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). account Pneumatopteris makes no to but from pretension finality; I hope it is a beginning which others can be

to useful lines of further guided investigation. Intensive studies in 'biosystematics' can be undertaken when of wider only a broadoutline a field has been effectively established; fields without it, useful narrower cannot be selected. An intensive study of the species of the Pacific but might be illuminating; good new collections from very many places would be needed.

KEY TO THE SPECIES OF PNEUMATOPTERIS

I. Veins anastomosing.

2. Stipe and rachis bearing short dark spines irregularly arranged. Indusia 3. lacking. with Cauda 4. Pinnae to 25x1.6—2.2 cm small 3 cm long; spores bearing many

wings I. P. glabra 296 BLUMEA VOL. XXI, No. 2, 1973

continuous and 4. Pinnae to 12x2 cm with cauda to 1J cm long; spores with wing cross-wings 2. P. sibelana

3. Indusia present. inverted (basal P. dicranogramma 5. Sori in an V ones divergent) 3.

5. Sori medial, lower ones not divergent I. P. glabra

2. Stipe and rachis lacking short dark spines.

6. Elongate aerophores present at bases ofpinnae.

7. Indusium lacking. hairs stalks acicular 8. Acicular on of sporangia; or capitate hairs near annulus; with continuous and spores wing cross-wings.

9. Sori supramedial 4. P. glandulifera

9. Sori medial.

Acicular hairs P. 10. on body of sporangia 5. parksii of obstructa 10. Capitate hairs on body sporangia 6. P.

of or cells stalk of hairs with 8. Hair 2 3 on sporangium; no near annulus; spores P. stokesii many wings 7. Indusium 7. present. minute rim in 11. Reduced pinnae consisting of a prominent aerophore with a place of lamina.

hairs with stalks of 8. callosa 12. 2- or 3-celled glandular tips on sporangia . P. stalks of 12. Acicular hairs on sporangia 9. P. mesocarpa with distinct lamina. 11. Reduced pinnae a

13. Pinnae crenate; veins c. 6 pairs 10. P. subappendiculata

lobed towards —16 II. 13. Pinnae | costa; veins 12 pairs P. superba swollen. 6. Aerophores not or slightly Indusia 14. lacking.

15. Lowest pair of pinnae not or little reduced. hairs between 16. Upper surface of lamina bearing a few thick veins; sporangia

bearing capitate hairs or setae 12. P. sandwicensis hairs 16. Upper surface lacking between veins; neither capitate hairs nor setae on sporangia.

17. Sori near costules, round, or at most basal ones a little elongate; a bud near

unita apex of rachis 13. P.

bud rachis 17. Sori ± medial, many spreading along veins; no on

14. P. subpennigera

15. Lower pinnae gradually reduced, or many much-reduced pinnae present. lower 18. 2 —5 pairs pinnae gradually reduced.

often with seta 19. Pinnae crenate; 2 pairs veins anastomosing; sporangia a near

annulus 15. P. oppositifolia

lobed or veins with or 19. Pinnae halfway more; 1 pair anastomosing; sporangia

without capitate hairs near annulus. lower 20. Sporangia lacking capitate hairs; surfaces of lamina quite glabrous

16. P. pennigera hairs lower surfaces short 20. Sporangia bearing capitate or setae; bearing very capitate and acicular hairs. with 21. Pinnae commonly 12—15 x 1.5—2.0 cm, thin; lower surface of rachis acicular hairs or without minute hairs; sporangia bearing capitate

17. P. costata R. E. HOLTTUM: Studies in theJamily Thelypteridaceae V 297

Pinnae when of 21. to 7X I cm, rigid dry; lower surface rachis densely covered

with hairs spreading i mm long; sporangia mostly bearing setae

18. P. latisquamata base of 18. Many much-reduced pinnae or tubercles, nearly to stipe.

22. Sporangia bearing capitate hairs; lower surface of costae and costules glabrous

6. P. obstructa

22. Sporangia sometimes bearing setae; lower surface of costae and costules short-

hairy 5. P. parksii

14. Indusia present. Rhizome fronds 23. long-creeping; at least 2—3 cm apart. Pinnae lobed less than 24. halfway to costa.

least of much 25. At 2 pairs basal pinnae reduced and auricled. afra 26. Pinnae crenate; sporangia copiously setiferous 19. P.

26. Pinnae lobed £; sporangia with few setae 19. X buchananii

25. One pair of basal pinnae a little reduced, narrowed towards the base

20. P. inclusa

24. Pinnae lobed more than halfway to costa 21. P. longipes Rhizome unknown. 23. erect or short-creeping; or

27. Veins 2—5 pairs; pinnae in most cases not over 12 cm long.

28. Lower pinnae gradually reduced; stipe 5—15 cm to lowest pinna. Basal auricled 29. pinnae conspicuously at acroscopic base.

30. Basal large pinnae narrowly cuneate at base on basiscopic side

22. P. egenolfioides

Basal base 30. large pinnae truncate at on basiscopic side.

Pinnae to cm to 8 mm wide above base lucida 31. 7 long, .... 23. P.

Pinnae cm wide above base. 31. to 4 long, to 11 mm hairs hairs 32. Brown on rachis, both sides; no capitate on sporangia

24. P. patentipinna

surface of rachis hairs on 32. Lower glabrous; capitate present sporangia

25. P. cheesmaniae Basal 29. pinnae not auricled on acroscopic base.

33. Basal pinnae narrowly cuneate at base on basiscopic side; pinnae to 15 cm cauda rhizome long including 4—5 cm; creeping (?) 26. P. angusticaudata Basal if 33. pinnae not narrowly cuneate at base onbasiscopic side; pinnae, 15 cm long, with shorter cauda; caudex erect. . . P. microloncha quite . 27.

28. brooksii Abrupt transition to reduced pinnae; stipe to 50 cm long . 28. P. Veins 27. 7—xo pairs or more; pinnae usually much longer. Basal little reduced. 34. pinnae not or of lobed less than 35. Pinnae well-grown plants 4 cm or more wide, halfway to

costa 20. P. inclusa

to lobed than 21. 35. Pinnae cm wide, more halfway . . . . P. longipes of basal 34. At least one pair pinnae conspicuously reduced. Middle and their of 36. upper pinnae very oblique, bases asymmetric; apices

pinnae long and very narrow.

37. Larger reduced pinnae and lower normal ones distinctly auricled at base

29. P. laevis auricled. 37. Lower pinnae narrowed towards base and not 298 BLUMEA VOL. XXI, No. 2, 1973

Pinnae lobed of 38. halfway to costa; c. 12 pairs reduced pinnae, largest 3 cm

long 30. P. obliqua

lobed less than reduced 6 38. Pinnae halfway; pinnae c. pairs, longest 5 mm

long 26. P. angusticaudata

36. Middle pinnae not very oblique, their bases ± symmetric. Reduced auricled side 39. pinnae, 1 or more pairs, conspicuously on acroscopic

at base.

Reduced basal normal ones auricled 40. pinnae 1 pair, not 31. P. basicurtata

Reduced at least 2 or basal normal also auricled. 40. pinnae 3 pairs; ones

41. Reduced pinnae 2 or 3 (or 4) pairs; lower surfaces glabrous

32. P. ecallosa of 41. Reduced pinnae 6 or more pairs; lower surface rachis at least hairy.

42. Reduced pinnae with large serrate auricle; rachis and costae closely hairs surface also short-pale-hairy beneath; on upper pale

33. P. auctipinna

Reduced with small entire 42. pinnae auricle; thick brown hairs on upper of and sometimes lower surface rachis, often on bases of costae beneath.

Indusia lobed less than basal 43. copiously short-hairy; pinnae halfway; veins P. uniting at a very obtuse angle 34. jermyi Indusia lobed basal 43. glabrous; pinnae to f; veins meeting at an acute

P. angle 35. papuana

Reduced not thus widened almost 39. pinnae auricled, in some cases equally

both sides at base.

Reduced 44. pinnae 1 or 2 pairs, minute (sometimes absent?).

Pinnae 1.6 basal both surfaces between 45. cm wide, ones narrowed to base;

veins, and indusia, glabrous 36. P. microauriculata

45. Pinnae 2.3 cm wide; basal ones not narrowed to base; both surfaces hairy

between veins; indusia hairy 37. P. christelloides

Reduced several 44. pinnae pairs, conspicuous. less than 46. Pinnae lobed distinctly halfway to costa.

Stiff of rachis and 47. curved brown hairs present on lower surface costae at

least near base of frond.

less 48. Pinnae crenate or lobed than J towards costa

10. P. subappendiculata

48. Pinnae lobed |—§ towards costa.

cm costules mm 49. Pinnae to 30 long; c. 4$ apart 38. P. remotipinna

to cm costules 6 11. P. 49. Pinnae 20 long; —7 mm apart . . superba stiff 47. No brown hairs onlower surface. reduced laticuneata 50. Pinnae io£ X ii cm; pinnae 2 or 3 pairs . 39. P.

much reduced more 50. Pinnae longer; pinnae numerous. of reduced 51. Gradual reduction of c. 4 pairs of lower pinnae +3 pairs

little narrowed to base pinnae; lower large pinnae not or

40. P. usambarensis

reduced 51. Rather abrupt transition to pinnae. little narrowed base. 52. Lowest pinnae not or to P. 53. Stipe-scales narrow, hairy 11. superba

53. Stipe-scales broad, thin, not hairy. least Reduced to at 10 lowest 54. pinnae pairs, upper cm,

2x2 cm 41. P. kerintjiensis R. E. HOLTTUM: Studies in thefamily Thelypteridaceae V 299

Reduced 8 54- pinnae c. 6 pairs, upper 2X2 cm, lowest mm long humbertii 42. P.

52. Lowest large pinnae much narrowed in basal part.

P. 55. Lower sori divergent 9. mesocarpa

sori 55. Lower not divergent.

56. No capitate hairs on body of sporangium; lower surfaces of costae and costules glabrous.

and 57. Pinna-lobes truncate toothed; largest pinnae 1.5—2.0 cm wide 43. P. michaelis

57. Pinna-lobes not truncate nor toothed; largest pinnae 2.5 cm or

more wide 44. P. novae-caledoniae

hairs on lower surfaces 56. Capitate present sporangium; ± hairy

45. P. truncata

46. Pinnae lobedat least halfway to costa.

from reduced 58. Lower sori gradually more divergent costule; pinnae many, small P. very 9. mesocarpa Lower sori little reduced small. 58. not or divergent; pinnae not very

acicular hairs P. 59. Some sporangia bearing 46. oxyoura hairs 59. Sporangia bearing capitate or none.

60. Thick curved brown hairs on lower surface of rachis and costae

P. 35. papuana

60. Thick curved hairs lacking.

61. No capitate hairs on sporangia.

62. Indusia small few very with a long hairs; stipe-scales narrow,

spreading 47. P. pergamacea

62. Indusia conspicuous, glabrous; stipe-scales broad, thin, adherent. with wide Largest fronds pinnae over cm P. sogerensis 63. 4 . 48.

63. Largest fronds with pinnae not over 3 cm wide.

64. Lower pinnae narrowed towards an abruptly dilated base Fernando ( Po) 49. P. venulosa narrowed not-dilated base. 64. Lower pinnae to a

65. Veins 6—8 pairs 50. P. prismatica

65. Veins 11 —13 pairs 51. P. macroptera hairs 61. Capitate present on sporangia near annulus.

66. Pinnae lobed f or more deeply.

Reduced all small P. tobaica 67. pinnae very ([Sumatra) .... 52. least 67. Upper reduced pinnae at i| cm long.

68. lobed to — from costa Pinnae 2 mm .... 53. P. incisa

68. Pinnae lobed from to c. 4 mm costa. towards costules 69. Basal large pinnae much narrowed base; os reduced largest pinnae not over 5 mm apart; pinnae 2—4 pairf

54. P. bryanii 69. Basal large pinnae little narrowed towards base; costules of

—8 reduced large pinnae 7 mm apart; pinnae 5 or 6 pairs

55. P. vaupelii Pinnae 66. lobed to about halfway to costa, or a little more deeply.

Lower surface short hairs small subsessile 70. bearing capitate or glands with without acicular indusia between and on veins, or hairs;

small 17. P. costata 300 BLUMEA VOL. XXI, No. 2, 1973

Lower surface rachis and 71. or costae glabrous or bearing minute acicular indusia hairs; inconspicuous var. costata surface of 71. Lower rachis and costae bearing minute acicular hairs and scattered indusia but long ones; small distinct, bearing short hairs var. hispida Lower surfaces 70. bearing short acicular hairs or none; indusium conspicuous. hairs 72. Indusia with a few in centre.

Costules commonly —6 hudsoniana 73. 5 mm apart ... 56. P. Costules 73. commonly 4—5 mm apart.

Fertile to wide 74. pinnae 3.0 cm 57. P. comorensis Fertile the 74. pinnae not over 2.5 cm wide (Malesia and Pacific).

75. Fronds dimorphic; fertile gradually decrescent at base, sterile abruptly 58. P. rotumaensis Fronds 75. not dimorphic.

76. Pinnae of largest fertile fronds more than 2 cm wide, lobed

more than to costa nitidula halfway 59. P. of fertile fronds 76. Pinnae not over 2 cm wide, lobed not more

than halfway to costa 60. P. glaberrima

72. Indusia glabrous.

77. Lower pinnae gradually reduced; slender erect hairs ± abundant

on lower surface between veins 61. P. rodigasiana

77. Transitionfrom normal to reduced pinnae abrupt or subabrupt;

no hairs between veins of lower surface.

78. Aerophores swollen; filiform scales abundant on lower surface

of costules, veins and lamina 62. P. micropaleata

minute filiform scales 78. Aerophores not swollen; very few.

