SMITHSONIAN CONTRIBUTIONS TO 

BOTANY 

NUMBER 8 

F. R. Fosberg Three Indo-Pacific 

Thelypteris Species and M.-H. Sachet 

Reinterpreted and a 

New African Species 

Described 

SMITHSONIAN INSTITUTION PRESS 

CITY OF WASHINGTON 

1972 
ABSTRACT 

Fosberg, F. R., and M.-H. Sachet. Three Indo-Pacific Thelypteris Species Reinterpreted and a New African Species Described. Smithsonian Contributions to Botany, 

number 8, 10 pages, 3 figures. 1972.-Two species of Thelypteris which have been 

confused and to which a number of names have been applied were found to be 

separable by a number of characters. They must now be called Thelypteris opulenta 

and T. wagneri. Their distribution is Indo-Pacific, extending to Africa, and T. opulenta introduced into tropical America. T. wagneri and an African species, T. 

fadenii, are here described as new, The name Thelypteris interrupta, heretofore 

applied to this complex, is shown to be the correct name for what has commonly 

been called Dryopteris gongylodes and Thelypteris totta. 

Library of Congress Cataloging in Publication Data 

Fosberg, Francis Raymond, 1908- 

Three Indo-Pacific Thelypteris species reinterpreted and a new African species described. 

BiblioaraDhv : D. 

(Smithsonian contributions to botany, no 8) 

-, . 1. ThGlypteris. I. Sachet, Marie-HClhe, joint author. 11. Title. 111. Series: Smithsonian 

Institution. Smithsonian contributions to botany, no. 8. 

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Official publication date ic handstamped in a limited number of initial copies and is recorded 

in the Institution’s annual report, Smithsonian Year. 

For sale by the Superintendent of Documents, U.S. Government Printing Office 

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F. R. Fosberg Three Indo-Pacific 

Th elyp teris S p ec ies and M.-H. Sachet 

Reinterpreted and a 

New African Species 

Described 

Among the common Indo-Pacific ferns are a pair 

of Thelypteris (Cyclosorus) species with lobed pinnae, nearly marginal sori, a sterile zone on both 

sides of the midrib of the pinna, and with at least 

a few, usually many, yellow particles (glands?) on 

the lower surface of the blade, especially on the lobes. 

They may be found in herbaria under many names, 

some properly applied to ferns of this relationship, 

others not. The present attempt to unravel the specific taxonomy and nomenclature of this group was 

stimulated by our realization that a plant of the 

Marquesas Islands, described by Dr. Elizabeth Brown 

as Dryopteris sulfurea, was apparently identical with 

what we had previously called Dryopteris extensa 

(Blume) Kuntze, or Dryopteris interrupta (Willd.) 

Ching, in Micronesia. 

Since the latter two species were originally described from Java and India, respectively, it seemed 

appropriate to examine past treatments to determine 

what plants were originally so named. This survey 

showed considerable variation in opinions as to 

whether one or several species are involved and as 

to the names to be applied to them. The criteria by 

which they were separated also varied considerably, 

from author to author. Most of the purported differences are rather intangible-depth and shape of 

lobing, ranges of thickness and habit of rhizome, 

F. R. Fosberg, Department of Botany, Smithsonian Znstitution, Washington, D. C. 20560. M.-11. Sachet, Department 

of Botany, Smithsonian Institution, Washington, D. C. 

20560. 

abundance of yellow glands, and details of venation. 

An attempt to sort the available material from the 

western Pacific in the United States National Herbarium according to these criteria was not very 

successful, except to suggest a geographical separation 

of two entities. A further attempt on these lines, 

separating the Malesian from the Micronesian and 

Polynesian material, showed a reasonably consistent 

difference in the shapes of the leaf apices of the 

plants from these two regions. The Pacific specimens 

have the apical segment broad at base, pinnatifid or 

at least deeply lobed in the basal half, gradually 

attenuate distally into a caudate tip. Most of the 

specimens from the Malesian region have the apical 

segment essentially similar to the pinnae, linearoblong, with one or two similar but reduced segments 

at its base, either separate from it or fused with it. 

These segments are abruptly much smaller than the 

next lower pinnae. 

It is surprising that such a clear-cut difference 

apparently has not been used before to distinguish 

these plants. Those with the pinnatifid tip include all 

material seen from Polynesia and Micronesia and a 

few specimens from the Philippines, Malacca, Pulu 

Penang, Java, Rawak, Thailand, Ceylon, and westward to East Africa, and those from tropical America 

(introduced) . They have most frequently been called 

Nephrodium extensum or Dryopteris extensa, but 

this epithet has also on occasion been applied to the 

other plant. 

The plant with the linear-oblong terminal segment 

1 
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

is found from New Guinea, the Malesian and Philippine Islands, westward to India and Ceylon, Central 

Africa and Fernando Po Island. It has commonly 

been called Nephrodium or Dryopteris pteroides, or 

more recently Dryopteris interrupta or Cyclosorus 

interruptus. 

Whether we are dealing with two species or only 

with varieties of one is a difficult question. Most 

pteridologists, including Copeland and Holttum, have 

maintained the two as separate species, though not 

always on the same grounds. Wagner, in a manuscript on file in the U. S. National Herbarium Division of Ferns but apparently never published, regarded them as varieties of one. They are big, 

unwieldy ferns and not always fairly represented by 

herbarium material. Unfortunately, the rhizome is 

lacking on most specimens, making it difficult to 

apply the criterion that one has a short, thick erect or 

ascending rhizome with congested fronds, whereas 

the other has a more slender, creeping rhizome with 

scattered fronds. Our present impression is that this 

is not a very reliable difference, but that it does represent a real tendency. 

Since the difference in the apical segment seems 

reasonably clear, and since that is supported by a 

number of less easy-to-use characters, and since the 

extensive geographical ranges have only partial overlap, we are for now regarding the two plants as 

separate species. Future studies on a more representative range of material may possibly show this to be 

wrong. In any event, the two form a closely related 

group, defined by yellow glands, lobing only part way 

to the costae of the pinnae, and marginal or submarginal positions of the sori, which are usually lacking in 

the united portions of the pinnae. The margins tend to 

be a bit revolute around the sori, making the lobes 

appear narrower and the sori more marginal than 

they really are. The indusia are well developed. The 

spores are bilateral and very deeply rugose with short 

curved or sigmoid ridges. The group seems related to 

the plant that is currently called Thelypteris totta 

(Thunb.) Schelpe or T. goggilodus (Schkuhr) Iwatsuki, which is discussed under Thelypteris interrupta. 

