomanes Trichomanes javani ω， m Hook. & Grev. Ic. Fil. pl. Boivin s.n. ， probably not a type of C. 240. 240. 1830 ，non Blume 1828; Cope l. Phi l. J. Sci. madag ωcariense. 67: 67: 67. 1938. - Trichomanes singaporianum Copeland (1933) distinguished this species from (v.d.B.) (v.d.B.) v.A.v.R. Bull. Jard. Bo t. Bui t. II. 20: 25. the other members of Cψ halomanes by broadly 1915; 1915; Cope l. Phi l. J. Sci. 51: 247. pl. 52. f. 5. 1933; cordate pinnae ， though this is not in the case of t. t. Holt Rev. Fl. Malaya 2: 103. f. 39. 1955. Type: this species. The pinnae 紅 e close together and Singapore ， Wallich (K). imbricate; the acroscopic base is auricled and Trichomanes Trichomanes christu Rosens t. Bull. J ard. Bo t. covers the rachis; the basiscopic base is cuneate ， Bui t. II. 2: 27. 1911 ，non Cope l. 1906. - not dimidiate as in the other species of subgen. Trichomanes Trichomanes rosenstockuv.A.v.R. Bul 1. Jard. Bo t. Cephalomanes. In imperfect collection from B 凶t. 7: II. 27. 1912. - Type: Borneo. Bourbon in P ， the pinnae are long-stalked and Trichomanes Trichomanes borneense v.A.v.R. Bull. Jard. deeply dissected. More collections are waited to Bo t. Bui t.‘ II. 20: 25. 1915. Type: 引に Borneo ， elucidate the structure of this species. There is a Teuscher Teuscher (BO). big disjunction between Himalaya and Mada- Distr. Distr. Malaya and Borneo. gascar ，and it is interesting to elucidate the taxo- Some Sabah collections (M. Hotta 20382 ， nomic structure of the Madagascarian plants. Kokawa & Hotta 245 ，1178 ，122α Tamura &
Hotta 88 乃 are the variants somewhat similar to 6. Cephaloman 回目邸側m (Cope l.) Price ，Cont r. C. C. densinervium in general frond-construction and Univ. Mich. Herb. 17: 268. 1990. - Trichomanes need need further research. They are different from a crassum Cope l. Phi l. J. Sci. 51: 256. pl. 54 ， 55. typical typical form of C. singapori αnum in having f. 3. 1933 ， Fern F1. Phi l. 1: 78. 1958. Type: Leyte ， overlapping overlapping pinnae with linear sterile lobes in Cabalian ， G. Lopes s.n. (B ur. Sci. 40804) (Lecto- soriferous soriferous area ，and the mouth of involucre often type in MICH; dup l. in P US). extruded extruded beyond the pinna-margin. The second collection bearing fertile fronds is: Samar ，M.G. Price & B.F. Hernaez 806 (KYO L 5. 5. Cephalomanes madagascariense v.d. Bosch ， MICH). Copeland had an impression that this was Ned. K r. Arch. 4: 35 1. 1859 ，Med. Rijksh. Leiden similar superficially to Athyrium porphyrorachis (ed. (ed. Goddijn) 17: 15. f. 7. 1913; Cope l. Phi l. J. and the species of Polypodium pectinatum group ， Sci. Sci. 67: 68 ， 1938. - Trichomanes madagas- and Price and Hernaez noted on their label that cariense cariense (v.d.B.) Moore Ind. Fi l. 230. 1861; C. their collection was taken on embarkment in Ch r. Dansk Bo t. Ark. 7: 5. 1932; Cope l. Phi l. J. damp shaded ravine in limestone area and was Sci. Sci. 51: 256. 1933; Tard. in Humbert ， Fl. Madag. Haplodictyum-like. This species is at a glance Com. 3: 29. f. II I. 7-8. 