Se Tie FERN GAZETTE is a journal of the British Pteridological Society and Ree ae ne +. a. aspects 0 pteridology. zt gas d, and books etc. sent for review, to: Prof. M. Gibby, aoe Royal Botanic Garden Edinburgh, 20A Inverleith _— Leagiim de EH3 5LR, UK ears 0131-248-2973 E-mail: FernGazette@eBPS.org.uk ions for authors are on page 283 of this volume and also available at es http://www.eBPS. cae Copyright © 2009 British Pteridological Society. All rights reserved. No part of this publication may be reproduced in any material form (including photocopying or storing it in any medium by electronic means) without the permission of the British Pteridological Society. THE FERN GAZETTE Volume 18 Part 4 was published on Ist December 2008 Published by eee eae eas sie AL SOCIETY Department The Natural History Cen cS met Rs em SW7 S5BD, UK Printed by Bishops Printers Limited Fitzherbert Road, Farlington, Portsmouth, PO6 1RU, UK www.bishops.co.uk Cover design by Hazel Sims MISSOURI BOTANICAL JUL 06 2009 A SUMMARY OF INDIAN CHEI AEARRERL [DBREARNS AND THE DISCOVERY OF NEGRIPT, RIDACEAE), AN AFRO-ARABIAN FERN GENUS NEW TO INDIA FERN GAZ. 18(5):216-229. 2009 216 C.R. FRASER-JENKINS! & C.S. DULAWAT? "Student Guest House, Thamel, P.O. Box no. 5555, Kathmandu, Nepal (Email: chrisophilus@yahoo.co.uk) “Bryology Laboratory, Dept. of Botany, University College of Science, M.L.S. University, Udaipur, Rajasthan - 313001, India (Email: csdulawat@rediffmail.com) Key words: Cheilanthes, Aleuritopteris, Notholaena, Negripteris, fern, Rajasthan, India ABSTRACT A summary of Indian cheilanthoid ferns treated under nine genera includes three new names, Notholaena dipinnata Fras.-Jenk., Cheilanthes bhutanica Fras.- Jenk. & Wangdi and Cheilanthes tibetica Fras.-Jenk. & Wangdi, and five new combinations, Cheilanthes nitidula Hook. subsp. henryi (Christ) Fras.-Jenk., Aleuritopteris bicolor (Roxb.) Fras.-Jen ulawat, Aleuritopteris subdimorpha (C.B.Clarke & Baker) Fras. Jai, and Notholaena muelleri (Hook.) Fras.-Jenk. Negripteris scioana (Chiov.) Pic.Serm. (Pteridaceae), a close relative of both A/euritopteris and Chrysochosma, was discovered by the second author in semi-arid conditions in the Kumbhalgarh and Sitamata Reserves of the Aravalli Hills in central Rajasthan, N.W. India, the first record for the Indian sub-continent. It was known previously only from N.E. Africa, Socotra and S. Arabia and is an Afro-Arabian species which, as now found, extends eastwards into the hills of the semi-arid region of W. India. INTRODUCTION The cheilanthoid ferns of India have at various times been placed in the families Sinopteridaceae, Negripteridaceae, Cheilanthaceae and Hemionitidaceae, but are now generally accepted as belonging to subfamily Cheilanthoideae, within Pteridaceae, with the other four families, among others, in its synonymy. The common term cheilanthoid is a vague and undefined one, though it should be recognisable to most fern-workers. The most obviously “cheilanthoid” ferns of India belong to nine genera, though various other genera are also allied or within Subfam. Cheilanthoideae, but are not dealt with here (see, for example, Tryon, Tryon & Kramer 1990). These nine genera with their Indian subcontinental species are as follows: 1. Notholaena R.Br., includes Cosentinia Tod., Paraceterach Copel. and Paragymnopteris K.H.Shing, perhaps also Chrysochosma (J.Sm.) Kiimmerle. This genus has long been accepted internationally as typified by N. marantae (L.) Desv., following Christensen (1905) and subsequently the carefully reasoned work of Pichi Sermolli (1981, 1989). An apparent lectotypification by Smith (1875) merely used the word “type” in the sense of “typical species”, or even “representative species” and in several other cases Smith cited species not included by the original author of a genus, or even cited more than one species after the word “type”. His misuse of the word 217 FERN GAZ. 18(5): 216-229. 