Pinnae to cm lobed little than 79. 2.5 wide, a more halfway to

costa; lobes rather thin, rounded at apices 63. P. magnifica

79. Pinnae rarely over 2.0 cm wide, lobednot more than halfway;

lobes firm, a little toothed at apices . 64. P. transversaria

1. Veins free.

16x6 entire 80. Largest pinnae mm, to slightly crenate; many lower pinnae gradually reduced 65. P. nephrolepioides

80. Largest pinnae larger, lobedat least halfway to costa; lower pinnae otherwise. oflower reduced. 81. 2—4 pairs pinnae gradually

82. Pinnaelobed about halfway; indusium lacking or very small . . 17. P. costata lobed 82. Pinnae almost to costa; indusium distinct 66. P. ligulata

81. Transition to smaller basal pinnae (if present) abrupt. Reduced small. 83. pinnae 2 or 3 pairs, very

84. Basal basiscopic segment oflower pinnae very small.

85. Pinnae to c. 23 X 4 cm 67. P. keysseriana

85. Pinnae to c. 8x 1.5 cm 68. P. caudata

84. Several basiscopic segments much reduced 69. P. excisa

83. Very small reduced pinnae none or more numerous.

86. Basal pinnae (1 pair) ± reduced.

P. 87. Pinnae 2 cm x 6 mm, lobed halfway to costa 70. versteeghii lobed almost 87. Pinnae to 12 X 2.2 cm, to costa 71. P. deficiens

86. Reduced pinnae at least 6 pairs. R. E. HOLTTUM: in Studies the family Thelypteridaceae V 301

88. Pinnae to 15 cm or more long. Reduced P. 89. pinnae 6 pairs, 1—1| cm long 72. regis lower much 89. Reduced pinnae many pairs, smaller, lowest close together

73. P. imbricata

88. Pinnae not over c. 10 cm long.

Pinnae lobed to costa 90. halfway 74. P. petrophila

90. Pinnae lobed almost to costa.

to basal free 92. Pinnae 3 cm long, segment 75. P. sumbawensis Pinnae free walkeri 92. 8— 11 cm long; no segment 76. P.

I. Pneumatopteris glabra (Copel.) Holtt., comb. nov.

Fern Cyclosorus glaber Copel., Philip. J. Sc. 81 (1952) 34; Fl. Philip, (i960) 363. — Thelypteris sevillana Reed,

Phytologia 17 (1968) 313. — T ype: Ramos BS 42976, Bohol, Sevilla River (UC; BM). Fern Ramos Cyclosorus viridis Copel., Philip. J. Sc. 81 (1952) 35, pi. 24; Fl. Philip, (i960) 364. — Type: &

Edaiio BS 37490 , Luzon, Mt Masigit (US; BO).

Rhizome short first reduced basal creeping. Stipes to pinna, 50 —100 cm to large pinna,

dark, glossy, bearing irregular spines to 1 mm long; reduced pinnae many pairs, consisting of conical and small lamina. Lamina a aerophore very to 120 cm long; largest pinnae to

1.6— lobed lobes costules with 25 X 2.2 cm, caudate-acuminate, J to costa, falcate, acute; small aerophores at their bases; veins 8—10 pairs; lower surface strongly pustular, costules and veins bearing minute capitate hairs only; upper surface hairy on costa only. Sori indusia hairs medial; none or very small; capitate on sporangia. Distribution. Luzon, Bohol.

Copeland's distinction of C. viridis on the colour of dried fronds is not valid; the colour depends on method of drying. The other differences between Copeland's types are slight, in width of pinnae and depth of lobing.

sibelana 2. Pneumatopteris Holtt., spec. nov.

Caudex ignotus; stipes 5 cm longus usque pinnam reductam infimam, 45 cm usque

evolutam aculeis mm multis pinnam infimam, gracilibus 1 longis armatus. Frons 65—70 cm longa, pinnis reductis exclusis; pinnae reductae 10—12-jugatae, semicirculares, z\ —3 mm diametro, aerophoris 1 mm longis praeditae; pinnae evolutae c. 28-jugatae, infimae basin versus leviter angustatae,non auriculatae, aerophoris 1 mm longis instructae;

maximae 1.8— acuminatae pinnae 12 X 2.0 cm, basi subtruncatae, apice (cauda 10—15 mm lobis longa dentata), margine J versus costam lobatae, falcatis subacutis integris; costulae inter 5 —si mm se distantes; venae 10-jugatae, graciles, concolores, 1—1 J-jugatis anasto- mosantibus, ad membranam sinus procurrentibus; rachis subtus pilis arcuatis brunneis brevibus costa costam vestita, minute puberula; pagina superior praeter glabra.

Sori mediales, inferiores non vel paulo divergentes; indusia nulla; sporangia eglandulosa; alatae. sporae Alston Type: 16942, Batjan, Mt. Sibela, 1500 m (BM).

comb. 3. Pneumatopteris dicranogramma (v. A. v. R.) Holtt., nov.

v. v. Bot. Dryopteris dicranogramma A. R., Bull. Jard. Btzg III, 5 (1922) 202.— Type: Biinnemeijer 10454, Kerintji, Sumatra (BO; L).

callosa short Closely related to P. (no. 8, infra), differing in copious dark spines on the in stipe and lower part ofrachis, and pinnae more deeply lobed. 302 BLUMEA VOL. XXI, No. 2, 1973

Distribution. known from G. about and G. Only Kerintji at 2200 m, on Dempo, The and described in Sumatra. specimen so named in Holttum, Rev. Fl. Malaya 2 (1954)

267, is P. callosa.

comb. 4. Pneumatopteris glandulifera (Brack.) Holtt., nov.

Brack, in U. S. — Goniopteris glandulifera Wilkes, Expl. Exp. 16 (1854) 29. Dryopteris subspinosa C. Chr.,

Ind. Fil. (1905) 296; Bull. Bishop Mus. 177 (1943) 92. — Polypodium brackenridgei Hook., Spec. Fil. 5

(1864) — Samoa, Tutuila (US). 9. Type: Brackenridge,

muricatum — T Polypodium Powell ex Bak., Syn. Fil. ed. 2 (1874) 506, tioti Linn. y p e: Powell 160, Samoa (K; B).

Rhizome decumbent or to first (Powell). Stipe 100 cm more large pinna, bearing many Lamina aerophores as P. callosa. to 120 cm long, pinnae to 40 pairs; largest pinnae 28 X lobed costules with small 2.0—2.3 cm, caudate-acuminate, not quite halfway to costa; veins lowest aerophores at bases; to 15 pairs, I—1J pairs anastomosing, 2—3 pairs to

sides of long sinus-membrane; lower surface almost glabrous apart from hairs on sinus- distal lower membrane and edges. Sori medial on veins, supramedial on ones; no in-

hairs dusium; capitate on sporangia, acicular hairs on receptacle or sporangia-stalks.

Distribution: Samoa, Solomon Is. (Guadalcanal, S. Ysabel), New Hebrides.

also described with cm Christensen var. angustata (I.e. 1943:93) pinnae 1.3 wide, veins Whitmee to 12 pairs; type: 212, Samoa (K).

comb. 5. Pneumatopteris parksii (Ballard) Holtt., nov.

Kew microsora Bull. Dryopteris parksii Ballard, Bull. (1937) 347; new name for Dryopteris Copel., Bishop

Mus. — Viti 59 (1929) 12, non O. Kuntze 1891. Type: Parks 20483, Levu, Fiji (UC; MICH).

Differs from P. glandulifera and P. obstructa in setose sporangia.

Distribution. Fiji.

6. Pneumatopteris obstructa (Copel.) Holtt., comb. nov.

Dryopteris obstructa Copel., Univ. Cal. Publ. Bot. 12 (193 1) 378. — Type: Parks 22536, Rarotonga (UC;

K).

Differs from all medial. have acicular hairs P. glandulifera in sori Sporangia 3 or 4 on

their stalks, sometimes also a hair of 2 cells, the end cell swollen.

stokesii comb. 7. Pneumatopteris (E. Brown) Holtt., nov.

E. 6. — Dryopteris stokesii Brown, Bull. Bishop Mus. 89 (193 1) 20, fig. Type: Stokes 39, Rapa (BISH).

from Differs P. glandulifera:: neither seta nor capitate hair near annulus of sporangium;

hair on stalk of small sporangium 3 or 4 cells; spores bearing many wings. Distribution. Rapa (several collections).

8. Pneumatopteris callosa (Bl.) Nakai

PI. P. callosa Bot. J — callosum Enum. (Bl.) Nakai, Mag. Tokyo 47 (1933) 79 • Aspidium Bl., Jav. (1S28) 152;

108. Mett. Farngatt. 4 (1858) — Nephrodium callosum Keys., Pol. Cyath. Herb. Bung. (1873) 48; Racib.,

Fl. — callosa Ind. Fil. Handb. Btzg 1 (1898) 192. Dryopteris C. Chr., (1905) 256; v. A. v. R., (1908) 220;

Varenfl. Backer & Posth., Java (1939) 62. — Type: Blume, Java (L, no go8. 539—365).

Goniopteris lobbiana Fée, Gen. Fil. (1852) 251. — Type: Lobb 276, Java (orig.?; BM, K).

Ann. Bot. Sarasin Aspidium multijugum Chr., Jard. Btzg 15 (1898) 135. — Type: 1323, Celebes (BAS). Fl. Cyclosorus dicranogramma auct. non (v. A. v. R.) Ching: Holtt., Rev. Malaya 2 (1954) 267. R. E. HOLTTUM : Studies in thefamily Thelypteridaceae V 303

60 reduced each Caudex massive, erect. Stipe —100 cm, bearing many pinnae consisting of with covered slime when lamina an aerophore a very small rim, with young; 100 cm or

— lower narrowed towards more long; pinnae commonly 15—25 x 2 2% cm, ones gradually their bases, acuminate, lobed J—J, lobes falcate, entire; small aerophores at bases of veins sinus costules; c. 12 pairs, pairs anastomosing, 2J—3 J pairs to sides of membrane; lower surfaces ofcostae and costules sometimes sparsely hairy. Sori in inverted V; indusium firm, glabrous; sporangia with gland-tipped hair on stalk and capitate hairs near annulus.

Distribution.W. Malesia, Celebes, Flores, Ceram.

comb. 9. Pneumatopteris mesocarpa (Copel.) Holtt., nov.

Mus. — Grant Dryopteris mesocarpa Copel., Bull. Bishop 93 (1932) 9, pi. 7A. Type: 5290, Society I., Huahine (BISH; UC).

of small Closely related to P. glatidulifera (no. 4, supra), differing in presence a indusium.

10. Pneumatopteris subappendiculata (Copel.) Holtt., comb. nov.

Dryopteris subappendiculata Copel., Univ. Cal. Publ. Bot. 18 (1942) 220. — Cyclosorus subappendiculatus Fil. Brass W. Copel., Gen. (1947) 143; Philip.J. Sc. 78 (1951) 449, pi. 30. — Type: 12247, New Guinea, Idenburg R. (MICH).

Reduced lowest deltoidand pinnae 5 —7 pairs, 5 mm long, uppermost 2 cm, broadly auricled on acroscopic side; lower large pinnae ± dilated at base; largest pinnae 15—18 X

veins 6 lower 2\—3 cm, crenately lobedto depth of 2 mm (sterile) 3 mm (fertile); pairs; surface of rachis with hairs some spreading red-brown like those on upper surface; sori medial; indusium thin with a few hairs; sporangia with hair of 3 cells on stalk, no hairs near annulus.

Distribution. Eastern half ofNew Guinea, at 1400—2200 m, in forest.

11. Pneumatopteris superba (Brause) Holtt., comb. nov.

NE. New Dryopteris superba Brause, Bot. Jahrb. 56 (1920) 105. — Type: Ledermann 11733, Scpik, Guinea (B)\

Dryopteris deltiptera Copel., Univ. Cal. Publ. Bot. 18 (1942) 142. — Cyclosorus deltipterusCopel., Gen. Fil.

Brass New (1947) 142; Philip. J. Sc. 78 (1951) 449, pi. 29. — Type: 11260, W. Guinea, Lake Habbema (GH).

Caudex erect; scales elongate, hairy. Stipe and rachis densely red-hairy, lower surface of rachis less reduced lowest densely; pinnae 7 or 8 pairs, spreading, broadly hastate,

wide above dilated acu- 3—5 mm; largest pinnae to 17 cm long, 3 cm base, very firm, minate, edges lobed J towards costa; aerophores distinctly swollen; costules 6—7 mm basal lobes rachis and beneath apart; veins 10 or 11 pairs (in to 16 pairs); costae ± hairy

(hairs stiff, red, \ mm), costules and surface not; sori medial; indusia rather thin, glabrous: sporangia lacking hairs near annulus. half Distribution. Eastern of New Guinea, at 2000—2500 m.

12. Pneumatopteris sandwicensis (Brack.) Holtt., comb. nov.

Stegnogramma sandwicensis Brack, in Wilkes, U. S. Expl. Exp. 16 (1854) 26, t. 4, f. 2. — Polypodium sand-

Am. — wicense Hook., Spec. Fil. 5 (1864) 5, non Hook. & 1832. Polypodium stegnogrammoides Bak., Syn.

Pl. nov. — Eaton in Enum. Haw. 218 Fil. (1867) 317. Polypodium microdendron Mann, (1868) (nom. super-

282. — fi-) - Thelypteris hawaiiensis Reed, Phytologia 17 (1968) Type: Brackenridge, Sandwich I. (US; K). BLUMEA 304 VOL. XXI, No. 2, 1973

& Bot. Sw. 1801. — T Polypodiumpolycarpon Hook. Am., Beechey Voy. (1832) 104, non ype: Beechey, Oahu (K).

Caudex erect. to at least scales Stipe 50 cm, broad, firm, short-hairy. Lamina c. 100 cm, basal middle lobed pinnae deflexed, hardly reduced; pinnae commonly 18x3 cm, hardly costules edges strongly cartilaginous; 5 mm apart, veins 10 pairs; lower surface pustular, costules and veins; surface thick hairs between short-hairy on upper bearing sparse long veins. Sori acicular elongate along veins, exindusiate; sporangia bearing many capitate or hairs; spores with a broad thin wing and a few cross-wings. Distribution.Hawaii.

unita comb. 13. Pneumatopteris (Kunze) Holtt., nov.

Gymnogramma unita Kunze, Linnaea 18 (1844) 115. — Type: Gueinzius, Natal (not seen).

Fil. — Natal Goniopteris patens Fee, Gen. (1852) 253. Type: Gueinzius, (not seen).