This may be only a superficial resemblance, due to 

the submarginal sori. The group with yellow glands 

has been confused with a number of other species, 

notably T. pennigera (Forst. f.) Allan and its close 

relatives. 

The rest of this paper will attempt to sort out the 

synonymy and establish the correct names for the two 

species under discussion, and to discuss two other 

species that enter into the problem. Unfortunately, 

several of the synonyms are hard to place without 

authentic material. These are indicated by a query. 

All that are definitely included in the two synonymies, 

without a query, and not as “sensu” particular 

authors, are plants with the above combination of 

features that characterize the group. 

The names that seem to have been based on plants 

of this affinity were all studied with reference to their 

original descriptions and types (when available), as 

well as to subsequent discussions of their typification, 

and in the light of the combinations of characters of 

the two species as indicated above. It was fairly easy 

to assign most of the synonyms to one or other of the 

two species, thus compiling the synonymies listed 

FIGURE 1 .-Thelypteris opulenta, portion of frond showing 

terminal segment and lobing of pinnae. 
NUMBER 8 3 

FIGURES 2 AND 3.-Thelypteris wagneri, portions of fronds showing terminal segment and 

lobing of pinnae. 

under the species. Names that have been misapplied 

to these species are indicated as “sensu” the misapplying author, “non” the correct author of the name. 

Herbaria where the specimens cited were examined 

are indicated by the standard Index Herbariorum 

abbreviations except those still in our possession and 

not yet deposited, which are marked (Fo) . 

Thelypteris opulenta (Kaulf.) Fosb., new 

combination 

Aspidium opulentum Kaulf., Enum. Fil. 238, 1824. Type: 

Polystichum pennigerum sensu Gaud., Bot. Voy. Uran. 328, 

Nephrodium impressum Desv., Prodr. 259, 1827. Type: 

Aspidium extensum Blume, Enum. PI. Jav. 156, 1828. 

Guam, Chamisso. 

1826, non Polypodium pennigerum Forst. f. 1786. 

Timor, Hb. Desv. (P) . 

Type: Pulu Pinang, Blume. 

Aspidium multijugum Wall., Cat. 348, 1829, nomen nudum. 

Aspidium prionophyllum Wall., Cat. 355, 1829, nomen 

Aspidium terminans Wall., Cat. 386, 1829, nomen nudum. 

Nephrodium cumingii J. Smith in Hook., J. Bot. 3:411, 

Nephrodium terminans J. Smith, Bot. Mag. 72 (Comp.) : 32, 

Lastraea malaccensis Presl, Epim. Bot. 35, 1849 [ 185 11. 

Nephrodium conioneuron Fee, Gen. Fil. 308, 1852? 

Nephrodium extensum (Blume) Moore, Ind. Fil. 91, 1858. 

Aspidium pteroides vars. a?, b (pro parte), Mett., Ann. 

Mus. Lugd. Bat. 1:231, 1864. 

Aspidium pteroides sensu Luerssen, J. Mus. Godeffr. 1 :55, 

1873, non Polypodium pteroides Retz. 1791. 

Nephrodium parasiticum var. multijugum C. B. Clarke, 

Trans. Linn. SOC. Bot. 1:533, 1880. Type: Penang, 

Wallich 348. 

Dryopterir extensa (Illume) Kuntze, Rev. Gen. P1. 2:812, 

1891. 

nudum. 

1841, nomen nudum. 

1846, nomen nudum. 

Type: Malacca, Cuming 391. 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Nephrodium wakefieldii Baker, Ann. Bot. 5:326, 1891. 

Type: Mombasa, Kenya, Wakefield. 

Dryopteris sulfurea E. Brown, Bish. Mus. Bull. 89:23, 1931. 

Type: Marquesas, Brown 293. 

Phegopteris pennigera sensu Hosokawa, Trans. Nat. Hist. 

SOC. Formosa 26:233, 1936, non Polypodium pennigerum Forst. f., 1786. 

Dryopteris impressa (Dew.) Posth., Verh. K. Akad. Wettensch. Amst. 36(5) : 14, 1937. 

Cyclosorus extensus (Blume) Ching, Bull. Fan Mem. Inst. 

Biol. Bot. 8:182, 1938. 

Cyclosorus interruptus sensu Wagner and Grether, Occ. 

Pap. Bish. Mus. 19:50, 1948, non Pteris interrupta 

Willd., 1794. 

Thelypteris extensa (Blume) Morton, Amer. Fern J. 49: 

113, 1959. 

Thelypteris interrupta sensu Stone, Micronesica 2 : 3, 1966, 

non Pteris interrupta Willd., 1794. 

Thelypteris sulfurea (Brown) Reed, Phytologia 17 : 3 18, 

1938. 

Rhizome tending to be rather thick and short; 

fronds broad, .oblong, pinnate; pinnae deeply lobed 

usually one-half to three-fourths the way to the midrib, the lobes tapering somewhat, the apex of fronds 

deeply pinnately lobed or pinnatifid in lower half 

strongly tapering to a caudate tip; sori submarginal 

around the lobes, appearing marginal because of revolute margins of lobes; yellow glands on lower surface 

of pinnae especially on veins and near tips of lobes. 

SPECIMENS EXAMINED.-~NSULAE OCEANI PACIFICI : Forster (BM) (a wilted specimen but probably this). 

TUAMOTU Is.: Bow Island [Hao Atoll], Barclay 165(?) 

MARQUESAS Is.: Dupetit-Thouars in 1843 (P, 2 sheets). 

Uapou I.: alt. 200 m, Jones 1105 (BISH, BM); Hakahetau valley above village, 25-250 m, common name: 

“Autapa’amei,’’ Decker 2283 (Fo, US), Uahuka I.: Hannay, 150 m, Jones 2662 (BISH); Jones 2665 (BISH); 

Upper Ha’avei valley, 200-250 m, Decker 2469 (Fo); intenor Vaipae’e Valley, 200 m, Decker 1537 (Fo), 2549 

(Fo, US) ; Upper Vaipae’e, 170 m, Decker 1982 (Fo, US) ; 

west side of Vaipae’e, 100 m, Decker 2793 (Fo, US). 

