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Pneumatopteris Nakai Probably the Most Sandwicensis,. Narrow Hairy

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Pneumatopteris Nakai Probably the Most Sandwicensis,. Narrow Hairy BLUMEA 21 (1973) 293 —323 Studies in the family Thelypteridaceae V. The genus Pneumatopteris Nakai R.E. Holttum Royal Botanic Gardens, Kew I re-described this in genus Blumea 19 (1971) 42. The principal characters there enu- merated are: rhizome-scales broad, thin, with marginal hairs (some bearing mucilage glands) but rarely superficial hairs; lower pinnae, usually several pairs, reduced, the transition reduced to pinnae abrupt or gradual; aerophores at bases of lower pinnae ± swollen andwhite on young fronds, sometimes much elongate; pinna-lobes with cartilag- inous which the margins are often toothed at ends of veins; veins usually anastomosing but free insome species which have deeply lobed pinnae; lamina ± pustular when dried; lower surfaces with short acicular never densely long-hairy, sometimes or capitate hairs; sessile spherical glands lacking on lamina, indusia and sporangia; body of sporangium often stalk of bearing club-shaped or capitate hairs, rarely setae; on sporangium a hair of or 4 cells, terminal cell but not brown with 3 enlarged spherical; spores light many small thin thus ± quadrate wings, spinulose in aspect. The of here included majority species in this genus conform to all the above characters, but there which differ are some species by showing one or two (rarely three) of the following characters: basal pinnae not or little reduced; scales on bases of stipes elongate and with continuous bearing many superficial hairs; spores a ± wing and anastomosing cross-wings. Almost all these aberrant species have distinctly pustular lower surface, and have the usual accompanying characters of rather thick veins and cartilaginous edges of pinna-lobes. the most is Probably aberrant species P. sandwicensis, which has narrow hairy scales, no . much-reduced basal and with continuous pinnae, spores a wing and some cross-wings. It also has thick hairs scattered the surface of the peculiar on upper lamina between veins. The only other Hawaiian of is P. which is in species Pneumatopteris hudsoniana, no way aberrant. Another aberrant Pacific is P. species costata, with two near allies in eastern Malesia. These lack pustular thickenings and have with continuous with spores one wing cross- but have the of and wings, they stipe-scales typical Pneumatopteris capitate hairs on their Their lower sporangia. pinnae are gradually reduced, giving the aspect of Christella, with which they have been confused in herbaria. P. afra, widely distributed in has Africa, a long-creeping rhizome and fronds that are A rhizome is known only slightly pustular. comparable only to occur in two other species in the assigned to this genus, Java and Sumatra. P. afra is one of few species examined and cytologically, is tetraploid. There are herbarium specimens which look like hybrids between it and Thelypteris dentata E. St. John (genus Christella of (Forsk.) my arrangement, I.e. 43) and one of them shows abortive Ghatak of the 1971, p. spores. Dr. J. University of Cape Ghana, informs that he has found such a with abortive Coast, me hybrid, spores 293 294 BLUMEA VOL. XXI, No. 2, 1973 in Ghana of the the and irregular meiosis, growing near plants two putative parents; rhizome. Ghatak that hybird can persist and spread because of its creeping Dr. also states considerable of that at meiosis a number pairs were formed, so that it is possible P. afra and dentata have Further T. (both tetraploid) some common ancestry. experiments might provide more definite evidence. The other aberrant species above mentioned have not been cytologically examined. also have and if would be In They might a hybrid nature, tetraploid self-perpetuating. and any case, cytological examination of more species, experimental hybridization, of the aberrant would the especially species, probably provide new insights into status of the genus in its relation to the other genera (or species-groups, for those persons who do not regard my groups as genera). Spores with a continuous wing and cross-wings are characteristic of Pronephrium (see Blumea but similar to those of and 20, 1972, 105 —126), spinulose spores, Pneumatopteris of of Sphaerostephanos, occur in some species Pronephrium. There are also other indications intergrading between Pronephrium and Sphaerostephanos. Such intergrading might occur between and do clear indication it. Pronephrium Pneumatopteris, but I not see any of unita and has reduced continuous P. (Madagascar Africa) no pinnae, spores with a wing and a bud on the of the rachis. has the broad scales and somewhat upper part But it pustular fronds of Pneumatopteris, and (apart from the absence of small basal pinnae), P. New P. shows a considerable resemblance to pennigera of Zealand. subpennigera appears the has of and to be closely allied to P. unita, but former spores typical Pneumatopteris no bud on the rachis. If one or both these fems are tetraploid, their inter-relations might be traced experimentally. found that of Manton a plant P. truncata in