Fil. Afr. — unita Goniopteris silvaticus Pappe & Rawson, Syn. austr. (1858) 39. Type: as Gymnogramma

Kunze.

Goniopteris madagascariensis Fee, Gen. Fil. (1852) 251. — Aspidium malagassium Kuhn 1868. — Type: Goudot, Madagascar (not seen).

Nephrodiumcostulare Bak., J. Linn. Soc. Bot. 16 (1877) 203. — Type: Gilpin, Madagascar (K).

Dryopteris gladiataC. Chr., Ark. for Bot. II, 14, no 19 (1916) 4, t. 1; Dansk Bot. Ark. 7 (1932) 52, pi. 12,

12. — T Palm & fig. y p e: Afzelius 333, Madagascar (BM).

Caudex with white erect. Young coiled fronds covered slime, linear aerophores and swollen. Lamina c. 100 cm; basal pinnae slightly reduced, deflexed, auricled on acroscopic

costules —6 side, narrowed basiscopically; largest pinnae to 25 X 3 cm, lobed J—J, 5 mm which apart, veins 8—12 pairs; broad thin scales at first present on lower surface of costae also bears short hairs; a gemma present near apex of rachis. Sori near costules, lower

hairs with continuous ones spreading a little, exindusiate; no on sporangia; spores a wing and cross-wings.

D istribution. Eastern Africa, Madagascar. Kunze's description is adequate for identification of this species.

comb. 14. Pneumatopteris subpennigera (C. Chr.) Holtt., nov.

Dryopteris subpennigera G. Chr., Dansk Bot. Ark. 7 (1932) $2, pi. 12, f. 1,2. — Type: Perrier de la Bathie 15620 Madagascar (P; BM). ,

Differs all from P. unita: no gemma on rachis; sori medial, elongate along veins; spores with small many wings.

comb. 15. Pneumatopteris oppositifolia (Hook.) Holtt., nov.

Polypodium oppositifolium Hook., Spec. Fil. 5 (1863) 8. — Cyclosorus oppositifolius (Hook.) Tard., Mem-

28 Ferns W. Tr. G. S- I.F.A.N. (1953) 128, tiomen tantum; Alston, Afr. (1959) 62. — Type: Mann, Thomd (K).

Rhizome lamina basal of short-creeping; stipe 75 cm; 75 cm, 2 or 3 pairs pinnae wide gradually reduced, lowest cm long, 2\ cm at cordate base; largest pinnae 22 x 2.3 auricled of cm, at base, edges crenate to depth i| mm; costules 3^—4 mm apart, veins short stiff hairs. 8 pairs; lower surface finely pustular, costae and costules with Sori in- sometimes framedial; no indusia; sporangia with a seta near annulus; spores with a ± continuous wing and cross-wings.

Distribution. S. Thome, Fernando Po, Annobon. R. E. HOLTTUM: Studies in the family Thelypteridaceae V 305

16. Pneumatopteris pennigera (Forst.) Holtt., comb. nov.

Kr. — Polypodium pennigerum Forst., Prodr. (1786) 82; Schkuhr, Gew. 1 (1804) 17, t. 22. Goniopterisforsteri

T. Ind. Fil. (1858) 99. — Type: Forster New Zealand. Moore, ,

lamina with Caudex erect; stipe-base bearing flat not-hairy scales; c. 4 pairs of lower reduced lobed costules pinnae gradually and auricled; largest pinnae c. 15x2 cm, halfway;

lower surface of rachis and costae ovate scales 5 —7 mm apart; veins to 10 pairs; bearing when sori young, also sparse brown acicular hairs and very short capitate hairs; inframe- hairs with small dial, exindusiate; sporangia lacking near annulus; spores many wings. Distribution: New Zealand, Australia (Victoria and Tasmania).

I have seen no Forster specimen, but Schkuhr based his description and figure on one, his and evidence is adequate. Swartz (Syn. Fil. p. 250) stated that a minute reniform-

is veins he peltate indusium present, and that the are pubescent beneath; must surely the under have seen another species, perhaps P. nymphale Forst. The scales on surface of

costae of fronds are like those of but the different. young P. unita, spores are

17. Pneumatopteris costata (Brack.) Holtt., comb. nov.

Goniopteris costata Brack, in Wilkes, U. S. Expl. Exp. 16 (1854) 28. — Type: Brackenridge, Fiji (US).

rurutensis Occ. Mus. — T St & Dryopteris Copel., Pap. Bishop 14 (1938) 55, pi. 7. y p e: John Fosberg 16590, Rurutu, Austral I. (BISH; K).

Dryopteris christophersenii C. Chr., Bull. Bishop Mus. 177 (1943) 87, pi. 3A. — Type: Christophersen 79, Samoa (BISH).

Lastrea cavitensis Copel., Philip. J. Sc. 81 (1952) 26, Fern Fl. Philip, (i960) 328. — Type: Mangubat BS

1302, Cavite, Luzon (MICH).

C. Ind. Fil. Dryopterispennigera sensu Chr., (1905) 284, p.p. in Dryopteris parasitica auct. non (L.) O. Ktze: C. Chr. Skottsb., Nat. Flist. Juan Fernandez & Easter I. (1920)

50.

Dryopteris truncata auct. non (Poir.) C. Chr.: C. Chr. in Rensch, Hedwigia 74 (1934) 232, quoad Rensch 761

tantum.

Caudex base of covered with broad flat oflower erect; stipe scales; 3 or 4 pairs pinnae

gradually reduced, not auricled, lowest c. 1 cm long; largest pinnae to 18 X 2 cm (much

smaller be lobed little veins ones may fertile), a over halfway to costa; to 8—9 pairs, basal pair anastomosing; all lower surfaces bearing minute acicular and capitate hairs old surface between veins sometimes (the latter often collapsed on specimens); upper

with short acicular and sori exindusiate with very capitate hairs; small, medial, or a very hairs annulus hair of small indusium; sporangia with capitate near and a 3 cells on stalk;

with continuous and spores a wing cross-wings. Distribution. Malesia (Bah, Sumbawa, Luzon, NE. New Guinea), N. Queens-

land, New Hebrides and New Caledonia to Easter I. and Young plants may have free veins, one such, already fertile, was the type ofJLastrea

cavitensis; M. G. Price has confirmed this. Specimens from Cook Is., Pitcairn and Easter I.

have acicular hairs lower surface of rachis and and small some longer on costa, a hairy

indusium, but otherwise do not differ; they deserve varietal rank.

18. Pneumatopteris latisquamata Holtt., spec. nov.

P. costata (Brack.) Holtt. affinis, differt: fronde multo minore, coriacea; pinnis maximis

0.8 costulis inter venis indusiis c. 3.5 X cm; 2 mm se distantibus; 4-jugatis; nullis; sporangiis setiferis.

Type: Millar & Holttum NGF 15777, NE. New Guinea, Morobe Distr., Edie Creek,

2000 m (K; LAE). 306 BLUMEA VOL. XXI, No. 2, 1973

afra 19. Pneumatopteris (Chr.) Holtt., comb. nov.

Soc. Bot. France Mem. — Dryopteris afra Chr., Bull. 55 (1908) 8b: 107. Cyclosorus afra Ching 1941; Alston,

Ferns W. Trop. Afr. (1959) 63; Tard. in Aubr. Fl. Cameroun 3 (1964) 250, pi. 37. — Lectotype:

Chevalier 5799, Oubangui (P).

Notes Pterid. — Le Testu Gabon Dryopteris dewevrei Chr. ex Bonap., 14 (1924) 207. Type: 1719, (P).

Nephrodiumabruptum (Bl.) Hook., Spec. Fil. 4 (1862) 77, p.p.

Rhizome long-creeping; basal 2 pairs pinnae much reduced, strongly auricled; largest of pinnae to 20X 2.5 cm, crenate to depth 2 mm; veins 10—12 pairs, 4—5 pairs anastomo-

surface of costae and costules sori medial, indusia hairy; sing; lower short-hairy; spo- with of cells their with rangia usually 2 or 3 setae, a hair 2 or 3 on stalks; spores many small wings.

Distributio n.West Africa (south to Angola); Uganda.

dentata S. Bot. T Probable buchananii Afr. id. — hybrid: Thelypteris var. Schelpe,J. 31 (1965) 265, fig. y p e:

Schelpe 5599, Mozambique(BOL).

towards of Rhizome long-creeping; pinnae lobed 1/3—2/5 costa; 2 pairs veins anasto- hairs all mosing; minute over lower surface; few sporangia setose; spores (where observed) abortive.

Distribution. Liberia, Ghana, Cameroons, Tanzania, Uganda, Kenya, Mozam- bique. Probably a hybrid between P. afra and Christella dentata (Forsk.) Holtt., comb. nov.

dentatum Fl. The have (Polypodium Forsk.,) Aegypt-arab., 1775, 185). presumed hybrids a than ferns have wider distribution is now known for P. afra; but Thelypteroid not yet been critically collected in Africa.

inclusa comb. 20. Pneumatopteris (Copel.) Holtt., nov.

Publ. Bartlett Dryopteris inclusa Copel., Univ. Cal. Bot. 14 (1929) 373, pi. 57. — T ype: 8376, Karo Plateau, Sumatra (UC).

Ark. Dryopteris berastagiensis C. Chr., Dansk Bot. 9, no 3 (1937) 59. — T ype: Ridley s.n., 1921, Berastagi,

Sumatra (K).

Rhizome diam. Frond with 8 creeping, I cm stipe 120—250 cm; pinnae —12 (—20) lower lowest little lobed pairs, ones 12 cm apart, a reduced, largest 25 x 4—3 cm, edges J towards costules lower surface costa; 6|—mm apart, veins 10—12 pairs; glabrous; sori medial, indusia large, thin, with a few hairs.

Distribution. Central Sumatra, several collections.

21. Pneumatopteris longipes (Bl.) Holtt., comb. nov.

Enum. — — Aspidium longipes Bl., PI. Jav. (1828) 155. Type: Blume , Boerangrang, Java (L, no 908. 333 1062).

lamina basal Rhizome long-creeping, I cm diam.; stipe c. too cm, c. 100 cm; I or 2 often pinnae ± reduced (variable, unequal); largest pinnae on type 17x2.2 cm, largest lobed than lobes seen 25x2.7 cm, caudate-acuminate, more halfway to costa, slightly costules falcate, rounded and slightly toothed; 4 mm apart, veins 10—11 pairs; costae and costules sparsely hairy beneath; sori medial; indusium small, hairy; sporangia with capitate hairs.

Distribution. Java, at 1400—1800 m; Sumatra (?).

22. Pneumatopteris egenolfioides Holtt., spec. nov.

Rhizoma fasciculatos procumbens, apice stipites gerens; stipites 10—17 cm longi, R. E. HOLTTUM: Studies in the family Thelypteridaceae V 307

vestiti. basin versus paleis tenuibus adpressis i| mm longis Lamina usque 45 cm longa; pinnae c. 28—jugatae, inferiores 3 —4-jugatae sensim reductae et remotiores, infimae 5 —7 laminae firmiter vel mm longae; textura herbacea rigida, in sicco pallide olivacea; pinnae basi maximae 2.4—3.2 cm longae, 6—8 mm latae, basiscopice anguste cuneatae, acrosco- pice late cuneatae et valde auriculatae, apice subtruncatae mucroneque minuto praeditae, costulae margine c. £ versus costam lobatae; lobi obliqui truncati, leviter dentati; usque inter 3 mm se distantes, angulo 45° vel paulo ultra e costa orientes; venae plerumque vel inferior 2-jugatae, infimae anastomosantes (prope apicem pinnae) liberae; pagina pilis unicellularibus destituta, costae costulaeque paleis reductis capillaceis praeditae; rachis

Sori basesque costarum supra hirsutae, pagina superior cetera glabra. prope costam utrinque uniseriati; indusia glabra; sporangia intcrdum pilo capitato ornata.

Brass W. New Guinea, 18 km NE. of Lake Habbema, 2200 on Type: 11497, m, limestone cliffs in forest (L; BO, BM).

lucida 23. Pneumatopteris (Bak.) Holtt., comb. nov.

Nephrodium lucidum Bak., Gard. Chron. N. S. 8 (1877) 456. — Type: cult. Hort. Bot. Reg. Kew, orig. 'Madagascar

Rhizome short-creeping; frond to 35 cm long; lower 1—3 pairs of pinnae gradually

0.8 base lobed less than reduced; largest pinnae 7X cm, auricled, apex acuminate, edges |; veins 3 —4 pairs except in basal auricle, 1 pair anastomosing; lower surface quite glabrous; sori with glabrous indusia; sporangia lacking hairs near annulus. This matches known from but similar no species Madagascar, is closely to P. laevis from think that the (no. 29, infra) Philippines. I cultivatedplant was probably mis-labelled; other Philippine plants were in cultivation at Kew at that time.

24. Pneumatopteris patentipinna Holtt., spec. nov.

Lamina Caudex brevis erectus; stipes 12—15 cm longus, glaber. 30—35 cm longa, inferiores pinnis 14—16-jugatis papyraceis constituta; pinnae 5-jugatae sensim reductae, fere basi infima 6 auricula oppositae, omnes acroscopice auriculatae, mm longa 5 mm alta praedita, sequentes 9 mm, 11 mm, 20 mm, 2.4 cm longae; pinnae maximae 4.0 x 1.1 breviter cm, alternae, recte patentes, basi truncatae et auriculatae, apice acuminatae, antice leviter margine usque J costam versus lobatae, lobis truncatis, acutis, dentatis, costulae ciliatis; 3 mm inter se distantes; venae usque 4-jugatae, concolores, unijugis anastomosantibus; rachis subtus pilis brunneis mm longis pilisque pallidis costae 5 vestita, brevipilosae, costulae pilis sparsis tantum praeditae; rachis supra pilis brunneis, costae pilis pallidis vestitae. Sori mediales; indusia tenuia, breviter pilosa; sporangia eglandulosa.

Type: T.G. Walker 8477, NE. New Guinea (BM).

from Eastern otherwise have of lower Specimens Highlands, similar, up to 10 pairs pinnae gradually reduced; they are almost equally dilated on both sides at their bases.