Hiva’oa I.: 300 m?, Brown 793 (BISH, type of Dryopteris 

sulfurea); central Puamau, 85 m, Decker 802 (Fo, US); 

Puamau village, 25 m, Decker 883 (Fo, US); SW Puamau, 

200 m, Decker 2012 (Fo); central Puamau valley, 2 km 

inland, 225 m, Decker 1080 (Fo) ; E. Puamau valley, 175 m, 

Decker 2233 (Fo) ; Puamau, along Puamau-Atuona trail, 

500-650 m, Decker 1200 (Fo). Nukuhiva I.: Hakaui, 

Brown 790 (BISH) ; Taiohae, Brown 31 (BISH, 2 sheets); 

Taiohae-Hatiheu trail, 450 m, Decker 21 70 (Fo, US). Fatuhiva I.: Omoa Valley, 500 m, Brown 987 (BISH); Hanavave, 100 m, Jones 2808 (BISH); s.l., Cheesman in 1925 

(BM). Motane I.: 100 m, Jones 2725 (US, BISH). Eiau 

I.: 700 m, Jones 1525 (US, BISH). Tahuata I.: Vaitahu, 

150 m, ]ones 2745 (BISH). 

(BM). 

SOCIETY IS.: Meetia I.: Fatia-Po to Fareura, 250 m, 

St. John 24289 (BISH). 

AUSTRAL Is.: Rimatara I.: Stream near Mutuaura, 20 

m, Fosberg 22053 (US, BISH). Rurutu.: Moerai, 10 m, 

St. John 26636 (BISH); Puputa Valley, above Moerai, 

25 m, Fosberg 12937 (BISH). 

SAMOAN Is.: Tau: 1 mi. S Siufaga, 60 m, Yuncker 9020 

(BISH) . BONIN Is.: Nakata-ioto, Tuyama (reported by Ito, 1937: 

714). 

MARIANAS Is.: Gaudichaud (P, 2 sheets) ; Gaudichaud 

in 1824 (P). Agrigan I.: canyon on E side, Fosberg 31633 

(US, BISH, Fo, MICH). Alamagan I.: s.1. Falanruw 1923 

(US) ; 60 m, Anderson 412 (US, BISH, Fo, MICH, IJ, 

NY). Saipan I.: s.1. Kanehira 2229 (NY); Hosokawa 

6659 (A) ; Raishingan, Hosokawa 8032 (BISH, A). Rota 

I.: Savana, Hosokawa 7593 (BISH, A), 7646 (A); Necker 

R 88 (US); 1300 ft, Kondo in 1952 (BISH); 800 ft, 

Grether 4470 (US) ; slopes above As Malote, S side of 

island, 250 m, Fosberg 32875 (US, BISH, Fo, MICH) ; 

NW coast of W end of main part of island, between Rota 

and Tataacho Point, Fosberg 25137 (US, BISH, Fo, 

MICH, IJ). Guam 1.: Mertens (BM); Wagner 3292 

(US); Connover 534 (US); Herb. Richard in 1824 (P); 

Marche 167 (P), 47 (P), 73 (P) ; Haputo Point, Johnson 

and Necker 2 (US) ; Mt. Santa Rosa, Glassman 272 (Fo) ; 

Hosaka 3251 (US, BISH, Fo) ; 1 mi. S of Taguac, on north 

plateau, Fosberg 35479 (US, BISH, Fo, MICH, IJ, NY) ; 

Agaiia, Safford and Seale 2086 (US), 2088 (US), 1089 

(US) ; Agania Swamp, E of Agniia, 1 m, Fosberg 31226 (US, 

BISH, Fo, MICH, IJ, NY); 5 m, Bryan 2084 (US, NY, 

BISH) ; Agania Marshes, Grether 3823 (US, BISH) ; lower 

Fonte River, 15 m, Fosberg and Evans 46297 (US, BISH) ; 

2 km S of Piti, Necker 201 (US) ; shores of Piti Harbor, 

Bartsch 202 (US), 110 (US); Yigo, 180 m, Bryan 1239 

(BISH) ; Lonfit River, Grether 3720-A (US) ; Sigua River 

Valley, Grether 3728 (US, BISH) ; near junction of Lonfit 

and Sigua Rivers, Grether 3817 (US, BISH) ; W of Mapao, 

3 mi WNW of Talofofo Bay, 100 m, Fosberg 35455 (US, 

BISH, Fo, MICH, IJ); Talofofo, Mt. Almzgosa, Hosaka 

3184 (US, BISH, Fo); Umata, Marche 243 (P). 

CAROLINE Is.: Sonsorol 1.: Berry 2 (US). Palau Is.: E 

end Koror I., Fosberg 50632 (US, BISH); Tor Uil I., between Koror and Malakal I., 15 m, Fosberg 47598 (US, 

BISH) ; Aulupse’el I., 10-15 m, Fosberg 47556 (US, BISH, 

Fo, MICH, IJ, NY, L); E end of Urukthapel I., road to 

lighthouse, 1-5 m, Fosberg 32036 (US, BISH, Fo, MICH). 

NEW GUINEA : Kaiser-Wilhelmsland, Walder von Woble, 

180 m, Schlechter 26349 (BISH). Rawak I.: Gaudichaud 

in 1819 (P); in 1845 (?) (P); Gaudichaud (P). 

TIMOR I.: Herb. Desvaux (P, prob. type of Nephrodium 

impressum). 

PHILIPPINES: Luzon I.: Prov. Laguna, Los Banos,ii 

Mount Maquiling, Elmer 27905 (BISH) ; Prov. Ilocos 

Norte, Ramos 7772 (BISH); Prov. Ilocos, Burgos, Ramos 

27212 (US). Balabac I.: Mangubat 415 (NY, US). Mindanao I.: Cuming 293 (BM); District of Zamboanga, 

Santa Maria, Reillo 26502 (US). 