2009 “type” is therefore not acceptable as effective lectotypification. Christensen’s proper lectotypification was also followed world-wide, including in N. America (Tryon 1956, 1964 et al.), but Tryon & Tryon (1980) subsequently reversed it in favour of his own local regional misapplication of the name Notholaena to the N. and C. American segregate-genus, Chrysochosma (J.Sm.) Kiimmerle, on the basis that Smith’s “lectotypification” was earlier, though that was then rejected by the International Committee for Nomenclature (Pichi Sermolli 1981), which Tryon did not mention again. He thus ignored the bulk of literature across four other continents of the Old World, which created considerable nomenclatural confusion, and was subsequently supported locally in the USA by Yatskievych & Smith (2003). The conservation of Notholaena with a conserved type, N. marantae, is therefore advisable to restore the status quo ante without any room for doubt and allow current Old World authors to continue using Notholaena in its usual sense. Neither the Australian genus Paraceterach, nor Shing’s genus, Paragymnopteris, are accepted as replacing Notholanea for the eight well marked Old World species. Notholaena, in its traditional and present sense, and Chrysochosma are very close, critical genera, very difficult to define separately, and maintained largely on molecular grounds of unknown and possibly minor taxonomic significance. Their separability at a rank as high as genus is somewhat doubtful, despite being accepted in the USA, but if maintained, Pichi Sermolli (1989) has already made the great majority of necessary combinations for the New World species within Chrysochosma. Notholaena species are frequently exindusiate (without a pseudoindusium) and usually markedly hairy/scaly on the surfaces and axes; they often have the sori spreading down the veins from the margin, as a derivative from the marginal condition in many other cheilanthoid ferns. The five Indian subcontinental species are: N. himalaica Fras.-Jenk. (1997) (syn.: Gymnopteris vestita (Hook.) Underw., shown to have been misplaced in that genus by Mickel (1974)). Simply pinnate, with slightly elongated-ovate, densely silky-hairy pinnae. Widespread throughout the W. and E. Indo-Himalayan region, except for the far west Himalaya; Tibet; and S.W. China. N. borealisinensis (Kitag.) Fras.-Jenk. (1997) (probable syn.: Gymnopteris bipinnata var. auriculata (Franch.) Ching). China; newly reported from the Indian subcontinent in Bhutan (Thimphu, Taba, CRFJ 31573, 11 Oct. 2005, TAIF). Simply pinnate; distinguishable from N. himalaica by its smaller, cordate-based pinnae, narrowing slightly more to their apices and thus hastate-sagitate in shape. Notholaena dipinnata Fras.-Jenk., nom. nov. for Gymnopteris bipinnata Christ, Not. Syst. (Lecomte) 1: 55 (1909), non N. bipinnata Liebm. The lower pinnae are again pinnate and the upper ones lobed. The relationship between this species and N. borealisinensis is unclear as some intermediate plants appear to connect them together, though not in the Indian subcontinent. They were therefore treated as two varieties by Ching, but it is not considered likely here that the two extremes could merely represent variation within a single species. The intermediates require further study. This species has not been found in India, as the record by Fraser-Jenkins (2008), from Arunachal Pradesh (Lohit. B. Krishna 48966, ASSAM) was in error for a scrumpled specimen of N. borealisinensis, but it is mentioned here to explain its pe from the previous species and because it occurs very near to the border in S.W. Chi N. marantae (L.) Desv. Sometimes stated to consist of three dienes in Europe and Macaronesia, subsp. marantae, subsp. subcordata (Cav.) Benl & Poelt and subsp. cupripaleacea (Benl) Rivas Mart. et al., but these merely represent minor variation FRASER-JENKINS & DULAWAT: INDIAN CHEILANTHOID FERNS 218 within the species and are not considered here to be appropriately ranked as subspecies, or needing recognition. Cape Verdes; Morocco; Algeria; Ethiopia; Macaronesia: across S. Europe; the Caucasus and Turkey; Cyprus; Lebanon; Syria; Iran; Yemen; from the west to east Indo-Himalaya; Tibet; and $.W. China. NV. marantae is diploid sexual, with n= 29, including from Uttarakhand, and a report by Khullar, Sharma & Verma (1988) of n = 58, from Jamunotri, Uttarakhand, is considered to have been in error (Fraser- Jenkins 1997: 184). N. lanuginosa (Desf.) Desv. ex Poir. (syn.: N. vellaea (Aiton) Desv., non R.Br.: Cosentinia vellaea (Aiton) Tod.). Pichi Sermolli (1985) treated this species as a separate genus, Cosentinia, which he revived from earlier Italian literature, on the basis of its more trilete spores, but this has seldom been accepted. Morocco; Tunisia: Libya: Macaronesia; across S. Europe; Turkey; Cyprus; Lebanon; Syria; Israel; Yemen; Sudan: Iran; Afghanistan; Pakistan; N.W. India (Himachal Pradesh). A typical Mediterranean European element in the W. Himalaya. There are two subspecies, the tetraploid subsp. lanuginosa, which occurs throughout the range, excluding Pakistan and India, and the very closely similar, and cryptic diploid subsp. bivalens Reichst. (Badré & Reichstein 1983), apparently confined to Macaronesia, Spain and the Indian subcontinent. A further species is the Chinese N. sargentii (Christ) Fras.-Jenk. (1997). Two additional Australian species, Notholaena muelleri (Hook.) Fras.-Jenk., comb. nov., basionym: Gymnogramma muelleri Hook., Sp. Fil. 5: 143, t. 295 (1864), and N. reynoldsii F.Muell., were formerly placed in the genus Paraceterach, which was treated as endemic to Australia, but N. marantae and four other species were placed within Paraceterach by Tryon (1986) in order to accommodate his misapplication of Notholaena. 2. Cheilanthes Sw., which includes Cheilosoria Trevis. and Mildella Trevis., normally consists of efarinose species in Asia, with generally narrow stipe-scales and small segments. The 11 species present in the Indian subcontinent are: C. pteridioides (Reichard) C.Chr. subsp. acrosticha (Balb.) O.Bolos (syn.: C. acrosticha (Balb.) Tod.,). W. Himalaya westwards to Mediterranean S.W. Asia and Europe, N. Africa and the Cape Verdes Islands. Subsp. pteridioides (syn.: C. fragrans (L.f.) Webb & Berth., non Sw; C. maderensis R.Lowe; Negripteris quezelii Tardieu) occurs from Mediterranean S.W. Asia, west across Mediterranean Europe to Macaronesia and across N. Africa, but is not present in the Indian subcontinent or near to it. The two subspecies are different cytotypes with a close reticulate relationship and are very similar to each other, differing only in the length of the indusium. They have thus been much confused in earlier literature, usually under the name C. fragrans, which is illegitimate. Following the rejection by Committee of Nardi & Reichstein’s (1986) proposal to reject Polypodium pteridioides Reichard, in order to preserve their nomenclature, the nomenclature of this critical complex is now stable at either the specific or subspecific rank. C. persica (Bory) Mett. ex Kuhn (syn.: C. szovitzii Fisch & C.A.Mey.). Far N.W. Himalaya, ?7Himachal Pradesh, W. Asia, E. Mediterranean Europe and N. Africa. The lamina bears many long, pale hairs beneath. C. tibetica Fras.