25. Pneumatopteris cheesmaniae Holtt., spec. nov.

Caudex ignotus; stipes 26 cm longus, glaber. Frons 26 cm longa, pinnis c. 14-paribus, inferioribus 3-jugatis sensim reductis constituta; pinnae infimae 3 —5 mm longae; pinnae basi breviter maximae 3.3 X 1.1 cm, sessiles, truncatae acroscopice auriculatae, apice den- acutae, margine §—| versus costam lobatae, lobis quadratis subtruncatis leviter

costulae inter in lobo tatis; 3 —3| mm se distantes, recte patentes; venae 3-jugatae (nisi inferiorum ubi infimae acroscopico pinnarum 4 —5-jugatae), anastomosantes, vena 308 BLUMEA VOL. XXI, No. 2, 1973

communi excurrenti brevi ad sinum terminata; pagina inferior omnino (membranis indusia sinuum inclusis) glabra. Sori inframediales; glabra; sporangia saepe glandulis pyriformibus ornatis.

T Cheesman forest y p e: L. E. 1269, Japan I., W. New Guinea, 300 m, on log in mossy on old coral limestone (BM).

26. Pneumatopteris angusticaudata Holtt., spec. nov.

Habitu habitudioni laevi differt: inferioribus mm et P. congruens, pinnis 6-jugatis 3—5 ultra longis; pinnis maximis basi non auriculatis, margine non 2 mm versus costam crenatis, apice longe caudatis, cauda 4—5 cm longa subintegra non dentata. rock-face Type: Brass 24984, Goodenough I., Papua, on a beside a stream (BM; LAE).

microloncha 27. Pneumatopteris (Chr.) Holtt., comb. nov.

Sc. Bot. Gen. Fil. Dryopteris microloncha Chr., Philip. J. 2 (1907) 202. — Cyclosorus microlonchus Copel.,

(1947) 143; Fern Fl. Philip, (i960) 366. — Lectotype: Mangubat 1304, Luzon (P).

Caudex lamina erect; stipe 4 —10 cm, glabrous; 20—50 cm long, pinnae 15—25 pairs, lowest lowest mm 3—6 pairs gradually reduced, 3 —5 long; largest pinnae 3—12 cm base little dilated fronds long, 0.6—1.3 cm wide, a both sides, apex on larger caudate, lobed less than lowest lower edges halfway to costa; veins 3 —4 pairs, anastomosing; surface throughout bearing short fine spreading hairs; upper surface hairy on costa only; sori medial, indusia short-hairy; sporangia with yellow capitate hairs. low Distribution. Luzon, altitudes, on rocky stream-banks.

Christ cited several specimens. As lectotype I select the best one now in the Paris herbarium. Christ also cited but this to have been a mixed collection, Cuming 31 7, appears and it is not represented in Christ's herbarium at Paris. It is possible that P. microloncha represents a stunted form of P. nitidula, due to habitat conditions.

28. Pneumatopteris brooksii (Copel.) Holtt., comb. nov.

brooksii — Bidi, Dryopteris Copel., Philip. J. Sc. 3 Bot. (1908) 345. Type: C.J. Brooks s.n., April 1908, Sarawak (MICH; BM).

Caudex erect to lowest reduced lamina or suberect; stipe 53 cm pinna; 72 cm long; normal pinnae c. 40 pairs, lowest 3 pairs abruptly reduced, 4—5 mm long; lowest pinnae narrowed towards their base bases; largest pinnae X 1.3 cm, abruptly broad-cuneate, lobed rather less lobes costules apex gradually attenuate, edges £ or deeply, oblique; veins with short sinus- 3 J—4 mm apart, 4 pairs, lowest anastomosing excurrent vein to membrane; both surfaces of pinnae glabrous; sori medial; indusia small, glabrous (not seen by Copeland but certainly present); sporangia lacking hairs near annulus. Distribution. known collections. Sarawak, on limestone; 3

laevis comb. 29. Pneumatopteris (Mett.) Holtt., nov.

Aspidium laeve Mett.,Farngatt. (1858) — Neotype: Samar (B, ex Herb. Mett.). 4 104. Jagort

Bot. — Lecto- Dryopteris luzonica Chr., Philip. J. Sc. 2 (1907) 196, incl. var. polyotis, excl. var. puberula.

pe: Loher s.n. 1906, Mt Makiling ty t Jan. (P).

Rhizome quite prostrate, rather slender, with closely seriate fronds; stipe 12 cm to first reduced pinna, 30 cm to lowest large pinna; lamina to 90 cm long including reduced lowest of lowest pinnae; 4 pairs pinnae sub-abruptly reduced, 7 mm long; largest pinnae R. E. HOLTTUM: Studies in the family Thelypteridaceae V 309

— middle and to 18 x 1.4 2.0cm, upper ones oblique, base asymmetric, auricled onacroscopic

side, narrower and rounded on basiscopic, apex caudate-acuminate, edges lobed halfway

costules veins lower sori to costa; to 4 mm apart; 5—8 pairs; surfaces quite glabrous; hairs medial; indusia glabrous or sometimes with a few hairs; sporangia usually lacking

near annulus.

Distribution. banks of Philippines; on rocky streams, fronds pendulous. of bis but such Mettenius cited as type A. laeve: 'Cuming 83 ex parte', no specimen

exists at Berlin; it was probably destroyed in the herbarium at Leipzig. In the Berlin

Herbarium are two sheets of Jagor's collection which were named A. laeve by Mettenius

and with his agree description; I select these as neotype.

Fern Fl. has A. laeve of Copeland, Philip, (i960) 364 as a synonym Cyclosorus nitidulus, luzonica Chr. and and appears to regard Dryopteris as partly C. nitidulus partly C. micro-

is Mr. G. Price lonchus; his account very confused. I am indebted to M. for ample speci- and also has mens living plants. A living plant sent to Kew some capitate hairs on the

sporangia; perhaps there has been crossing with P. microloncha, but in general the two

are and I have species quite distinct, seen many specimens. luzonica Chr. which Dryopteris var. puberula is Nephrodium philippinense Bak., belongs in to the genus Sphaerostephanos my arrangement.

30. Pneumatopteris obliqua Holtt., spec. nov.

Ann. Bot. Aspidium externum auct. non Bl.: Chr., Jard. Btzg 15 (1898) 133.

Rhizoma vel ultra reductam breve, repens; stipes 40 cm usque pinnam infimam, 90 infimam cm usque pinnam evolutam, glaber. Lamina 65 cm longa, pinnis reductis exclusis; reductae pinnae c. 12-jugatae, superiores usque 3 mm longae, ovatae; pinnae evolutae inferiores basin 6-jugatae versus multo angustatae (basi 3—4 mm latae), non

16 auriculatae; pinnae mediales et superiores valde obliquae; pinnae maximae c. X 1.9 cm, basi asymmetricae, apice caudatae (cauda 3—4 cm longa subintegra), | costam versus lobatae, lobis apice falcatis, margine leviter dentatis; costulae 5 mm inter se distantes, sub subtus angulo 50° abeuntes; venae 7—8-jugatae, pallidae vel rufescentes, infimis latere anastomosantibus; costae utroque sparsim et breviter pilosae, pinnis cetera glabris. Sori mediales; indusia tenuia, glabra vel pilis brevibus paucis praedita; sporangia interdum pilis capitatis ornata. Celebes Other collections from Type: P & F. Sarasin 127, Masarang, (BAS). N. Celebes: Posthumus Alston 2434; 15693, 15729, 16370.

basicurtata 31. Pneumatopteris Holtt., spec. nov.

basin stramineus. Caudex ignotus; stipes 42 cm longus, praeter fuscam Frons 50 cm valde longa, pinnis c. 15-jugatis constituta, pinnis infimis unijugatis reductis, 2 cm longis, infimae 12 mm latis, acroscopice auriculatis; pinnae evolutae leviter remotae, deflexae, basin maximae basiscopice versus valde acroscopice parum angustatae; pinnae 12 cm longae, 2.4 cm latae, basi truncatae non dilatatae, apice breviter acuminatae, margine

paullo ultra dimidio lobatae, lobis falcatis integris subacutis; costulae 5 — mm inter se infimae distantes; venae 9—10-jugatae, pallidae, prominentes, anastomosantes; pagina Sori inferior praeter pilos paucos submarginales glabra. mediales, inferiores leviter

interdum divergentes; indusia magna, pilis brevibus paucis praedita; sporangia pilis brevibus capitatis ornata. Korinchi Type: H. C. Robinson & C. B. Kloss 148, Peak, Sumatra, 2290 m (BM). Also from same locality, Biinnemeijer 10296, pinnae to 19x2.7 cm.

309 310 BLUMEA VOL. XXI, No. 2, 1973

ecallosa comb. 32. Pneumatopteris (Holtt.) Holtt., nov.

Bull. Fl. Cyclosorus ecallosus Holtt., Gard. Sing. II (1947) 269; Rev. Mai. 2 (1954) 272, fig. 156. — Type:

Holttum 31294, Cameron Highlands, Malaya (SING; BO, K). Also cited Holttum 23427.

from P. truncata in Differs (no. 45, infra) short-creeping rhizome, 2—4 pairs of basal pinnae abruptly reduced and subequal, strongly auricled, lowest large pinnae slightly bases narrowedto their with a large lobed acroscopic auricle; lower surfaces quite glabrous.

Distribution. Only known from Cameron Highlands; now abundant near

at m where forest has been cleared. streams 1500

Holtt., 33. Pneumatopteris auctipinna spec. nov.

Ann. Bot. Aspidium truncatum sensu Chr., Jard. Btzg 15 (1898) 133, p.p.

Caudex brevis, erectus. Stipes c. 20 cm longus, pallidus, glabrescens. Frons c. 150 cm inferiores longa, basin versus subabrupte angustata; pinnae 35-jugatae, oppositae; pinnae

infimae 15 mm longae, sequentes 8 pares sensim longiores earum superiores 2\—4 cm

longae, omnes basi acroscopica auriculatae; pinnae intermediae 2 —3 pares et pinnae

maximae inferiores omnes basi auriculatae; pinnae suprabasales c. 18x2 cm, basi trun-

catae (non auriculatae), apice anguste acuminatae, margine —§ versus costam lobatae, inter leviter costulis 4—4! mm se distantibus, lobis subtruncatis, margine dentatis; venae infimis 8—9-jugatae utrinque prominentes, anastomosantibus, paribus 2—sequentibus

membranam pellucidem elongatam tegentibus (venis in auriculis pinnarum inferiorum in saepe furcatis, ut Haplodictyum anastomosantibus); rhachis costaeque subtus dense

breviter pilosae, pagina laminae ± pustulosa, pilis brevibus paucis praedita; costa supra

cetera Sori indusia breviter antrorse hirsuta, pagina superiore glabra. inframediales; parva,

pilosa; aporangia pilis brevibus capitatis prope annulum omata. P. & F. Sarasin N. Celebes, Tomohon(BAS). Ty p e: 119, Distribution. Celebes, E. Java, Flores, Ambon, Buru.

This species is strikingly distinct in the form of the base of the frond. The species most nearly resembling it in its auricled lower pinnae is P. ecallosa from Malaya, but in the latter of basal reduced. species only 2 or 3 pairs pinnae are

34. Pneumatopteris jermyi Holtt., spec. nov.

reductam Caudex brevis, erectus; stipes 23 cm usque pinnam infimam, 50 cm usque pinnam evolutam infimam, breviter hirsutus, basi paleis latis tenuibus vestitus. Frons reductis reductae infimae 60 cm longa, pinnis exceptis; pinnae 6-jugatae, alternae, ro- tundatae triangulares, basi 2 —5 mm longae, superiores 15—20x8 mm, truncatae; basin leviter basi pinnae evolutae c. 22-jugatae, infimae versus angustatae, acroscopice basi caudato-acumi- paulo auriculatae; pinnae maximae c. 13 X 1.8 cm, truncatae, apice

— cm costam lobis vix natae (cauda 3 longa), 2/5 —1/2 versus lobatae, subtruncatis, fere dentatis, breve ciliatis; costulae 4 mm inter se distantes, rectae; venae usque 9-jugatae, ad membranam 1—ij-jugatae anastomosantes, sequentibus unijugatis sinus longam rachis arcuatis procurrentibus; subtus pilis brunneis rigidis f mm longis vestita; costae subtus pilis brunneis vel pallidis vestitae, costulae minute hirsutae; rachis basesque

brunneis costulas non indusia costarum supra pilis vestitae. Sori inframediales, tegentes; copiose brevi-hirsuta; sporangia eglandulosa. forest Type: Jermy 3739, NE. New Guinea, Butemu, in secondary (BM).

Morobe has to Brass 29766, Distr., Kaindi, pinnae 14x2.5 cm. R. E. HOLTTUM: Studies in thefamily Thelypteridaceae V 311

35. Pneumatopteris papuana Holtt., spec. nov.

fere P. jermyo aflinis, difFert: frondibus majoribus; pinnis maximis usque 18 x 3.5 cm, f costulis inferioribus versus costam lobatis, 5 —6 mm inter se distantibus, venis prope sinum sub angulo acuto conniventibus vel latera membranae sinus tegentibus, rachi

cetera indusia subtus basin versus tantum brunneo-hirsuta, pagina inferiori glabra, glabra.

2000 (LAE; L). Also Brass Type: Brass 22778, Mt Dayman, m, Papua BM, 12273,

12221, 31641.

36. Pneumatopteris microauriculata Holtt., spec. nov.

Rhizoma breviter cm longus, glabrescens, repens, 5 mm diametro; stipes 50—70

18— paleis tenuibus latis basin versus vestitus. Frons 35 —40 cm longa, pinnis evolutis jugatis pinnisque reductis minutis remotis unijugatis constituta; pinnae infimae leviter basi reductae, e medio basin versus angustatae, 8 mm latae, non auriculatae; pinnae maximae basi truncata leviter acuminatam 12 x 1.6 cm, auriculatae, apicem versus sursum lobis costulae arcuatae, f- costam versus lobatae, falcatis obtusis integris; 3^—4 mm infimae inferior inter se distantes; venae 8-jugatae, concolores, anastomosantes; pagina

costarum et membranas sinuum Sori praeter pilos minutos prope apicem glabra. paulo inframediales; indusia magna, tenuia, glabra, caduca; sporangia eglandulosa. Also Clemens Type: Clemens 27137, Mt Kinabalu, Sabah, 850 m (BM). 30463.

christelloides 37. Pneumatopteris Holtt., spec. nov.