MALAY PENINSULA: Ayer Hitam: Matthew in 1908 (K, 
NUMBER 8 5 

2 sheetsj; G. Pardok, Matthew in 1911 (KO). NegriSembilan: Lukut Fort, sea level, Hose 272 (K). Malacca: 

Cuming 391 (K, 3 sheets, one young, doubtful, US, P, 

isotypes of Lastrea malaccensis Presl, photo of Paris sheet 

in US, Morton Neg. No. 17450) [cited by J. Smith (1841 : 

411) as his Nephrodium cumingii]. Penang: Wallich 1829 

(K), 348 (K, P, photos in US, basis of Aspidium multijugum Wall., Morton Neg. Nos. 17451, 17452), 355 (P, 

photos in US, basis of Aspidium prionophyllum Wall., 

Morton Neg. Nos. 17453, 17454) ; Curtis 3367 (K). Pulu 

Penang: Blume (photo in US of specimen in L, type of 

Aspidium extensum Blume) . Kedah: Pulau Langkawi, Holtturn 17414 (US); West Coast, Pulau Dayang Bunting 

(Pulau Langkawi) Henderson 29132 (US). Johore: Vesterdal 295 (US). 

SINGAPORE I.: Blakang Mati I.: Sea level, MolesworthAllen 2523 (US). 

JAVA: Goenung Parang, Blume (BM). 

THAILAND (SIAM): Murok Lek, Marcan 806 (BM); 

Bandon, Surat, near sea level, Smith 3004 (BM), 3007 

(BM, US); Setun Puket P. Siam, Smith 2469 (BM, US), 

2459 (US). 

CEYLON: G. Wall in 1884 (PDA); Thwaites (BM); 

“Gandola,” Herb. 1. Smith 2206 (BM) ; Kandy, Chevalier 

[?] (BM), [Thwaites] C. P. 975 (PDA); s.1. ex Herb. 

William Ferguson 238 (US, 2 sheets). 

CHAGOS Is.: Diego Garcia: Minni Minni, Stoddart 832 

AFRICA: Kenya: Mombasa, Wakefield (K, type of 

Nephrodium wakefieldii, 3 sheets) ; Tanzania (as Tanganyika Terr.) : Pangani, Tanga Prov., Tanner 2836 (K) ; 

Ulugurus, Kimbosa, 1000 ft, Bruce 133 (US, BM). Pemba 

I.: Vaughan 456 (K). Mozambique: Manica e Sofala: 

Cheringoma, Inhaminga, Mendoca 4377 (BM) ; Zambezia, 

Mendoca 1448 (BM). 

COSTA RICA: Prov. Puntarenas: 6 km S of airstrip, Osa 

Peninsula, 500 ft, Mickel 2770 (ISC, LP, US). 

MARTINIQUE : Stehle 4986 (US). 

BRITISH GUIANA: Maguire and Fanshawe 22983 (US). 

COLOMBIA: Putumayo, 15 km NW of Puerto Asis, 1000 

ECUADOR : Santiago-Zamora, Taisha, 1500 ft, Caralet 

(K). 

ft. King & Gueuara 6222 (US). 

& Pennington 7752 (US). 

We have not been able to examine Chamisso’s 

specimen, described by Kaulfuss as Aspidium opulentum, but the description is very clear and fits the 

species under discussion very well. This is the only 

species in Guam that corresponds to the Kaulfuss description, and there seems no reasonable doubt that 

the epithet opulenta is correctly applied here. 

The type of Nephrodium impressum Desv., from 

Timor, is probably a specimen from the Desvaux 

herbarium in Paris which is this species. 

Blume’s description of Aspidium extensum generally fits; a specimen in Leiden, named in Blume’s 

hand, photographed by C. V. Morton (Neg. No. 

1128), and considered to be the holotype by Morton, 

has the caudate pinnatifid apex of T. opulentum. 

Nephrodium conioneuron Fee has usually been 

referred to Dryopteris pteroides (Retz.) 0. Kuntze or 

D. interrupta (sensu auct., non Polypodium interruptum Willd.) . The type has not been located by us. 

A comparison of the description of this and Nephrodium oreopteris Fee, published together, shows that 

N. conioneuron should have more deeply lobed pinnae than N. oreopteris. Isotype material of the latter 

represents one of the few intermediates between T. 

opulenta and T. wagneri. Its pinnae are more deeply 

lobed than those of T. wagneri. Since the pinnae of 

N. conioneuron are described as still more deeply 

lobed, it seems safe to refer N. conioneuron tentatively here. 

Aspidium pteroides var. a Mett. was based on 

Nephrodium conioneuron Fee, and var. b Mett. was 

based on three synonyms, two of which, Aspidium 

opulentum Kaulf. and A. extensum Blume, belong 

here. 

No descriptions were provided for Aspidium terminuns, Nephrodium terminans, and Nephrodium cumingii. One of the two specimens cited with the 

latter, Cuming 391 from Malacca, is T. opulenta. The 

other, Cuming 293, from Mindanao, is probably T. 

wagneri, but the US specimen of this number is fragmentary and the determination doubtful. We have 

not seen the Wallich specimen of Aspidium terminuns. Smith merely cites A. terminans Wall. and N. 

cumingii J. Smith as the protologue of N. terminans, 

hence it is also a nomen nudum, unless provided with 

a description in some publication missed by the indexers. 

Ching refers Nephrodium parasiticum var. multijugum C. B. Clarke, Wall. Cat. 348, from Penang, 

to Dryopteris extensa (= opulenta), and our examination of Wallich’s material confirms this. 

The type of Nephrodium wakefieldii, from East 

Africa, certainly belongs here, but the material seen 

from there is very luxuriant and the pinnae are very 

deeply lobed. N. wakefieldii was reduced by Schelpe 

( 1965 : 266-267) to Thelypteris extensa (Blume) 

Morton, which belongs here. 

We have seen ample material of Dryopteris sulfurea from the Marquesas, including that studied by 

Dr. Brown, and find no distinction from T. opulenta. 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

References of Guam material of this species to derivatives of Polypodium pennigerum Forst. f., of P. 

pteroides Retz., and of Pteris interrupta Willd. are 

simply misidentifications. The two latter are discussed 

elsewhere in this. article. Polypodium pennigerum is 

a Thelypteris that superficially looks a bit like T. 

opulenta, but lacks yellow glands, has the sori differently disposed, and does not actually seem very closely 

related. 