-Jenk. & Wangdi, nom. nov., for Pellaea straminea Ching, Bull. Fan Mem. Inst. Biol., Bot., 2: 203, t. 17 (1931), non Cheilanthes straminea Brause [= Woodsia indusiosa Christ] (syn.: Mildella straminea (Ching) C.C.Hall & Lellinger). S. and S.E. Tibet (type) and Bhutan. Newly reported here from the Indian subcontinent in 219 FERN GAZ. 18(5): 216-229. 2009 W. Bhutan (Chele La Pass, Ha District, 77 Wangdi, CRFJ no. 31625, 12 Oct. 2005, THIM, TAIF, det. CRFJ). Specimens from S. Tibet by the N. Sikkim border and from Lhasa in Herb. Lloyd Botanic Garden, Darjeeling (!). Lamina pale-green, with long rather unlobed pinnules, a pale-brown stipe and rachis and hair-tipped, linear red-brown scales up the stipe; fronds deltate-lanceolate, bipinnate; indusia both short and long, arising at the curled over laminar edge rather than shortly within it as in other species formerly placed in Mildella. C. nitidula Hook. subsp. nitidula (syn.: Pellaea nitidula (Hook.) Baker; Mildella nitidula (Hook.) C.C.Hall & Lellinger), from the W. Himalaya only, reaching eastwards to W. Nepal. C. nitidula subsp. henryi (Christ) Fras.-Jenk., comb. nov., basionym: Pellaea henryi Christ, Bull. Herb. Boissier 7: 7 (1890) (syn.: Mildella henryi (Christ) C.C. Hall & Lellinger). N.E. India (Arunachal Pradesh), Bhutan (Thimphu, Taba, CRFJ 31496, 8 Oct. 2005, THIM, TAIF), S.E. Tibet, S. China (Yunnan (type), Szechuan, Kweichow, Kwangtung, Fukien), Taiwan, Vietnam. Reported from Namdapha, Arunachal Pradesh, by Singh & Panigrahi (2005) sub Aleuritopteris albomarginata (C.B.Clarke) Ching in error. Their drawing shows 4. albomarginata from some other source, that being a higher altitude species they could not have found in the lower reaches of Namdapha they visited. However their photograph is of C. nitidula subsp. henryi, whose presence at Namdapha has been verified by CRFJ, from specimens previously cited by Chauhan (1996), but misreported by them as the Afro-Arabian C. farinosa (Forssk.) Kaulf. Very close to subsp. nitidula and differing only in the rachis hairs being denser and more prominent and sometimes extending further around the sides of the rachis and the smaller fronds with slightly less lobed pinna and pinna-lobes. This subspecies is a vicariant of subsp. nitidula and replaces it in the eastern part of its range. Previous records and collections of C. nitidula from further east referred to the present subspecies, which merges into it morphologically. The interesting bicentric range of this species is a well known distribution-pattern for species that prefer a somewhat drier climate and thus occur scattered behind the Himalayan line in the otherwise wetter parts of the central and east Himalaya. C. opposita Kaulf. (syn.: C. mysurensis Wall. ex Hook., C. fragrans Sw.). Sri Lanka, S. India (N. to Orissa), S. China (Yunnan, Hainan, Kwangtung, Fukien), Taiwan, Myanmar and the Philippines. This beautiful species with narrow, upright and finely dissect, bright green leaves has been nomenclaturally confused until recently (see Fraser-Jenkins 1997), though its nomenclature was clarified by Alston (1936) and in detail by Fuchs (1961). Its range has sometimes been given as throughout China and other areas in error for the closely related, but less dissect C. chusana Hook. C. tenuifolia (Burm.f.) Sw. Sri Lanka, through most of the Indian subcontinent except the N.W., China, Taiwan, S.E. Asia, Australia, New Guinea and Oceania. Although reported as diploid apomict by Verma (1961) it is now known that this species is tetraploid sexual in India and Sri Lanka, but like apomictic species produces only 32 spores per sporangium, as do several other sexually reproducing Cheilanthes species. C. hancockii Baker. Newly reported from the Indian subcontinent, from Taba, Thimphu, Bhutan, CRFJ 31497, 8 Oct. 2005, THIM, TAIF; previous G0 from the E. Indo- Himalaya were in error for Aleuritopteris bicolor. S.W. China. ?N. Thailand (C. delicatula Tagawa & K.Iwats., which may otherwise be pe C. belangeri (Bory) C.Chr. (syn.: C. varians Hook.). N.E. India, from Nepal eastwards; Bangladesh; Myanmar; S.E. China; Thailand; and S.E. Asia to the Philippines. FRASER-JENKINS & DULAWAT: INDIAN CHEILANTHOID FERNS 220 Reported from S. India by Beddome (1864) in error for C. thwaitesii (below), which he included within his concept of this species. C. trichophylla Baker (syn.: Pellaea trichophylla (Baker) Ching; Cheilanthes undulata C.Hope & C.H.Wright). S.W. China. Reported tentatively by S.C. Verma in Mehra & Bir (1964), from the Kyangnosla pass, below Chhangu [Tsomgo] Lake, E. Sikkim (Verma, pers. comm. to CRFJ, 2004), but the specimen lost and unverified, though perhaps correctly recorded due to its distinct appearance, as recalled by Verma and illustrated in a drawing in his unpublished Ph.D. thesis of 1962 at Panjab University, Chandigarh. Distinctive zig-zag rachides and the axes and deeply tripinnatifid lamina densely covered in short yellowish-brown hairs. C. bhutanica Fras.-Jenk. & Wangdi, nom. nov., for Pellaea yunnanensis Ching, Act. Phytotax. Sinica 20(2): 235 (1982), non Cheilanthes yunnanensis Brause [= Aleuritopteris subvillosa (Hook.) Ching]. China (Yunnan (type), Szechuan), Bhutan. Newly discovered from the Indian subcontinent by CRFJ as a common species of rocks and walls around Thimphu and elsewhere in W. Bhutan (Serbithang, CRF 31487, 8 Oct. 2005, with 7. Wangdi; Taba, CRFJ 31570, 11 Oct. 2005; Paro, CRFJ 31596, 12 Oct. 2005, with R. Pradhan & T. Wangdi; Chuzzom, CRFJ 31676, 15 Oct. 2005, with T. Wangdi, THIM, TAIF). A small plant similar to a coarser-segmented and slightly less dissect C. nitidula with more deltate fronds and no hairs above the axes; sori and indusia continuous around the edges C. thwaitesii Mett. ex Kuhn (syn.: Cheilanthes laxa T.Moore ex Bedd., nom. superfl. for C. thwaitesii; Cheilanthes keralensis N.C. Nair & S.R. Ghosh, Aleuritopteris thwaitesii (Mett. ex Kuhn) Saiki). Sri Lanka (type) and South India; a Hindu-Lankan endemic. Type from Sri Lanka. This often overlooked species is important in illustrating that the presence or absence of farina is not confined to A/euritopteris, but also occurs in some true Cheilanthes species. 3. Aleuritopteris Fée, formerly treated by most authors as Cheilanthes subgen. Aleuritopteris (Fée) W.C.Shieh, and all the species, except A. stenochlamys Ching ex S.K.Wu, have names in Cheilanthes. But it was recognised generically by Ching in Ching & Wu (1981, 1983) and subsequent Chinese workers and Saiki (1984). It appears likely that it constitutes a distinct, if probably heterogeneous entity, which being easily recognisable is practicably and usefully recognisable as a genus. The species are usually well-marked and readily distinguishable by their stipe-scales (bicolorous or concolorous) and their distribution up the axes, though most were at one time referred to the African and Arabian species, 4. farinosa (Forssk.) Kaulf., which does not occur in Asia. The genus includes Sinopteris C.Chr. & Ching and Leptolepidium Ching & S.K.Wu (in the sense of its type species and also in the sense of Ching & Wu, which was based on a misapplication of the name Cheilanthes dalhousiae Hook.). It also has two sections, the first being the less dissect, more palmate species related to A. argentea (below) and the rest being the more pinnate species along the pattern of A. farinosa. Morphologically Aleuritopteris is slightly difficult to define in a way that distinguishes it from some species of Doryopteris, which are rather close to the A. argentea group, but are perhaps more likely to be related to the imparipinnate genus, Pe/laea. Most species have rather wide segments and a strong white or yellow farina, with wide stipe- scales. Due to considerable lack of clarity and confusion, the cytology and reported aneuploid base-numbers for various species need to be carefully reinvestigated with accurate identification and preservation of voucher-specimens. The 18 species present prs | FERN GAZ. 18(5): 216-229. 