195 1. Type: Madagascar ， similar to Trichomanes subgen. Trichomanes from June 1991 Journal of Japanese Botany Vo l. 66 No. 3 145 tropical tropical America ，though our species has Queensland. Austr. J. Bo t. 23: 509 ・547. anadromous branching in frond-architecture and Holttum R. E. 1955. Revised Flora of Malaya II. unistratose unistratose laminae except for the veins and Ferns of Malaya. Govern. Prin t. o ffice ， doubtlessly doubtlessly belongs to the Old World Singapore. Cψ 加 lomanes. The frond is pinnatifid nearly Iwatsuki K. 1984. Studies in the systematics of throughout ，and this characteristic ，rather peculiar filmy ferns VII. A scheme of classification among the species belonging to Cψ halomanes ，is based chiefly on the Asiatic species. Acta constant constant throughout the collections from the two Phytotax. Geobot. 35: 165-179. islands. islands. Copeland hesitated to combine this species Manton 1. 1955. Appendix: Cytological notes on with with Cephalomanes as he never saw the fertile the hundred species of 恥1alayan ferns. In R.E. frond ， but there is no doubt that this species Holttum ， Revised Flora of Malaya II. Ferns of belongs belongs to Cψ halomanes ， because of similarity in 民1alaya ， 623-628. pls. I-III. the the features including those found in the fertile Mitui K. 1976. Chromosome numbers of some fronds. fronds. The description of the fertile fronds reads: ferns in the Ryukyu Islands. J. Jpn. Bo t. 51: Fertile Fertile fronds about 8 cm long ，1 cm wide ， 33-41. pl. 1. (sterile (sterile fronds of the same plants 10-17 cm long ， 恥 10rton C. V. 1968. The genera ，subgenera ，and
up to 4 cm wide); pinnae round to subquad 幽 sections of the Hymenophyllaceae. Cont r. U .S. rangular ，more or less dissected ， each bearing 1-6 Na t. Herb. 38: 143-214. sori; sori; sori cup-shaped ， not immersed nor stalked ， Prantl K. 1875. Untersuchungen zur Morphologie about about 3 mm long ，1 mm in diameter at mouth ， der Gefaesskryptogamen 1. Hymenophyllaceen. subtruncate subtruncate or more or less blunt at mouth ， Tardieu-Blot 恥1. L. 1941. Hymenophyllacees distinctly distinctly winged; the receptacles long-extruded. nouvelles ou litigienses de Madagascar. No t. Sys t. Paris 10: 90-93. References References Tindale M. D. 1963. Hymenophyllaceae. Cont r. Braithwaite Braithwaite A. F. 1969. The cytology of some N.S.W. Na t. Herb. Fl. Se r. 201: 1-49. Hymenophyllaceae Hymenophyllaceae from the Solomon Islands. 要旨 Bri t. Fern Gaz. 10: 81-9 1.
一一一 -1975. Cytotaxonomic observation on some コケシノブ科の分類系は本シリーズの前報(植 Hymenophyllaceae Hymenophyllaceae from the New Hebrides ，F 祖i 物分類地理35 巻)所載のように 8 属にまとめられ and New Caledonia. Bo t. J. Li nn. Soc. 71: る.そのうち，ソテツホラゴケ属は広義にまとめ 167-189. 167-189. て，かつて，キクモパホラゴケ属，カンシノブホ Christensen Christensen C. 1932. The Pteridophyta of Mada- ラゴケ属など独立属とされていたものを包含する gasca r. Dansk Bo t. Ark. 7: 1-253. ことになるが，狭義のソテツホラゴケ類は亜属の Copeland E. B. 1933. Trichom αnes. Phi l. J. Sci. 階級でまとまっている.本報ではソテツホラゴケ 51: 51: 119-280. 一一一一 1938. Genera Hymenophyllacearum. Phi l. 亜属の再検討を行ない， 6 種を認めた.日本では， J. J. Sc i. 67: 1-110. 八重山諸島に 2 種知られる. 1 つはソテツホラゴ 一一一一 1947. Genera Filicum. Chron. Bo t.， ケで ，Ceph αlom αnesJ αvαnicumvar. αsplenioides Waltham ， Mass. CC. Chr.) K. Iwats. である.もう 1 つのサキシ Croxall Croxall J. P. 1975. The Hymenophyllaceae of マホラゴケは，西表島で，胞子嚢群が葉の頂部に 146 146 植物研究雑誌第66 巻第3 号 平成 3 年 6 月
集まる特徴で区別されたが，この性質はソテツホ 群が葉の頂部に集まる性質でサキシマホラゴケに ラゴケ亜属では他の種でも平行的に見られる現象 あてられた標本のうちには，包膜の唇部が広くな で，分類群を区別する指標とはなり難い.ただし， るものはなく，今のところ，サキシマホラゴケは このタイプ標本では，包膜の唇部が広くなり，ソ 日本ではタイプ標本以外には確認していない. ソ テツホラゴケとは異なっている. この性質は， c. テツホラゴケ亜属は世界で 6 種にまとめられ，ソ αtrovirens に典型的であり サキシマホラゴケは， テツホラゴケとサキシマホラゴケは共に変異があっ フィリピンから，台湾をとばして西表島に分布す て，いくつかの種内分類群が識別された. る例をもう 1 つ増すことになる.ただし，胞子嚢