Thelypteris opulenta var. hirsuta (Ching) Fosb., 

new combination 

Cyclosorus interruptus var. hirsutus Ching, Bull. Fan Mem. 

Inst. Biol. 8: 186, 1938. Lectotype: Thailand, Rock 

1716. 

This differs from var. opulenta in the short pale pubescence of the underside of the frond. This pubescence is most marked in the Rock 1716 specimen 

cited, here designated as lectotype. The Smith 473 

specimen is intermediate between T. opulenta and 

T. wagneri (see discussion below under T. wagneri) . Thailand, Prov. Chiengmai, Chieng Rai, lower 

slopes of Doi Chang Mt. near Hue San (420-1, 260 

m, Rock 1716 (US, lectotype, annotated by Ching). 

Thelypteris wagneri Fosberg and Sachet, new species 

Polypodium pteroidcs Retz., Obs. Bot. 6:39, 1791, type: 

Ceylon, Konig; non Thelypteris pteroidea (Klotzsch) 

Tryon, Rhodora 69:8, 1967, which was based on Polypodium ptcroidcum Klotzsch, 1847 ; non P. ptcroides 

Lam., 1778. 

Aspidium ptcroidcs (Retz.) Swartz, Schrad. J. Bot. 1800 

(2) :33, 1802. 

Ncphrodium orcopteris Fee, Gen. Fil. 304, 1852, type: 

Philippines, Cuming 48; non Thelypteris orcopteris 

(Ehrh.) Rydb., 1917. 

Aspidium pteroidcs var. b Mett., Ann. Mus. Lugd. Bat. 

1:231, 1864, pro parte. 

Dryoptcris ptcroidcs (Retz.) Kuntze, Rev. en. P1. 2:813, 

1891. 

TYPE.-Java, Raciborski in 1897. 

Rhizoma repens, glabrum; frons oblonga, ad apicem abrupte contracta, segment0 terminali pinnam 

simulante, pinnis lobatis lobis brevibus vix ad dimidium costae fissis, infra flavo-granulatis, soris submarginalibus. 

Rhizome slender, 3-4 mm thick, creeping, shortly 

but rather densely hirtellous, covered by brown linearlanceolate caducous scales ; fronds widely scattered, 

long-stipitate, up to 1 m or more long, the stipes 

dark brown, minutely retrorse-hirtellous, clothed on 

basal several cm with brown linear-lanceolate scales, 

the blades pinnate, oblong, up to 50 cm long, abruptly 

contracted toward apex, the rachis lighter brown than 

stipe, hirtellous, the pinnae lance-linear to linear oblong, tapering in distal third to an acuminate tip, 

13-28 cm long, 1-2.5 cm wide, the width and length 

varying independently, well separated, alternate, the 

lower sometimes subopposite or even opposite, the 

lowermost somewhat to rather strongly reduced on 

some specimens, these apparently early fronds in the 

dcvelopment of the plant (Smith 190, 299, Molesworth-Allen 2844, Merrill 594, Williams 2218, 

Otanes, 18728, Topping 659), pinna lobed to about 

half or less the distance to the midrib, lobes obliquely 

rounded or obtuse to apparently acute but this because of the margins curving round sori, lobes appearing slightly falcate, pinna sparsely pilose above 

and beneath, especially on the veins, and densely so 

on upper side of costa, underside with yellow bulbous 

or globose particles or “glands” scattered on veins and 

especially near tips of lobes, 3 veins from each lobe 

uniting in a vein running to the sinus between 2 

lobes, sori submarginal but appearing marginal because of the somewhat revolute margins, sometimes 

1-3 pairs below sinuses but a broad sorus-free band 

along pinna-midrib, this rarely with a few scattered 

sori, distal two or three pinnae somewhat reduced, 

terminal one essentially identical with lateral ones, 

little dilated basally; indusium orbicular, somewhat 

hirtellous; annulus with 15 or 16 cells; spores bilateral, very deeply rugose with short curved or sigmoid 

ridges. 

DISTRIBUTION.-New Guinea to Ceylon, westcentral Africa (Oubangi), and Fernando Po Island 

in the Gulf of Guinea; most abundant in the Philippines, Malesia, and Indo-China. 

SPECIMENS EXAMINED.-NEW GUINEA : Papua: Central 

Distr., near Douramoko Village, Vanapa River, 50 ft, 

Shoddc 2637 (US). 

HAINAN: Wang 33474 (US; very young wilted specimen, 

determination doubtful). 

PHILIPPINES : Cuming 48 (US, isotype of Ncphrodium 

orcopteris Fee?) . Luzon I. : Rizal Prov. : Montalban, Lohcr 

944 (US), Ramos 5203 (US); Antipolo, Ramos and 

Edoiio 29542 (US). Tayabas Prov.: Edaiio 26947 (US); 
NUMBER 8 7 

Atimorian, Topping 1310 (US), 1315 (US). Pangasinan 

Prov. : Umingan, Otanes 18278 (US). Laguna Prov. : 

McCregor 2311 I (US) ; Los Baiios, Topping 659 (US), 

680 (US); Mount Maquiling, Elmer 17905 (US). Ilocos 

Norte Prov.: Ramos 7756 (US). Mindoro I.: Puerto 

Galera, Ramos 46426; Calapan, Merrill 984 (US) (in most 

respects this species, but the terminal segment slightly 

broader and more lobed at base than usual, thus tending in 

this character toward T. opulenta). Balabac I.: Mangubat 

420 (US). Culian I.: Merrill 484 (US), 594 (US). Mindanao I.: Gulf of Leyte, Cuming 293 (US, doubtful, specimen fragmentary) ; Davao Distr. : Copeland 636 (US) : 

Todaya, Mount Apo, Elmer 11, 111 (US, BISH). Zamboanga Prov.: Binalanay, Copeland in 1917 (US); San 

Ramon: Copeland 1604 (US), Williams 2436 (US); Sax 

River, Williams 2218 (US). Palawan I.: San Antonio Bay, 

Merrill 5254 (US). Sulu Is.: 8 mi SE of Cagayan, Santos 

4677 (US). 