Caudex ignotus; stipes 40 cm longus, pinnas minutas remotas 2-jugatas ferens, stramineus, paleis latis tenuibus adpressis vestitus. Fronsc. 60 cm longa, pinnis 20-jugatis;

infimae basi maximae pinnae paulo reductae, nec angustatae nec auriculatae; pinnae breviter lobi 14.5x2.3 cm, basi truncatae, apice acuminatae, f costam versus lobatae; obtuso inter subtruncati, apiculo antrorso praediti; costulae 4 mm se distantes; venae

excurrenti 11—12-jugatae, concolores, 2-jugatis anastomosantibus, vena composita inferior flexuosa, 1^—2-jugatis ad membranam sinus procurrentibus; pagina costae cos- tularum laminaeque omnino pilis pallidis patentibus gracilibus vestita; pagina superior leviter inferioribus inter venas pilis multis minutis suberectis praedita. Sori supramediales, infimis leviter divergentibus, interdum contiguis; indusia parva, pilis multis brevibus vestita; sporangia eglandulosa.

Type: Clemens 27341, Sabah, Mt Kinabalu, Dallas, 1000 m (K). The specific epithet christelloides is given owing to the resemblance of this species to the fromall of Christella in its broad flat scales and in its genus Christella Lev. It differs species

which have small spores, many wings.

comb. 38. Pneumatopteris remotipinna (Bonap.) Holtt., nov.

Ark. C. Dansk Bot. (1932) f. 10. Dryopteris remotipinna Bonap., Notes Pterid. 5 (1917) 57; Chr., 7 50, pi. 12,

— Type: Perrier 6072, Madagascar (P). in — sambiranensis C. Chr. Dryopteris longifolia Bonap., Notes Pterid. 5 (1917) 55, non Hieron. 1907. D.

Dansk Bot. Ark. — Perrier, Cat. PI. Madag. (1932) 25, nom. nov.\ 7 (1932) 50, pi. 12, f. 11. Type:

Perrier 7665, Madagascar (P).

Caudex lowest reduced base covered with thin flat erect; stipe 25 —45 cm to pinna,

8 of reduced and scales; frond to 250 cm long; at least pairs lower pinnae widely spaced, lowest and not 5—6 mm long wide, 5thpair2.5 X 1.3 cm, triangular, crenate, auricled; largest BLUMEA VOL. No. 312 XXI, 2, 1973

—28x2.0 lower narrowed their pinnae 25 —2.5 cm, ones not to bases, apex narrowly lobes costules attenuate, lobed to c. §, oblong with rounded tips; 4J mm apart; veins sides of 10—11 pairs, 1—2 pairs anastomosing, 1 pair to sinus-membrane; some stiff red- hairs and brown on lower surface of costae, also short pale hairs on costae, veins, some-

times on surface between veins. Sori near costules, lower ones somewhat divergent;

indusia glabrous; sporangia bearing large yellow capitate hairs; red glandular cell at end of stalk. hair on sporangium Distribution. Madagascar.

Christensen distal of the lower of which apparently only saw a pinna type, pinnae are much his The larger than figure 10. type has slender hairs all over lower surface, but

much less other Madagascan specimens are hairy. The yellow glandular hairs on sporan-

red end-cells of hairs on their stalks gia, and are distinctive, and there are always some short red-brown hairs.

laticuneata 39. Pneumatopteris Holtt., spec. nov.

Caudex reductam basi latis brevis, erectus; stipes 22 cm usque pinnam infimam; paleis

60 tenuibus vestitus. Frons c. cm longa; pinnae reductae 2-jugatae, superiores 15—17x6 normales mm, basi cuneatae, inferiores 5 —6 mm longae; pinnae 18-jugatae, inferiores basi latae basin versus angustatae 1 cm non auriculatae; pinnae maximae 10.5 X 1.6 cm, late basi late patentes, cuneatae, apicem versus semsim angustatae, £ costam versus lobatae, inter lobis obliquis, truncatis, integris; costulae 3 J mm se distantes; venae 8-jugatae, ad membranam i£-jugatae anastomosantes, unijugatae sinus procurrentes; pagina inferior rachis omnino glabra; costaeque supra sparsim hirsutae. Sori inframediales; indusia glabra, sporangia eglandulosa.

Type: Jermy 8044, NE. New Guinea, Bewani Mts, 250 m (BM).

usambarensis 40. Pneumatopteris Holtt., spec. nov.

Rhizoma breve repens; stipes 30 cm longus, basi paleis parvis adpressis vestitus; frons inferiores 160 cm longa; pinnae 3-jugatae 5 —8 mm longae, pinnae sequentes 4J,

basi cm steriles 10, 15 cm longae, omnes truncatae; pinnae maximae 30 longae, usque

fertiles basi leviter costam 2.6 cm latae, 2.3 cm, dilatatae, apice acuminatae, —| rotundatis versus lobatae, lobis fertilibus subtruncatis leviter dentatis, sterilibus integris; late costulae 5 —5J mm inter se distantes, patentes; venae 8—10-jugatae, 2 —jugatae

anastomosantes; pagina inferior omnino glabra. Sori paulo inframediales; indusia tenuia

glabra vel pilis paucis praedita; sporangia pilis capitatis omata.

Faden et al. E. Usambara frond Type: 70/293, Tanzania, Mts, 1000 m (EA, one on Also Usambara 7 sheets). on Mts Greenway 4794, A. Braun 1427, A. Peter 159, 9999, 16859, C. Hoist 4263, 2266.

41. Pneumatopteris kerintjiensis Holtt., spec. nov.

Caudex ignotus; stipes 5 cm usque pinnam reductam infimam, 75 cm vel ultra usque infimam. pinnam evolutam Frons 120 cm longa pinnis reductis exclusis, textura rigide

reductae 10 inferiores 2x2 x papyracea; pinnae —12-jugatae, cm, supremae 3! 3J cm, evolutae infimaebasin basi triangulares, margine crenatae; pinnae versus non angustatae, basi dilatatae; aerophorae dilatatae, non elongatae; pinnae maximae 25 x 2| cm, truncatae,

apice acuminatae non caudatae, J—| costam versus lobatae, lobis rotundatis integris;

costulae inter 4J mm se distantes; venae usque 12-jugatae, graciles, 2-jugatae anastomo- R. E. HOLTTUM: Studies in thefamily Thelypteridaceae V 313

ad membranam inferior valde santcs, i—if-jugatae sinus procurrcntes; pagina pustulosa, minutis costa costulae venaeque subtus pilis praeditae; pagina superior praeter costam

hirsutam glabra. Sori paulo inframediales, inferiores non divergentes; indusia tenuia,

glabra; sporangia pilis capitatis ornata.

Type: Alston 14149, Sumatra, Sungei Kering, Kerintji (BM). Other collections from central Alston Sumatra are: 13096, Biinnemeijer 3575, 4172, 5031. The large number of large deltoid reduced pinnae and widened base of lower normal like those of lack pinnae are distinctive. Sterile pinnae look very much P. callosa, but aerophores.

humbertii 42. Pneumatopteris Holtt., spec. nov.

Cydosorus subpennigerus (C. Chr.) Tard, in Humbert, Fl. Madag. Polypod. i (1958) 298, p.p.

Caudex ignotus; stipes 20 cm longus usque pinnam reductam infimam; frons usque

100 cm longa. Pinnae inferiores 6-jugatae reductae, infimae 8 mm longae, superiores 2 cm vel ultra evolutae longae, basi truncata 2.2 cm latae; pinnae inferiores 20 cm longae,

basin vel basi non 2.5—3.0 cm latae, versus non paulo angustatae, acroscopice auriculatae, acuminatae lobis leviter apice non caudatae, | costam versus lobatae, falcatis, integris, interdum apiculatis; costulae 5 mm inter se distantes, venae 8—10-jugatae, crassae,

sinuatam pallidae, 2% —3-jugatae anastomosantes, venam excurrentem valde facientes; costa subtus pilis brevibus rigidis praesertim distaliter vestitae, costulae venae laminaque

brevibus Sori pilis sparsim praeditae; costa supra hirsuta, costulae minute setiferae. inferiores leviter indusia mediales, interdum divergentes et elongatae; magna, glabra; sporangia eglandulosa.

Type: Humbert 31679, Madagascar, partie occidentale du Massif de Marojejy.

1400 m (P). Also Humbert 22463 (K).

43. Pneumatopteris michaelis Holtt., spec. nov.

Caudex reductam infimam erectus; stipes usque pinnam 17 cm longus, usque pinnam infimam evolutam 50 cm, glaber, viridis. Lamina usque 60 cm longa, pinnis reductis

reductae 12x8 exclusis; pinnae 4 —5-jugatae, superiores mm, triangulares, basi truncatae non auriculatae; aerophorae in vivo dilatatae; pinnae intermediae 1—2-jugatae interdum basin basi praesentes; pinnae inferiores versus angustatae (parte angusta fere integra),

8 latae maximae —1.8 mm non auriculatae; pinnae usque 22 cm longae, 1.5 cm (fertiles) basi cm (steriles) latae, subtruncatae, apicem versus sensim angustatae, J costam versus costulae sub lobatae, lobis obliquis truncatis, dentatis; 4 mm inter se distantes, angulo

6o° vel ultra egredientes; venae 7—8-jugatae, 1 —i|-jugatae anastomosantes; pagina inferior, marginibus loborum inclusis, glabra. Sori inframediales; indusia tenuia, glabra; sporangia pilis non praedita.

M.G. Price Mt Also from Type: 317, Luzon, Makiling, 1050 m (K). same locality

Price Sabah: Kloss near Sandakan 337, 2418, 2177. From SFN 19113, Bettotan, (K).

novae-caledoniae 44. Pneumatopteris Holtt., spec. nov.

Fl. Nouv.-Cal. Cyclosorus truncatus sensu Brownlie, no 3 (1969) 219, p.p.

P. truncata et P. sogerensi similis; a P. truncata differt lobis pinnarum nec truncatis nec subtus dentatis, costis fere glabris, sporangiis eglandulosis; a P. sogerensi differt pinnis numquam usque dimidium lobatis, pinnis reductis basi angustatis. BLUMEA VOL. No. 314 XXI, 2, 1973

Lecard Type: s.n. Oct. 1876, New Caledonia, 'prenant un tres-grand developpement le bord des sur sources d'eau', o—400 m (P, a single frond on 5 sheets).

45. Pneumatopteris truncata (Poir.) Holtt., comb. nov.

truncatum Polypodium Poir., Encycl. Meth. 5 (1804) 534. — Type: Brezil, no collector (P).

Aspidium abortivum Bl., Enum. PI. Jav. (1828) 154. — Type: Blume, 'ad pedem Boerangrang', Java (L,

no 90S. 337—855)-

Aspidiumabruptum Bl., PI. — Bot. Enum. Jav. (1828) 154. Nephrodium abruptum J. Sm., Hook. J. 3 (1841) 411;

Fil. — T & Hook., Spec. 4(1862) 77, p.p., nott. 241B. yp e: Kuhl van Hasselt, Java(L, no 908. 337—817).

eusorum Thw., Enum. PI. — T C. P. Aspidium Zeyl. (1864) 391. ype: Thwaites, 3069 (K).

C. ex Notul. — Cyclosorus lepidopodus Chr. Tard. & C. Chr., Syst. 7 (1938) 73. Type: Eberhardt 5252, Tonkin, Bac Kan (BM).

non truncatum auct. Presl: Fil. — Nephrodium (Gaud.) Bak., Syn. (1867) 294,p.p.; Racib., Fl. Btzg 1 (1898) 190. O. Ktze: A. Handb. Backer Dryopteris truncata auct. non (Gaud.) v. v. R., (1908) 227, p.p.; & Posth.,

Varenfl. Java (1939) 54.

Caudex lowest reduced 8 erect; stipe to pinna 5 —10 cm; reduced pinnae to c. pairs, lower normal in largest c. 2 cm long, obovate; pinnae narrowed basal part, narrow part

to base rather shallowly lobed; largest pinnae c. 25 X 3 cm, broadly cuneate, apex abruptly little less than acuminate, lobeda halfway to costa; lobes subtruncate and slightly toothed; costules 4—s mm apart, veins to 10 pairs, 1^—2 pairs anastomosing, next pair to sides of surface of sinus-membrane; lower rachis, costae and costules usually ± hairy. Sori lower with inframedial, ones not divergent; indusia glabrous; sporangia small colourless

capitate hairs near annulus.

Distribution. Ceylon & S. India, NE. India to S. China;Western Malesia and

Philippines.

The the lower of It resembles type represents large pinnae a fertile frond. specimens

from Ceylon and Malaya and cannot have come from Brazil. As mentioned in the intro-

that ductory part of the present paper, Manton found a Ceylon plant was tetraploid, and

subsequently Loyal found a diploid near Darjeeling. The latter is sufficiently distinct to justify recognition as a variety as follows.

Pneumatopteris truncata var. loyalii Holtt., var. nov.

lobis valde A typo speciei differt; pinnis angustioribus; pinnarum dentatis; sporangiis glandulis luteis magnis fere globosis ornatis.

T D. S. 600 y p e: Loyal s.n. Aug. 1956, Manjitar-Teesta Road, Darjeeling, m (Herb. Several other Panjab Univ. no. 3172). specimens, from various places in Assam, in Kew

Herbarium, agree with Loyal's specimen in the details described. Biol. Ching has described var. angustipinna from Hainan (Bull. Fan Mem. Inst. 8, 1938, has small colourless hairs much of 218); it glandular on sporangia, as in Ceylon specimens

P. truncata, but much narrower pinnae. It might be another diploid.

In Sabah, on Mt Kinabalu, is a possibly distinct variety with pinnae to 4 cm wide but no other obviously distinctive character. The of abortivum Bl. and Bl. differ from each types Aspidium A. abruptum slightly other, but other specimens from Java appear to be intermediate. There may be a distinct mountain ofthe in but field is to show its distinc- variety species Java, new study necessary tive characters.