Ceylon was tetraploid (Phil. Trans. R. Soc., B, 238:138). No related species occurs there, and herbarium specimens are rather uniform, resembling closely specimens from Malaya. But in north India D. S. Loyal found that Ind. plants near Darjeeling were diploid (Proc. 48th Sci. Congr. Part III, 1961, 266—7). has his and Dr. Loyal kindly sent me a pinna from dried specimen, it matches several from Assam in Kew Herbarium. There be it be collections ought to another diploid; may found in the region from NE. India to southern China. Some indication of known below the of this variability in P. truncata is given in systematic part paper (species 45). the and Throughout Malayan region, in the Pacific, there are local species which are clearly related to P. truncata but which are distinct in various combinationsof characters: number and form of reduced pinnae, shape oflowest large pinnae, depth of lobing of frond and hairs pinnae, pubescence on on indusia, presence or absence of capitate on When is the of its sporangia. only a single specimen known, one may query significance peculiar assemblage of characters, but in many cases further specimens from the same neighbourhood have shown that the peculiar form is constant and distinctive. It wouldbe rank such local varieties of P. if this possible to some taxa as truncata, but one does one finds it impossible to delimit anenlarged P. truncata. One cannot give an adequate account of the into and I genus without taking such local taxa consideration, think the best treat- them all indeed ment is to regard as species (as many are already so recognized). Rhizome characters are always important, and unfortunately are often not recorded; related which in P. laeve they may serve to distinguish species grow the same habitat, as (creeping rhizome) and P. microloncha (erect caudex) among rocks by streams in the of it is also to have information Philippines. For adequate description species, necessary about all of frond. Individual herbarium all data, parts a specimens rarely supply necessary R. E. HOLTTUM: in the V Studies family Thelypteridaceae 295 and it is be only piecing together evidence from different specimens that one can arrive at a full This has its and lead description. process limitations, may to false conclusions. Fortunately I have seen the whole of some recent collections, and this has been a great help. I am particularly indebted to Mr. M. G. Price for excellent series of Philippine far collections which have enabled me to understand Philippine species better than would otherwise been have possible. Similarly the collections ofDr. T. G.Walker and Mr. A. C. Jermy in Celebes and New Guinea, Dr.W. A. Sledge in Samoa, Dr. A. F. Braithwaite in the Solomon Islands and New and Dr. G. Brownlie in have been Hebrides, Fiji of great also other collections of of which value; recent years I have seenrepresentatives in various herbaria. I have examined, at various times, all specimens of Old World Thelypteridaceae in the following British herbaria: Kew, Museum, Leiden, Paris, Berlin, Utrecht, Geneva, National Florence, Edinburgh, U.S. Herbarium, Nairobi, Sydney, Brisbane, Lae, Bogor, and have loan Singapore, and Calcutta, received on at Kew types of specimens from other herbaria. To the Directors of all these institutions I express my grateful thanks, especially to which has been base of Kew my operations since 1954. In Blumea 19 (1971) 42 I included Pseudocyclosorus Ching tentatively in Pneumatopteris remarks lines but (see on p. 43, 21—25), in the present account it is excluded. The dif- believe and ferences are slight, but I distinctive, there are differences in distribution. almost confined Pseudocyclosorus is to mainland Asia, with an extension into Africa; it is absent from the Malay Peninsula and is only represented in Malesia by P. tylodes (Kunze) in northern Luzon. I here in three Ching include Pneumatopteris free-veined species of New Guinea which in 1965 I placed in Pseudocyclosorus. Two of them (P. caudata and are related and P. excisa) certainly closely to P. ligulata P. keysseriana, having a thin lamina which is pustular when dried; the third (P. petrophila) is related to P. costata. of individual in the but I Descriptions species present paper are brief, have tried to indicate what are the important characters; these have often been omitted from earlier publications. I have not cited all earlier published descriptions because few are satis- cited of factory. As regards synonymy, I have place publication of all basionyms, in- those cluding regarded as synonyms. I have cited also the of all basionyms, and have them all the few types seen except in cases where the contrary is indicated, but descriptions have sometimes been amplified by addition of details from other which believe to be specimens I conspecific with the types. It is that in some I have erred in the addition possible cases my judgement by of characters shown the have not by types, but I believe I included nothing which is not true of the that types (except in some cases I have given larger dimensions of pinna-size, where it seemed obvious that the was taken from small This of type a plant).
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