MALAY PENINSULA: Kedah: Alorstan (Alor Sta.) , Ridley 14764 (K); West Coast, Dayang Bunting, Henderson 

29144 (K). Malacca: Crifith (K). Perlis: Kalu Bukit, 

Kiah 35250 (K); Bukit Ketri, Matthew in 1914 (K). 

Pahang: Ayer Besar, P. Tioman, Henderson 18767 (K, 

US). Penang: Molesworth-Allen 2844 (US). 

SINOAPORE I. : Lobb 24 (K). 

INDONESIA: Borneo I: Banguey Is: Boden Kloss 19234 

(US, BM). Java 1.: Prov. Tegal: Between Pogal and Gerbo, 

800-900 m, Mousset 15 (US, BISH); Tengger Mountains, Pogal-Gerbo, Moussct [Ros. Fil. Exs. 831 (US); 

Lebaksioe, Raciborski in 1897 (US, 2 sheets, type and isotype), in 1898 (US). Prov. Batavia: Buysman 124 (US); 

Dessa Dramaga near Buitenzorg, Schifner in 1893 (BISH). 

Celebes I.: Near Menado, Nielsen 754 (US, doubtful, very 

fragmentary). 

INDO-CHINA: Pierre (US, 4 sheets) ; (poor material, determination doubtful). 

CAMBODIA: Angkor, H. M. Smith 299 (US). 

THAILAND: Singora: H. M. Smith 190 (US). Chantabun: Vesterdal 477 (fragment, US). Gulf of Siam: Koh 

Tao Island, H. M. Smith 263 (US). Phitsanolok: Tung 

Salaeng Luang National Park, 500 m, Tugawa 11795 (US). 

CEYLON: Ferguson 137 (US). 

AFRICA : Republique Centre-Africaine: Oubangui-Chari, 

region de M’Baiki et Boukoko, Tisserant 1170 (P). Fernando Po I., Vogel (K) [possibly introduced]. 

Names based on Polypodium pteroides Retz. 

( 1791) have usually, until recently, been applied to 

this species, and correctly. Then Ching ( 1933 : 566- 

568) confused the application of the epithet pteroides 

of Retzius by saying he had examined the type of 

Polypodium pteroides Retz., a specimen from Madras 

collected by Konig, which he says “really represents a 

species of the group of Aspidium sagenioides Mett. 

(Dryopteris sagenioides 0. Ktze.) , now considered as 

a free-veined tectarid fern, nearest to, or perhaps 

identical with Aspidium fuscipes Wall. or Nephrodium membranifolium Presl, both common ferns in 

Tropical Asia.” This, however, ignores the statement 

in Retzius’ description that the sori are marginal, and 

the fact that Retzius cited a Konig collection from 

Ceylon, not the one from Madras. Therefore, except 

for the possibility of regarding Polypodium pteroides 

as a later homonym of P. pterioides Lam. ( 1778) , the 

epithet pteroides would have been available until 

1967, when Rolla Tryon transferred P. pteroideum 

Klotzsch to Thelypteris. Perhaps it would still be the 

correct epithet if the plant were regarded as belonging to another genus. However, it is unclear from Art. 

73 of the Code whether Polypodium pterioides Lam. 

and P. pteroides Retz. (and P. pteroideum Kl.) are 

or are not homonyms. If they had not been described 

in the same genus it might be possible to regard the 

epithets as different. However, in one genus they will 

certainly be confusing, and we choose to regard the 

three epithets as orthographic variants or at least as 

too close to be maintained under Art. 75. 

If the Cuming 48 duplicate in the U.S. National 

Herbarium may be taken as an isotype of Neplzrodium oreopteris Fee this name can be referred here 

with some hesitation. It seems rather intermediate 

between T. ulagneri and T. opulenta, having the 

abrupt terminal contraction of the frond of T. wagneri, the pinnae more deeply cut but rather nearer T. 

wagneri in the shape of the lobes, and the terminal 

segment somewhat more deeply lobed at base than is 

usual for this species. In any event, the use of this 

epithet in Thelypteris is precluded by the combination T. oreopteris (Ehrh.) Slosson ex Rydb. (1917). 

Mettenius based his Aspidium pteroides var. a on 

A. conioneuron Fee, his var. b partly on P. pteroides 

Retz. and partly on plants here referred to T. opulenta, and var. c on A. obtusatum Swartz, so it is just 

as well he did not name the varieties. 

Since the names applied to this species are either 

unavailable or incorrectly applied to it, it seems necessary to propose a new name. We take great pleasure 

in dedicating it to Prof. W. H. Wagner, Jr., outstanding and profound student of ferns and teacher of 

botany. To avoid confusion, we have proposed it as a 

new species, rather than as a renaming of one of the 

old ones, thus attaching it to an extant type of which 

we are certain. 

The boundaries between T. wagneri and T. opulenta are unclear in Thailand. The rhizomes of the 

plants cited as T. opulenta in Thailand tend to be 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

more slender and creeping than is usual in that species. The specimens cited below are variously intermediate in the shape of lobes and depth of lobing of 

the pinnae, the abruptness of contraction of the terminal portion of the frond, and the shape and degree 

of cutting of the terminal segment. 

THAILAND: Wang-Djao, near Koh-Yai, Hosseus 89 

(BM); Raheng, Nakawn Sawan, 150 m, E. Smith 3018 

(US) ; Tap Sai, Chantabun, 200 m, Kerr 9604 (US) ; 

Prov. Udon, H. M. Smith 424 (US); Gulf of Siam, Koh 

Tao Island, H. M. Smith 256 (US). Khun Tai Mts., H. M. 

Smith 473 (US; annotated by R. C. Ching as Cyclosorus 

interruptus var. hirsutus) ; Sriracha, near sea level, E. 

Smith 3011 (BM); Sriracha, Nong Kaw, Marcan 1218 

Thelypteris interrupta ( Willd.) Iwatsuki 

Thelypteris interrupta (Willd.) Iwatsuki, Jap. J. Bot. 38: 

314, 1963. 

Pteris interrupta Willd., Phytogr. 13, Tab. X, 1794. Type: 

India, Klein. 

Polypodium tottum Thunb., Prodr. F1. Cap. 172, 1800. 

Type: Cape, Thunberg. 