A plant from Sabah, cultivated at Kew, has densely short-hairy indusia, also short erect hairs on lower surface oflamina; it couldbe ofhybrid origin.

Therehas been confusionin the literature betweenPolypodium truncatum Poir. andPolysti- chum the latter here of truncatum Gaud.; is placed as a synonym Pneumatopteris glaberrima. R. E. HOLTTUM: in Studies the family Thelypteridaceae V 315

comb. 46. Pneumatopteris oxyoura (Copel.) Holtt., nov.

Sc. Dryopteris oxyoura Copel., Philip. J. 60 (1936) 107, pi. 12. — Type: Brass 2696, Solomon Is., San Cristoval (BRI).

Caudex erect; stipe 50 cm or more long; reduced pinnae very small, I or 2 pairs; frond several of towards their 100 cm or more long; pairs lower pinnae narrowed bases; lobed largest pinnae 25 x 3 cm, caudate-acuminate, a Httle more than J; lobes subentire; costules to mm sides 5 apart, veins 10—12 pairs, I—1| pairs anastomosing, 1—1| pairs to of sinus-membrane; short stiff spreading hairs on lower surface of costae. Sori medial; indusium small with a few short hairs; sporangia setose.

Distribution. San Cristoval, two collections.

47. Pneumatopteris pergamacea Holtt., spec. nov.

Caudex suberectus; stipes 10 cm usque pinnam reductam infimam, 45 cm usque pin- evolutam omnino nam infimam, pilis brevibus palbdis basi paleis patentes 10X 1 mm setiferis vestitus. Frons 50 cm longa, pinnis reductis exclusis; pinnae reductae 11—15- infimae basi late jugatae, 8x6 mm, hastatae, superiores usque 2.4x1.5 cm, cuneatae

evolutae infimae margine lobatae; pinnae c. 20-jugatae, leviter deflexae, medio 3.4 cm

basin late cuneatam sensim fertiles maximae latae, versus angustatae; pinnae 23 X 2.8 cm interdum leviter (steriles latiores), acuminatae, | costam versus lobatae, lobis falcatis

costulae inter sursum paulo angustatis, subintegris; usque si mm se distantes; venae infimae 11—12-jugatae, concolores, tantum anastomosantes, sequentibus 1—i^-jugatis ad membranam sinus procurrentibus; rachis costae costulaeque subtus minute et sparsim hirsutae; pinnae reductae supra inter venas pilis pallidis adpressis vestitae, pinnae evolutae Sori supra glabrae. mediales, inferiores leviter divergentes; indusia minuta, pilis paucis praedita; sporangia eglandulosa. cult. Type: Jermy 7877, origin New Ireland, Hort. Bot. Kew. (123/70, no IOJJ).

found in Danfu limestone Original plant river valley, 630 m, onmossy boulder in forest.

48. Pneumatopteris sogerensis (Gepp) Holtt., comb. nov.

Dryopteris sogerensis Gepp, J. Bot. 61 (1923) Suppl. 61 — Lectotype: Forbes 446, Papua, Sogeri (BM).

Caudex erect; stipe to 30 cm to basal reduced pinna; reduced pinnae c. 6 pairs, largest with 2—4 cm long, ± triangular broad base; basal large pinnae narrowed towards their

— than bases; largest pinnae commonly 27 X 2\ 3 cm (to 36 X 4J cm), lobed more halfway to costa smaller fronds to about costules mm veins (on £); commonly 4—5 apart; 10—14 pairs, 1 pair anastomosing, next I or 2 veins to sides of sinus-membrane; lower surface of sometimes with hairs costa usually glabrous, sparse spreading to | mm long. Sori lowest indusia inframedial, not divergent; glabrous; sporangia lacking hairs near annulus.

Distribution. Moluccas, New Guinea, Bismarck Arch., Solomon Is. in low country, frequent; Queensland.

venulosa 49. Pneumatopteris (O. Kuntze) Holtt., comb. nov.

Desv. Nephrodium venulosum Hook., Spec. Fil. 4 (1862) 71, non — Aspidium elatum Mett. ex Kuhn, Fil. Afr.

— is O. Rev. Gen. — (1866) 131, non Bojer. Dryop ter venulosa Ktze, PI. 2 (1891) 814, nom. nov. Dryopterts

C. Soc. Brot. elata Chr., Ind. Fil. (1905) 263. — Cyclosorus elatus Alston, Bol. 2, 30 (1956) 13; Ferns W.

Trop. Afr. (1959) 63. — Thelypteris elata Schelpe, J. S. Afr. Bot. 31 (1965) 265. — Thelypteris venulosa Mann Po Reed, Phytologia 17 (1968) 323. — T y p e: G. s.n., Fernando (K). BLUMEA 316 VOL. XXI, No. 2, 1973

to at least to basal reduced reduced Stipe 40 cm pinna; pinnae c. 4 pairs, lowest 1.5 cm

to auricled lower long, highest 5 cm, on acroscopic base, largest pinnae 20 x 2\—3 cm, narrowed in basal lobed little ones 4 —5 cm, base 2 cm wide, acuminate (not caudate), a

than lobes toothed costules more §, oblique, oblong, ± at vein-ends; 5 mm apart; veins lowest 9 pairs, pair anastomosing, one pair to sinus-membrane; rachis and costae ± hairy beneath. Sori indusia inframedial; thin, glabrous; sporangia lacking hairs near annulus.

Distribution. Fernando Po & S. Thome.

Christensen for this regarded A. elatum Mett. as correct basionym species, believing that A. elatum but Bojer (1837) was a nomett nudum; Bojer printed a brief description, so

that A. elatum Mett. is a later homonym.

50. Pneumatopteris prismatica (Desv.) Holtt., comb. nov.

Mem. Linn. Soc. Paris — collector Nephrodium prismaticum Desv., 6 (1827) 256. Type: Mauritius, no named (P).

Linnaea caudiculatum Sieber Kunze, 280. — caudiculatum Sieber Aspidium ex 24 (1851) Nephrodium ex Presl,

Bot. — T Mauritius Epim. (1851) 46. ype: Sieber 47, (dupl. K).

Rhizome short-creeping; stipe c. 20 cm to lowest reduced pinna; reduced pinnae 3 —4 pairs, ± orbicular, 5 —10 mm long and wide; 1—2 pairs pinnae of intermediate length sometimes lower much narrowed their present; large pinnae to bases; largest pinnae c. base of 17—23 x2.2—2.5 cm, middleones sometimes dilatedonbasiscopic side, acuminate, lobed little than lobes to a more halfway to costa, slightly oblique, apices rounded to subtruncate costules 6—8 and ± dentate; 4—5 mm apart; veins pairs, 1 pair anastomosing, next pair to sides of sinus-membrane; lower surfaces glabrous. Sori medial; indusia gla- brous; sporangia lacking capitate hairs. Distribution. Mauritius, Bourbon.

A further possible is Kaulf. ed. synonym Aspidium mascarenense ex. Spr., Syst. Veg. 16, 'Ins. Mascaren.' 4 (1827) 101. No type was indicated, only

51. Pneumatopteris macroptera (Copel.) Holtt., comb. nov.

Univ. Cal. Bot. Parks Dryopteris macroptera Copel., Publ. 12 (1931) 392. — T ype: 16177> Tonga (UC; US).

reduced and Stipe 70 cm to lowest normal pinna; lowest 4 pairs pinnae gradually more widely spaced, not auricled; lowest normal pinnae narrowed towards their bases; largest lobed than costules pinnae 25 —28x1.8—2.4 cm, more halfway to costa; 4 mm apart; veins n—13 pairs, lowest pair anastomosing, next pair to sinus-membrane; lower surface and of rachis costae slightly hairy. Sori medial, lower ones not divergent; indusia small, hairs annulus. glabrous; sporangia lacking near

Distribution. Tonga.

52. Pneumatopteris tobaica Holtt., spec. nov.

Caudex ignotus; frons typi imperfecta (basi stipitis et dimidio apicali laminae deficienti-

reductae c. X non bus); pinnae plurijugatae, 7 7 mm, auriculatae, inter se 5 —6 cm distantes; evolutae sensim multo maximae pinnae inferiores plures basin versus angustatae; pinnae

24 x 2.8 cm, acuminatae, fere f costam versus lobatae, lobis oblongis subtruncatis, dentatis; costulae infimae 5 —5J mm inter se distantes; venae 11—12-jugatae, anastomosantes, ad ceterae omnes ad marginem procurrentes; pagina inferior praeter pilos paucos rachin, membranas inferiores sinuum et marginem glabra. Sori inframediales, non divergentes; R. E. HOLTTUM: Studies in the family Thelypteridaceae V 317

indusia firma, fusca, pilis brevibus praedita; sporangia pilis capitatis prope annulum ornata.

Surbeck side of Lake m Type: 14, NE. Sumatra, S. Toba, 1900 (L; BO).

The pinnae of this specimen are much like those of P. longipes, but the reduced pinnae are distinctive; the rhizome might show another difference.

53. Pneumatopteris incisa Holtt., spec. nov.

Caudex ignotus; stipes atroviridis (Alston), glabrescens, 15 cm longus usque pinnam reductam infimam, 50 cm usque pinnas evolutas; pinnae reductae 7-jugatae, infimae evolutae 5 —6 mm longae et latae, superiores IJX 10 mm, triangulares, crenatae; pinnae inferiores basin versus sensim angustatae, non auriculatae; pinnae maximae 21 X2.7 cm, i—costam lobis leviter acuminatae, versus lobatae, falcatis, integris, apice rotundatis; costulae 5 mm inter se distantes; venae usque 12-jugatae, graciles, infimae tantum anasto- mosantes;rachis subtus sparsim hirsuta, costae costulae venaelaminaque subtus pilis gracili- erectis vestitae. Sori costulas indusia bus pallidis sparsim inframediales, non tegentes; tenuia, pilis gracilibus praedita; sporangia pilis capitatis ornata.

Type: Alston 16623, Ternate, G. Gamalama (BM). Also Pleyte 397, Halmahera,

G. 600 in forest Sembilan, m, by stream (L).

54. Pneumatopteris bryanii (C. Chr.) Holtt., comb. nov.

Mus. — T Dryopteris bryanii C. Chr., Bishop Bull. 177 (1943) 89, pi. 4A. y p e:Bryan 160, Samoa (BISH; K).

Caudex basal of erect; stipe 30—40 cm, minutely hairy; 2—4 pairs pinnae abruptly much much reduced and widely spaced, lowest 1 cm long; lower large pinnae narrowed towards their bases; largest pinnae 25 x 2.8 cm, acuminate, lobed f or more deeply, lobes

and costules to slightly falcate, tips broadly pointed slightly toothed; 5 mm apart; veins to 14 pairs, 1—pairs anastomosing; lower surfaces glabrous. Sori near costules; indusia annulus. small, glabrous; sporangia with capitate hairs near

55. Pneumatopteris vaupelii (C. Chr.) Holtt., comb. nov.

C. Mus. Samoa Dryopteris vaupelii Chr., Bishop Bull. 177 (1943) 89, pi. 3B. —-Type: Vaupel 228, (B).

Caudex frond all Basal of not known; in 250 cm long. 4 or 5 pairs pinnae 1—2 cm long, then orbicular, c. 3 pairs transitional to normal pinnae; largest pinnae 30X 3.5 cm, lobed veins c. f, lobes somewhat narrowed to subtruncate apex; costules 7 mm or more apart;

1 to sides of lower surface 10—12 pairs, 1 pair anastomosing, or 2 veins sinus-membrane; glabrous. Sori inframedial; indusia glabrous; sporangia with capitate hairs.

hudsoniana 56. Pneumatopteris (Brack.) Holtt., comb. nov.

Nephrodium hudsonianum Brack, in Wilkes, U. S. Expl. Exp. 16(1854) '88, t. 25. — Typ e: Brackenridge s.n.,

Sandwich I. (US; K).

first reduced first 6 Stipe c. 17 cm to pinna, 45 cm to large pinna; reduced pinnae c. lowest under pairs, 1 cm long, upper ones not auricled; basal large pinnae not narrowed to lobed little half- bases; largest pinnae 20—2J cm long, 2\—3 cm wide, acuminate, a over

—6 veins way to costa; lobes oblong with minutely dentate apex; costules 5 mm apart; to

next 1 2 veins to sides of sinus-membrane; lower 9 pairs, basal pair anastomosing, or surface from hairs. lowest not indusia glabrous apart marginal Sori medial, divergent; with with a few hairs; sporangia capitate yellow hairs near annulus. Distribution. Hawaii. 318 BLUMEA VOL. XXI, No. 2, 1973

57. Pneumatopteris comorensis Holtt., spec. nov.

Caudex verisimiliter erectus; stipes 30 cm longus usque pinnam reductam infimam, reductae infimae paleis adpressis vestitus; pinnae 4-jugatae, 1 X 1 cm, superiores usque infimae evolutae basin 2x1.5 cm, crenatae,non auriculatae; pinnae versus non angustatae; maximae pinnae 33x3.0 cm, basi truncatae, caudato-acuminatae (cauda 2\ cm longa), dimidio vel paulo ultra versus costam lobatae; lobi non falcati, integri, apice rotundati, ciliati; costulae 4%—j(—6) mm inter se distantes; venae usque 12-jugatae, I—- anastomosantibus; rachis subtus sparsim hirsutae, costae copiose costulae sparsim pilis patentibus vestitae. Sori mediales; indusia tenuia, longe-pilosa; sporangia pilis capitatis elongatis omata.

Comoro Type: Hildebrandt 1782, I., Johanna, 200—800 m (K; BM, B, L, FI).

58. Pneumatopteris rotumaensis (St John) Holtt., comb. nov.

rotumaensis Mus. I. St Occ. — T St Rotuma Cyclosorus John, Pap. Bishop 21 (1954) 180. y p e: John 19159, (BISH; K).

Caudex tall. Fronds sterile with erect, to 20 cm dimorphous, subabrupt transition to lower basal reduced pinnae, large pinnae narrowed to their bases, largest to 24 X 2.7 cm, little lobed a more than fertile frond with basal pinnae, c. 9 pairs, gradually reduced, lowest narrowed their costules —6 large pinnae not to bases, largest 20x 2.2 cm; 5 mm

8 —10 rachis beneath apart, veins pairs (sterile) 5 —7 pairs (fertile); minutely hairy; sori slightly inframedial; indusia large, thin, with a few hairs; sporangia bearing capitate hairs.

Distribution. Rotuma I., New Hebrides. Perhaps only a fully developed form of P. glaberrima (no. 60, infra).

nitidula comb. 59. Pneumatopteris (Presl) Holtt., nov.

Nephrodium nitidulum Presl, Epim. Bot. (1851) 46, Holtt., Novit. Bot. Inst. Bot. Univ. Carol. Prag. 1968

— Fern omn. T (1969) 40. Cyclosorus nitidulus Copel., Fl. Philip, (i960) 364, sytt. excl. — y pe: Cuming

s.n., Philippines (PRC).

Caudex reduced lowest erect; pinnae 5 —7 pairs, c. 6x6 mm, orbicular, uppermost

cm cm more wide at truncate basal not 3 long, 2 or base, edges crenate; large pinnae or little narrowed their of sterile sometimes to bases; largest pinnae type 24 x 2.2 cm, ones slender lobed little lobes with rounded to 3 cm wide, evenly attenuate to tip, a over £,

costules — lowest tips; 4% mm apart; veins 9—10 pairs, concolorous, slender, pair anastomosing, next pair to sinus-membrane; lower surfaces with minute erect hairs.

Sori slightly inframedial; indusia hairy; sporangia with capitate hairs.

Distribution. Philippines, Luzon to Mindanao.

60. Pneumatopteris glaberrima (Richard) Holtt., comb. nov.

Sert. Astrol. xviii. — T Durville 1827, W. New Aspidium glaberrimum Rich., (1834) y pe: s.n., Guinea, Port Dorei (P).

Polystichum truncatum Gaud, in Freyc. Voy. Bot. (1827) 332. — Aspidium truncatum Gaud., ibid. pi. 10. —

T Notul. — Gaudichaud Rawak Cyclosorus truncatus Tard., Syst. 7 (1938) 78, nomen tantum. ype: , (Lawak) I., W. New Guinea (P; B).

Caudex reduced —6 lowest probably erect; pinnae 4 pairs, c. 1 x 1 cm, uppermost c.

2x1 cm, base truncate not auricled; largest pinnae probably c. 16 x 1.6 cm, lobed halfway

veins —8 with minute or rather less deeply; 7 pairs; lower surface glabrous (or some erect few short hairs. hairs). Sori medial; indusia with a hairs; sporangia bearing capitate R. E. HOLTTUM Studies in : thefamily Thelypteridaceae V 319

Distribution. New Guinea, Bismarck Arch.

Better specimens are needed for clear characterization of this species. It is possible that the Gaudichaud's species is distinct; if so, it cannot bear name truncata in the genus Pneuma- topteris.

comb. 6l. Pneumatopteris rodigasiana (T. Moore) Holtt., nov.

T. Moore in Hort. T Nephrodium rodigasianum Linden, 111. 29 (1882) 27, pi. 442. — ype: cult., origin Samoa (K).

Caudex basal 8 of lowest apparently erect; pairs pinnae very gradually reduced; large pinnae not narrowed to their bases; largest pinnae (type) 15x2 cm, acuminate, lobed almost halfway to costa, texture thin, lobes with rounded tips slightly toothed; costules

4 —4J mm apart; veins to 9 pairs, very oblique except lowest, 1 pair anastomosing; lower surface of rachis and costae closely very-short-hairy, some short hairs on lamina also. Sori inframedial; indusia thin, glabrous; sporangia bearing capitate hairs.

Distribution. Samoa, Solomon Is., Bismarck Arch.

I am not sure whether this is distinct from P. rotumaensis.

62. Pneumatopteris micropaleata Holtt., spec. nov.

infimam infimam Caudex erectus; stipes usque pinnam reductam 10 cm, usque pinnam infimae evolutam 50 cm; pinnae reductae 5 mm longae, superiores 10 mm; pinnae evolutae

maximae 1.8 inferiores basin versus leviter angustatae, non auriculatae; pinnae 17 X cm, ad basin aerophoris tumidis hemisphaericis praeditae, basi truncatae, apice caudatae, dimidio vel leviter ultra fere costam versus lobatae, lobis rectis non falcatis apice late rotundato leviter costae fertilium inter dentatis; pinnarum 3 —3J mm se distantes; venae solum membranam 7—8-jugatae, infimae anastomosantes, unijugatae ad sinus procur- rentes; pagina inferior rachidis pilis tenuibus rectis paleisque minutis praedita, costarum, costularum, venarum laminaeque paleis per-reductis filiformibus multis ornata. Sori paulo inframediales, infimi non divergentes; indusia glabra; sporangia pilis capitatis ornata.

Type: Holttum 58, Sabah, MtKinabalu, 1800m, on wet rocks near waterfall, Nov.

1972 (K; SING).

comb. 63. Pneumatopteris magnifica (Copel). Holtt., nov.

Mus. — Dryopteris magnifica Copel., Bull. Bishop 59 (1929) 11. Type: Parks 20187A, Fiji, Viti Levu,

1000 m (UC; BISH).

Caudex frond all reduced lowest erect; in 200 cm long; pinnae c. 5 pairs, 5 mm, upper- lowest little narrowed their most 1J mm long, not auricled; large pinnae a at bases;

X base lobed a little than largest pinnae 28 2.4 cm, truncate, apex acuminate; more J, costules veins lobes oblong, distinctly oblique, apex rounded entire; 5 mm apart; 7—9 pairs, lowest anastomosing, next to sinus-membrane; lower surface with slender short hairs on rachis and bases of costae. Sori medial or inframedial; indusia glabrous; sporangia

hairs. bearing many capitate

Distribution. Fiji.

comb. 64. Pneumatopteris transversaria (Brack.) Holtt., nov.

Nephrodium transversarium Brack, in Wilkes, U. S. Expl. Exp. 16 (1854) 187. — Dryopteris transversaria

Bot. Mus. 88. — Brause, Jahrb. s6 (1920) 104; C. Chr., Bull. Bishop 177 (1943) Type: Brackenridge,

Samoa (US). 320 BLUMEA VOL. XXI, No. 2, 1973

Caudex 6 of reduced lowest then erect; c. pairs pinnae, I cm, uppermost 3 cm long, lowest 1 or 2 pairs intermediate; large pinnae gradually narrowed to base, largest pinnae

1.6 lobed 20 X —2.0 cm, base broadly cuneate, apex evenly attenuate, about halfway to lower costa, lobes firm, slightly narrowed distally, toothed; veins 7 pairs; surface hairy hairs. on costae. Sori near costules; indusia small, firm, glabrous; sporangia bearing capitate

Distribution. Samoa, Fiji.

65. Pneumatopteris nephrolepioides (C. Chr.) Holtt., comb. nov.

Dryopteris nephrolepioides C. Chr., Brittonia 2 (1937) 268, 295, fig. 1, c, d. — Type: Brass 5354, Papua,

Mafulu, Centr. Div., on limestone, 1700 m (BM; NY).

Rhizome short, fronds closely tufted; stipe 3—4 cm, slender, scales ovate, 1 mm long; frond cm lower more to 40 long, pinnae 40 pairs; pinnae gradually reduced, widely

—6 spaced and deflexed; largest pinnae 13—16x5 mm, edges sinuous to crenate; veins

—8 in each basal white scales 7 pairs pinna, oblique, simple except acroscopic one, with at their tips on upper surface; minute capitate hairs on costae and veins beneath; short hairs surface small erect on upper throughout. Sori medial on veins, exindusiate; very capitate hairs on sporangia. Distribution. also collection Morobe Known from one in District, Sambanga:

Clemens 7207 (BM, B).

comb. 66. Pneumatopteris ligulata (Presl) Holtt., nov.

Presl, Bot. Fern Fl. — Lastrea ligulata Epira. (1851) 35; Copel., Philip, (i960) 327. Type: Cuming 74, Philippines (PRC; K, E).

Lastrea philippina Presl, Epim. Bot. (1851) 36. — Type: Cuming 393, Zebu (PRC). Linn. Soc. Bot. I. Nephrodium luerssenii Harr., J. 16 (1877) 29. — Type: Steere, Bulukai (K).

Dryopteris foxii Chr., Philip. J. Sc. 2 Bot. (1907) 208. — Lectotype: Copeland 940, Mindanao (US).

Rhizome basal scales frond short-creeping; stipe to 40 cm, narrow, hairy; to 70 cm, well basal pinnae c. 20 pairs, spaced, 2—4 pairs reduced, variable; largest pinnae to lobes side all shorter than base 20 X 3 cm, on basiscopic on acroscopic, unequally cuneate,

lobed to from lobes oblique, sometimes toothed apex caudate-acuminate, \ mm costa; fine short distally, sinuses broad; veins 9—12 pairs, free; both surfaces bearing spreading hairs. Sori submarginal; indusia short-hairy or glabrous; sporangia sometimes with capitate hairs.

Distribution. Philippines, Talaud I., beside rocky streams.

The of much smaller than that of. but all ofintermediates type D. foxii is P. ligulata, grades occur. The lectotype of D.foxii has glabrous indusia. Christ cited several specimens under

D. without in his herbarium Paris and foxii specifying a type; they are not at were pre- sumably on loan to him from Manila, now destroyed; the lectotype is from a Copeland collection cited by Christ.

67. Pneumatopteris keysseriana (Rosenst.) Holtt., comb. nov.

Dryopteris keysseriana Ro:enst., Fed. Repert. 10 (Feb. 1912) 333. — Type: Keysser 23, Sattelberg 1911 (not seen).

New Bot. — Schultze NE. Guinea, Dryopteris schultzei Brause, Jahrb. 49 (Aug. 1912) 19. T y p e: L. 253, Sepik (B).

Rhizome base with thin short-creeping, stipes tufted; stipe to 70 cm, ovate appressed scales; frond to 80 cm, pinnae 20 pairs or more; basal 2 or 3 pairs pinnae abruptly reduced in V R. E. HOLTTUM: Studies the family Thelypteridaceae 321

and remote; largest pinnae to 23 x 4 cm, base unequally broadly cuneate, basal basiscopic

lobed to mm from lobes segment reduced, apex caudate-acuminate, if —2 costa; slightly

wide costules —6 veins 16 tapering, separated by sinuses; 5 mm apart, to pairs; rachis and beneath. costae ± hairy Sori supramedial; indusia small with short hairs; sporangia sometimes bearing capitate hairs.

Distribution. NE. New Guinea, by streams in lowland forest. the ofBamler I have not seen type; at Kew is a specimen L. 56, Logaueng, also cited by

Rosenstock. The type of D. schultzei is firmer in texture, with short hairs between veins on lower surface.

68. Pneumatopteris caudata (Holtt.) Holtt., comb. nov.

caudatus Hort. Bot. Kew Blumea — cult. Pseudocyclosorus Holtt., 13 (1965) 133. Type: 545/68, no 12, New origin near Lae, Guinea, leg. Holttum.

of small Caudex erect; stipe 15 cm or more long, bearing c. 2 pairs very reduced pinnae, basal scales frond small, ovate; 30 cm, pinnae 15 pairs, lower ones slightly reduced, largest basal lobe 8x1.5 basiscopic minute, abruptly caudate, lobed to 1 from cm, apex mm costa, lobes oblique with rounded costules mm at veins apices; 3f apart, 45°; 9 pairs, free; lower surfaces of costules and with rachis, costa, veins minute acicular and capitate hairs. Sori supramedial; indusia short-hairy; sporangia eglandular.

Distribution. NE. New Guinea.

excisa comb. 69. Pneumatopteris (Holtt.) Holtt., nov.

Pseudocyclosorus excisus Holtt., Blumea 13 (1965) 133. — Type: Millar & Holttum NGF 18625, NE. E. New Guinea, Kerowagi, Highlands, 2000 m (K; L).

Differs from P. caudata: several basiscopic basal lobes in each pinna much reduced; lower sori medial, filling surface at indusia The maturity; minute, glabrous. type has

to iox Brass also from E. has 1.8 pinnae 1.5 cm; 31505, Highlands, pinnae to 14 x cm; there are two other collections.

70. Pneumatopteris versteeghii Holtt., spec. nov.

Rhizoma 2 arete breve-repens, mm diametro, stipites seriatos gerens. Stipes 8 cm longus, paleis minutis adpressis basi lamina translucens, —18 vestitus; 15 cm longa, pinnis usque infimae laminae 20-jugatis; pinnae ± abbreviatae, deflexae, remotiores; apex acuminatus, maximae basi valde profunde lobatus; pinnae 20 x 6 mm, asymmetricae (basiscopice anguste cuneatae acroscopice truncatae et auriculatae, auricula dentata), apice obtusae, marginibus dimidio costam lobis interdum leviter versus lobatae, dentatis; venae liberae, auriculis vulgo 2-jugatae (in pinnarum majorum 4-jugatae), tenues, fuscae, vena infima

basi sinus inferior omnino acroscopica terminata; pagina glabra; costa supra basin versus pilis praedita. Sori costam paucis prope utrinque uniseriati; indusia pilis brevibus multis vestita; sporangia pilis brevibus capitatis ornata.

BW New Div. Genofa Type: Versteegh 10259 >W. Guinea, Fak-Fak, Mts., 1100 m, in forest on steep rocky soil (L).

71. Pneumatopteris deficiens Holtt., spec. nov.

Caudex brevis, cm erectus; stipes usque 40 longus, glaber, paleis tenuibus ovatis vestitus; lamina cm infimae usque 35 longa, pinnis 8-jugatis; pinnae reductae, interdum 5 mm vulgo inferiores basin longae, majores; pinnae maximae versus sensim angustatae; pinnae 322 BLUMEA VOL. XXI, No. 2, 1973

fere ad lobis usque 12 X 2.2 cm, basi asymmetrice truncatae, acuminatae, costam lobatae, obliquis acutis leviter dentatis breviciliatis; costulae 5 —6 mm inter se distantes; venae subtus usque 8-jugatae, liberae, utroquepallidae prominentes; costa pilis minutis praedita. leviter indusia brevissimis Sori inframediales; parva, pilis paucis praedita; sporangia pilis capitatis destituta.