Aspidium obtusatum Swartz, Schrad. J. 1800(2) :33, 1802. 

Type: no specimen is mentioned, but probably the 

pinna in BM thus labeled by Swartz and discussed below 

is a fragment of what he described. 

Aspidium goggilodus Schkuhr, 24 K1. Krypt. Gew. 1:193, 

t. 33c, 1809 [spelled goggylodus on plate, usually corrected to gongylodes]. Type : Essequebo, from Gartner. 

Aspidium resiniferum Kaulf., Enum. Fil. 237, 1824. Type 

OWahu [Hawaiian Is.], Chamisso. 

Cyclosorus gongylodes (Schkuhr) Link, Hort. Berol. 2 : 128, 

1833. 

Dryopteris gongylodes (Schkuhr) 0. Ktze, Rev. Gen. P1. 

2:811, 1891. 

Thelypteris gongylodes (Schkuhr) Small, Ferns SE States 

248, 475, 1938. 

Thelypteris totta (Thunb.) Schelpe, J. So. Afr. Bot. 29:91, 

1963. 

Names based on Pteris interrupta Willd. have commonly been applied to one or the other of the two 

species described above, distinguished most readily 

from other similar Thelypteris (Cyclosorus) species 

by the tiny yellow particles or globules (glands?) on 

the lower sides of the lobes of the pinnae. However, 

while trying to separate and determine the correct 

names for these two plants, our suspicions were 

aroused by the similarity of Willdenow’s plate to the 

common pantropical marsh fern usually called Dryopteris (or Cyclosorus) gongylodes (Schkuhr) Kuntze 

or more recently Thelypteris totta (Thunb.) Schelpe. 

This possibility was dismissed as unlikely, since no one 

had ever suggested it in print, so far as we could determine, and the epithet interrupta was in common 

use for the other plants mentioned above. 

While looking atplants of this affinity in the British 

Museum in 1968, we encountered a sheet on which 

were mounted three separate pinnae from the Swartz 

herbarium. One of these is labeled, in Swartz’ hand 

“pinna ex Pteridis interrupta Willd. (ab ipso!)- 

mihi aspid. obtusatum.” Another is labeled, also by 

Swartz, “Aspidium obtusatum Sw. prodr.” Both of 

these are labeled in pencil, probably by Gepp, “D. 

gongylodes var. proliferum.” Very careful examination shows no globular yellow particles, the venation 

of the lobes is of the peculiar ascending type and the 

undersurface has the sessile glistening resinous glands 

characteristic of what has been called Dryopteris gongylodes. These two pinnae belong to the hirsute form 

that is common over most of the range of the species. 

It seems clear that the pinna from Willdenow must 

be a fragment of the specimen on which he based his 

Pteris interrupta, and which he sent to show Swartz 

what this species was. This specimen was, presumably, 

one collected by Klein in India. 

Dr. D. E. Meyer has very kindly sent a photograph 

of the Klein specimen in the Willdenow Herbarium 

in Berlin. This photo shows a plant with exactly the 

appearance of “Dryopteris gongylodes” except that 

the large pale scales sometimes found on the rachis do 

not show, unless some rather shadowy grayish 

blotches are the scales. The sheet bears an annotation, 

possibly in Willdenow’s hand “Pteris sp. n.? Aug. 

1794. Auf den Reise nach Madras. (Klein)” and 

another “Klein Ind. 1797 W,” as well as annotations 

by Ching and Hieronymus. This sheet and another 

without a label but probably a duplicate are both 

numbered 19770 in the Willdenow Herbarium. They 

both bear the annotation in the upper right corner 

“Asp. obtusatum” “la” and “lb,” presumably added 

by Willdenow when informed by Swartz of their identity with his species in the genus Aspidium. Interestingly enough, the sheet marked “la” has one pinna 

missing, probably that sent to Swartz. 

Dr. Meyer also kindly sent a photo of the type sheet 

of Aspidium goggylodus Schkuhr, from the Schkuhr 

Herbarium. This consists of the drawing reproduced 

as t. 33c of Schkuhr (1809) and a pocket containing 

a single pinna, with the note “Durch Gartn. aus 

Essequebo. Eine pinne habe an Mohr und Sw. ge- 
NUMBER 8 9 

sandt.” Although Dr. Meyer says that “Aspidium 

goggylodus Schkuhr und Willdenow No. 19770 sind 

nicht identisch,” they appear to belong to the same 

species. Schkuhr apparently later regarded his A. 

goggilodus as the same as A. obtusatum Swartz, since 

on a small ticket attached to the sheet bearing the 

drawing for his Tab. 33b he has crossed out “gongylod” and written “obtusat.” Why this is on the 

sheet of this plate, which is his Aspidium serra is not 

clear. 

In any event, there seems little doubt that the 

original material of Pteris interrupta Willd., Aspidium obtusatum Swartz, and Aspidium goggilodus 

Schkuhr belong to the same species, the common pantropical marsh fern. Willdenow’s epithet was earlier 

and must replace the several later ones that have been 

applied to it, e.g., totta, obtusata, and goggilodus 

(goggylodus, gongylodes). The necessary combination 

was made by Iwatsuki and later by Stone (1967: 

135), both under the impression that Willdenow’s 

Pteris interrupta was the widespread species now to 

be known as Thelypteris opulenta. 

For a pantropical species, Thelypteris interrupta 

shows remarkably little variability over most of its 

range. However, East African specimens exhibit considerable variation in depth of lobing of pinnae, scaliness, and the hairiness of the underside of the fronds. 

Material from Mafia Island is unusually hirsute; 

glabrous plants are found in Tanganyika, Zanzibar, 

and Pemba, as well as in South Africa, New Caledonia, and Tahiti. 

Other African Species with Yellow Glands 

The presence of yellow particles or glands on the 

underside of the fronds is apparently a rather uncommon feature in Thelypteris, so much so that their 

presence may cause people to refer quite unrelated 

plants to the species complex discussed above. Thelypteris microbasis (Baker) Tard. has abundant yellow 

glands, especially on the indusia. It is a plant with 

much more delicately cut fronds and does not seem 

especially related to any of the species treated in this 

paper. The only specimen we have examined is from 

Liberia, in the vicinity of Mount Coffee, Cook 22 

Specimens from Uganda, at first referred to Cyclosows quadrangularis, were, when found to have such 

yellow glands, tentatively placed by the collectors in 

(US) ’ 

Cyclosorus extensus. The glands on this material, 

however, are not identical with those of the T. opulenta-T. wagneri complex. Mr. Robert Faden and Dr. 

K. E. Lye, of the East African Herbarium and 

Makerere University, respectively, have kindly placed 

the available material of this species at our disposal 

for study. This has enabled us to distinguish it clearly 

from the species discussed above and, hopefully, to 

indicate its relationship. Since it does not seem to 

have been described it is proposed as a new species. 

Thelypteris fadenii Fosberg and Sachet, new species 

TYPE.-Uganda, Lye 51 86. 

Planta terrestris, rhizomatibus repentibus, frondibus hirsutis ovatis ad basim non contractis, pinnis 

lobatis, loborum costulis infra glandulis clavatis vel 

fusiformibus aureis instructis, indusiis valde hirsutis. 

Terrestrial plant with a creeping rhizome about 3 

mm thick, this shortly hirsute and very scaly with 

lance-linear caudate caducous brown scales; fronds 

up to 100 cm long, irregularly disposed along rhizome 

from congested to 2-3 cm apart, the stipe pale dull 

brown, shorter than, to rarely as long as, blade, the 

longest stipe seen 50 cm, the bases of stipes hirsute 

and scaly with large glossy brown caudate caducous 

scales, the scales at first abundant, on old fronds 

scarce; blades up to 48 cm long, narrowly ovate in 

outline, broadest at base or the basal pinnae slightly 

shorter and spreading to more or less deflexed, without 

reduced pinnae, narrowed apically to a caudate basally pinnatifid terminal segment, fertile almost to tip 

in large fronds; pinnae 16-18 well-developed pairs, 

subopposite to alternate, linear-lanceolate, acuminate, 

up to 13.5 cm long, up to 1.8 cm wide, lobed to 7 mm 

or even deeper, the lobes very slightly falcate, 

rounded to almost obliquely truncate at apex, hirsute 

on both surfaces, beset beneath, especially along 

costae, with abundant or rather sparse clavate or fusiform to rarely bulbous or globular golden brown to 

rarely pale yellowish glands, these glossy and translucent, appearing resinous; one pair of veins united 

below the sinus into a single vein ending in the sinus; 

sori about one-third in from margins of lobes but 

sometimes appearing submarginal because of the revolute lobe margins, rather closely set along the sides 

of the lobes, absent from the tips, sparsely scattered 

to abundant on united part of pinna, on most fronds 

reaching to near midrib, on some reduced to one or 
10 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

two pairs per lobe, these then near midrib; indusium 

well developed, orbicular-reniform, prominently long 

white hirsute. 

Found in Uganda, on forest floor in West and East 

Mengo districts ; one specimen, incomplete but probably this, from St. Helena Island. 

AFRICA: Uganda, West Mengo District, Kyadondo 

County, Kampala, Makerere University Hill, 1200 m, K. E. 

Lye 5186 (MHU, holotype, isotypes, US, EA, NY, UC, 

GH, BM, IS, P, 0, B, MO) ; Cooper 2 (MHU, sterile, 

yellow glands very pale); Mawakota County, Mpanga 

Forest Reserve and Research Station, 1150 m, Faden, 

Evans ti3 Lye 69/998A (EA), Lye et al. 3822 (MHU; 

yellow glands pale and not abundant). East Mengo District: Kyagwe County, Ssezzibwa Falls, 10 km W of 

Lugazi, 1200 m, Faden &, Evans 69/985 (EA, US). 

ST. HELENA I.: Quelle, Rosemary, lust in 1901 (US). 

The St. Helena specimen consists of only two 

blades and agrees in all respects, including the clavate 

golden glands, with T. fadenii, although the distribution seems unlikely, 

This species seems without doubt closest to Thelypteris quadrangularis (Fee) Schelpe (Dryopteris contigua Rosenst.), in fact, so close that it might be regarded as a variety of it. It differs in its creeping 

rather than erect rhizome, more scattered fronds, 

shorter and broader lobes of the pinnae, and especially in the clavate to fusiform golden glands. As 

species go in tropical Thelypteris, this seems sufficient 

to justify specific rank for T. fadenii. A considerable 

series of Brazilian T. quadrangularis, determined by 

Alan Smith, and a series from Africa, not so uniform 

but agreeing in most essential characters, were examined, especially for glands. Some of the Brazilian 

specimens, especially three from Vicosa, Minas Gerais 

(Mexia 4487-a, 4760, and 5498-a, all US) and one 

from Sao Leopoldo, Rio Grande do Sul (Leite 4282, 

US), have scattered to abundant tiny white particles 

on the undersurface of the lobe tips. A sheet from 

Sao Bento, Bahia (Luettelberg 271a, US), has such 

particles and a few tiny globose pale to brownish 

glands. These could be regarded as approaching T. 

fadenii, but the “glands” are much smaller and not 

the same shape, and the other characters are those of 

T. quadrangularis. 

These two species do not seem particularly related 

to the T. opulenta-T. wagneri group, nor to T. intarrupta (T. goggilodus, T. totta), but perhaps more so 

to T. dentata (Forsk.) E. St. John and T. parasitica 

(L.) Tard. 

Literature Cited 

Ching, R. C. 

1933. Notes on the Herbarium Willdenow. Lingnan 

Journal of Science, 12: 565-570. 

Ito, H. 

1937. Filices Japonenses. VI. Botanical Magazine 

(Tokyo), 51 :709-714. 

Schelpe, E. A. C. L. E. 

1965. A Review of the Southern African Species of 

Thelypteris. Journal of South African Botany, 

3 1 : 259-269. 

Schkuhr, C. 

1809. Vier und zwanzigste Klasse des Linneischen 

Pfianzensystems odes Kryptogamische Gewachse. 

1 : 1-212, t. 1-173, lb-56bJ Wittenberg. 

Enmeratio filicum Philippinarum. Hooker’s lournal of Botany, 3:392-422. 

Smith, J. 

1841. 

Stone, B. C. 

1967. Further Additions to the Flora of Guam, 111. 

Micronesica, 2: 133-141. 

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