Posthumus of wet on earth Type: 3183, Flores, S. Roeleng, 1300 m, shady place banks in ravine (BO, K). Also Posthumus 3323, 3293 (BO, L).

comb. 72. Pneumatopteris regis (Copel.) Holtt., nov.

Univ. Bot. — Lastrea Dryopteris regis Copel., Cal. Publ. 18 (1942) 220. regis Copel., Philip. J. Sc. 78 (1951)

— C. 425, pl. 19. Type: King 486, Papua, mountains behind Wedan (MICH).

Rhizome and stipe not known; frond 120 cm long; reduced pinnae 6 pairs, 1—1 | cm and base little long wide, deeply lobed; largest pinnae 15 x 2.5 cm, truncate and a dilated

(most on acroscopic side), acuminate, lobed to 1 mm from costa; lobes 13x4 mm, costules slightly oblique, entire, tips rounded; 5 mm apart; veins 8 pairs, oblique, free; surfaces lower glabrous. Sori small, inframedial; indusia lacking (sori of type are old);

hairs few sporangia seen, no capitate observed on them.

imbricata 73. Pneumatopteris Holtt., spec. nov.

infimam Rhizoma ignotum. Stipes usque pinnam reductam 5 cm longus, usque lobatae pinnam evolutam infimam 45 —60 cm; pinnae reductae c. 1 cm longae, profunde

inter (lobis basalibus majoribus lobulatis), oppositae, superiores 3 —4 cm se distantes, inferiores reductis cm 2 cm. Lamina (pinnis exclusis) 100 longa, pinis 35-jugatis, oppositis; pinnae inferiores basi dilatatae, lobis infimis dentatis, acroscopicis rachin subtus, basi- medianae basi scopicis supra imbricantibus; pinnae usque24 x 3 cm, truncatae ± auriculatae,

fere ad costam sinubus late apice anguste acuminatae, marginibus lobatae, rotundatis, lobis leviter mm costulae mm oblongis, obliquis, falcatis, integris, in sicco 3 latis; 4\—5! inter infimaeambaead basin se distantes; venae liberae, usque 12-jugatae, marginem supra inferior membranas sinus protentae; pagina praeter margines et sinuum glabra. Sori leviter inframediales, parvi, exindusiati; sporangia pilis brevibus capitatis interdum ornata. Brooks river Type: 17775, Amboina, Hila, 200 m, on rock by (BM; L, BO). Distribution. Halmahera (Pleyte 370)-, Solomon Is., Guadalcanal ( Womersley &

Whitmore BSIP 1038). No other specimens seen.

74. Pneumatopteris petrophila (Copel.) Holtt., comb. nov.

Dryopteris petrophila Copel., Univ. Cal. Publ. Bot. 18 (1942) 220. — Lastrea petrophila Copel., Gen. Fil.

Blumea (1947) 139; Philip. J. Sc. 78 (1951) 424, pi. 18. — Pseudocyclosorus petrophila Holtt., 13 (1965) U3-

— Brass W. New NE. Lake moist rock Type: 11326, Guinea, Bele river, of Habbema, 2200 m, on

(MICH; L, BM).

Caudex —6 at base frond cm long, short, erect; stipe 5 cm, small appressed scales; 40 lower of pinnae c. 20 pairs, almost opposite throughout; 8 —10 pairs pinnae reduced, lowest and base 3 mm long wide, upper ones lobed; largest pinnae 4x0.8—0.9 cm, broadly cuneate and sometimes distinctly auricled on acroscopic side, short-pointed, lobes narrowed little blunt costules lobed about halfway to costa, falcate, a to tips; 3 mm slender and lower surface of rachis apart; veins 3 —4 pairs, free, prominent; copiously short-hairy; onrest oflower surface minutehairs, acicular and capitate; short hairsonupper with hairs annulus. surface between veins. Sori medial, exindusiate; sporangia capitate near R. E. HOLTTUM: Studies in thefamily Thelypteridaceae V 323

Also known from Distribution. several collections in NE. New Guinea, some

& NGF Clemens T. G. Walker with pinnae to 9 x i.o cm: Millar Holttum 15842, 7147 ;

7744, 8371, 8372, 8510.

sumbawensis comb. 75. Pneumatopteris (C. Chr.) Holtt., nov.

in — Dryopteris sumbawensis C. Chr. Rensch, Hedwigia 74 (1934) 231, tab. vii, fig. 1. Type: Rensch 578

Batu in Sumbawa, Dulang, 900 m, on rocks forest (BO; B).

frond Rhizome not known. Stipe 2 —3 cm long; 30 cm long, pinnae 35—40 pairs; lower 6 pairs oflower pinnae gradually reduced, lowest 3 mm long; pinnae ± deflexed,

a little narrowed to base on basiscopic side; largest pinnae 2.5—3.ox 1.0 cm; basal acro-

then or of adnate costules scopic segment free, 2 3 pairs segments separately to costa; or of free veins bases segments to 3 mm apart; 3 —4 pairs, dark but not prominent beneath, basal acroscopic one sometimes forked, rest simple and free; lower surfaces of all parts surface with short hairs bearing minute capitate hairs; upper very acicular and capitate between veins. Sori supramedial to medial, exindusiate; no hairs on sporangia. from Only known original collection. In its copious capitate hairs on lower surfaces matches also in this species P. costata, which occurs Sumbawa.

walkeri 76. Pneumatopteris Holtt., spec. nov.

Caudex brevis, erectus vel suberectus; stipes usque pinnam reductam infimam 20 cm, usque pinnam evolutam infimam 45 —50 cm, minute hirsutus, basin versus paleis latis lamina reductis adpressis vcstitus; pinnae reductae 2 —3 mm longae, alternae; (pinnis exclusis) 55 cm longa, pinnis 20-jugatis, alternis; pinnae inferiores 1—2-jugatae leviter

8—11 reductae; pinnae maximae cm longae, 1.1—1.9 cm latae, basi acroscopice truncatae, caudato-acuminatae basiscopice anguste cuneatae, apice (cauda 3 cm longa, dentata, sorifera), fere ad costam lobatae, lobis acroscopicis patentibus quam basiscopicis long- ioribus, lobis basiscopicis valde obliquis, lobis omnibus apice dentatis; costulae 4 mm subtus inter se distantes; venae usque 9-jugatae, liberae, pallidae, utroque prominentes; rachis subtus pilis erectis minutis praedita, pagina inferior cetera glabra. Sori mediales; indusia tenuia, glabra; sporangia glandulis destituta.

Type: T. G. Walker 9980, NE. New Guinea, Eastern Highlands, Waisa (BM).

DOUBTFUL SPECIES

Cyclosorus blastophorus Alston

C. Bol. Brot. Fl. W. Afr. 62. — T & blastophorus Alston, Soc. II, 30 (1956) 12; Trop. (1959) ype: Savory Keay FHI 25062, S. Nigeria (BM).

This species appears to be closely allied to Pneumatopteris unita (Kunze) Holtt. (no. 13

from I supra) which it differs in almost entire pinnaeand medial sori. have not seen spores,

the similar from Africa nor any good specimen except type. Some very specimens West have costular sori. XXI, No. 324 BLUMEA VOL. 2, 1973

Index

serial numbers of in the of basionyms, and of all binomials in Pneumatopteris (numbers refer to the species

are in bold present paper; new names type).

Aspidium (Dryopteris cont.)

sumbawensis C. 75 abortivum Bl.: 45 Chr.: Brause: abruptum Bl.: 45 superba 11

callosum Bl.: 8 vaupelii C. Chr.: 55

Ktze: caudiculatum Kunze: 50 venulosa O. 49

elatum Kuhn: 49 Goniopteris Brack.: eusorumThw.: 45 costata 17 T. Moore: 16 glaberrimumRichard: 60 forsteri Brack.: laeve Mett.: 29 glandulifera 4

lobbiana Fee: 8 longipes Bl.: 21 Fee: malagassium Kuhn: 13 madagascariensis 13 Fee: multijugum Chr.: 8 patens 13

Christella silvaticus Pappe & Rawson: 13

dentata sub (Forsk.) Holtt.: 19 Gymnogramma

Cyclosorus unita Kunze: 13

blastophorus Alston: doubtful Lastrca

ecallosus Holtt.: 32 cavitensis Copel.: 17

Presl: 66 glaberCopel.: 1 ligulata 66 lepidopodus Tard. & C. Chr.: 45 philippina Presl:

rotumaensis St John: 58 Nephrodium

viridis Copel.: 1 caudiculatum Presl: 50 costulare Dryopteris Bak.: 13

afra Chr.: 19 hudsonianum Brack.: 56 Bak.: berastagiensis C. Chr.: 20 lucidum 23 Harr.: 66 brooksii Copel.: 28 luerssenii Presl: bryanii C. Chr.: 54 nitidulum 59

christophersenii C. Chr.: 17 prismaticum Desv.: 50

T. Moore 61 deltipteraCopel.: 11 rodigasianum : transversarium 64 dewevrei Bonap.: 19 Brack.:

venulosum Hook.: 49 dicranogrammav. A. v. R.: 3 foxii Chr.: 66 Pneumatopteris

afra Holtt.: gladiata C. Chr.: 13 (Chr.) 19 26 inclusa Copel.: 20 angusticaudataHoltt.: Holtt.: keysseriana Roscnst.: 67 auctipinna 33 longifoliaBonap.: 38 basicurtata Holtt.: 31 brooksii 28 luzonica Chr.: 29 (Copel.) Holtt.: Chr.) macroptera Copel.: 51 bryanii (C. Holtt.: 54 8 magnifica Copel.: 63 callosa (Bl.) Nakai: caudata Holtt.: 68 mesocarpa Copel.: 9 (Holtt.) cheesmaniae 2$ microloncha Chr.: 27 Holtt.: christelloides Holtt.: microsora Copel.: 5 37 nephrolepioides C. Chr.: 65 comorensis Holtt.: 57 Holtt.: obstructa Copel.: 6 costata (Brack.) 17 deficiens Holtt.: oxyoura Copel.: 46 71

v. R.) parksii Ballard: $ dicranogramma(v. A. Holtt.: 3 ecallosa (Holtt.) Holtt.: petrophila Copel.: 74 32

regis Copel.: 72 egenolfioides Holtt.: 22

remotipinna Bonap.: 38 excisa (Holtt.) Holtt.: 69 60 rurutensis Copel.: 17 glaberrima(Richard) Holtt.:

sambiranensis C. Chr.: 38 glabra (Copel.) Holtt.: 1

schultzei Brause: 67 glandulifera(Brack.) Holtt.: 4

sogerensis Gepp: 48 hudsoniana (Brack.) Holtt.: 56 humbertii stokesii E. Brown: 7 Holtt.: 42 imbricata subappendiculata Copel.: 10 Holtt.: 73 C. incisa Holtt.: subpennigera Chr.: 14 53 inclusa subspinosa C. Chr.: 4 (Copel.) Holtt.: 20 R. E. HOLTTUM : Studies in thefamily Thelypteridaceae V 325

(Pneumatopteris cont.) (Pneumatopteris cont.)

jermyi Holtt.: 34 stokesii (E. Brown) Holtt.: 7

kerintjiensis Holtt.: 41 subappendiculata(Copel.) Holtt.: 10

keysseriana (Rosenst.) Holtt.: 67 subpennigera (C. Chr.) Holtt.: 14

laevis (Mett.) Holtt.: 29 sumbawensis (C. Chr.) Holtt.: 75

laticuneata Holtt.: 39 superba (Brause) Holtt.: 11

latisquamataHoltt.: 18 tobaica Holtt.: 52

ligulata(Presl) Holtt.: 66 transversaria (Brack.) Holtt.: 64

longipes (Bl.) Holtt.: 21 truncata (Poir.) Holtt.: 45

lucida (Bak.) Holtt.: 23 var. loyalii Holtt.: 45

macroptera (Copel.) Holtt.: 51 unita (Kunze) Holtt.: 13

magnifica(Copel.) Holtt.: 63 usambarensis Holtt.: 40

Holtt.: mesocarpa (Copel.) Holtt.: 9 vaupelii (C. Chr.) 55

michaelis Holtt.: 43 venulosa (O. Ktze) Holtt.: 49

microauriculata Holtt.: 36 versteeghiiHoltt.: 70

microloncha (Chr.) Holtt.: 27 walkeri Holtt.: 76

micropaleata Holtt.: 62 Polypodium

nephrolepioides(C. Chr.) Holtt.: 65 brackenridgei Hook.: 4

nitidula (Presl) Holtt.: 59 dentatum Forsk.: sub 19

novae-caledoniae Holtt.: 44 microdendron Eaton: 12

obliqua Holtt.: 30 muricatum Bak.: 4

obstructa (Copel.) Holtt.: 6 oppositifolium Hook.: 15

oppositifolia(Hook.) Holtt.: 15 pennigerumForst.: 16

oxyoura (Copel.) Holtt.: 46 polycarpon Hook. & Arn.: 12

Holtt.: Bak.: 12 papuana 3 5 stegnogrammoides

parksii (Ballard) Holtt.: 5 truncatum Poir.: 45

patentipinnaHoltt.: 24 Polystichum

pennigera (Forst.) Holtt.: 16 truncatum Gaud.: 60

pergamacea Holtt.: 47 Pseudocyclosorus

petrophila(Copel.) Holtt.: 74 caudatus Holtt.: 68

prismatica (Desv.) Holtt.: 50 excisus Holtt.: 69

regis (Copel.) Holtt.: 72 petrophilus (Copel.) Holtt.: 74 remotipinna (Bonap.) Holtt.: 38 Stegnogramma

rodigasiana (T. Moore) Holtt.: 61 sandwicensis Brack.: 12

rotumaensis (St John) Holtt.: 58 Thelypteris

sandwicensis (Brack.) Holtt.: 12 hawaiiensis Reed: 12

sibelana Holtt.: 2 sevillana Reed: I

sogerensis (Gepp) Holtt.: 48

325