2009 in the Indian subcontinent have mostly been detailed by Fraser-Jenkins (1992, 1993 and 1997) and are: A. argentea (S.G.Gmel.) Fée (syn.: A. flava (Ching & S.K.Wu) S.R.Ghosh, non Saiki, nec sensu Ghosh [= A. subargentea]). Far N.E. India, a single collection of J.D. Hooker & T. Thomson’s from the Khasi Hills, Meghalaya (K!), might be correctly localised, but it has never been collected since in this generally well known area and is normally a higher-altitude, and usually higher latitude species. A similar case occurs with their collection at Kew of Lycopodium annotinum L. subsp. alpestre (Hartm.) A. Love & D. Léve, said to be from Khasia, but almost certainly from N. Sikkim. However 4. argentea (S.G.Gmel.) Fée, which otherwise occurs in W., S.W., C., N. and E. China; Tibet; Taiwan; Korea; Japan; and Siberia, has now been discovered in Bhutan (Taba, near Thimphu, CRFJ 31500, 8 Oct. 2005, THIM, TAIF), and was previously collected from N. Lohit, at the Tibetan border in or on the border of Arunachal Pradesh (F- Kingdon-Ward, BM!). A further species, close to 4. argentea, but without white farina and with small, narrow, less lobed segments has also turned up in N.E. Arunachal Pradesh (Changlang, Namdapha, Shirung to Hunung, c. 1100 m., B.K. Shukla 88207, 7 Feb. 1986, ASSAM, det. CRFJ). It appears to belong to the efarinose Chinese species, A. shensiensis Ching, though further comparative study may be required. A. subargentea Ching ex S.K.Wu. Similar to 4. argentea but slightly more dissect and the frond slightly more pinnately arranged, less palmate. Newly discovered in the Indian subcontinent (Marpha to Tukuche, Mustang, N.C. Nepal, CRFJ 30509, 27 June 2004, with G. Tamang, TAIF); Songgong, Sikkim, 1,400ft, Ribu & Rhomoo Lepcha, for GH. Cave 7634, 4 Oct. 1923 (Herb. Lloyd Botanic Garden, Darjeeling!, det. CRFJ). Tibet; S.W. China. A, tamburii (Hook.) Ching. N.C. Nepal; N. Sikkim; Meghalaya; Tibet and S.W. China. A very distinctive, large species of the 4. argentea group, with wide, coarse lobes and bright white farina. A, subvillosa (Hook.) Ching (syn.: A. tenella (Ching & S.K.Wu) Saiki). W. to E. Indo- Himalaya; Tibet; S.W. China. Efarinose, but closely related to the farinose A. caesia (Christ) Ching and A. kuhnii (Milde) Ching, from Tibet and S.W. or C. to N.E. China; Japan (the latter reported from India in error for 4. dalhousiae by Dixit (1984)). A. duthiei (Baker) Ching. W. Indo-Himalaya (very rare); N.W. Nepal; Bhutan. Efarinose, related to 4. /eptolepis (Fras.-Jenk.) Fras.-Jenk., but with a wider-triangular and more dissect lamina; both have wide, pale stipe-scales. A. leptolepis (Fras.-Jenk.) Fras.-Jenk. (2008) (misapplied name: A. dal/housieae (Hook.) Ching, nom. utrique rejic.). The totally efarinose, higher-altitude species long known as 4. dalhousieae [* bor ieger *| in error due to Hooker’s confusing the efarinose gps Nae rronds of A. albomarginata (below) with this species. W. to E. Indo- Himalaya; nai Tibet: S.W. China. Misreported sub “C. dalhousiae” sensu auct. Ind. by Dixit (1996) from Orissa, in error for 4. bicolor with the powder washed off by alcohol during herbarium-poisoning. A. rufa (D.Don) Ching. Common from the W. to E. Indo-Himalaya; also in S.W. China; Myanmar; Thailand. The fronds are lanceolate, narrowing slightly to the base, and the axes are densely covered with narrow, hair-like, fibrillose scales. 4. dubia (C.Hope) Ching (syn.: A. subrufa (Baker) Ching; C. /eveillei Christ; A. humatoides Saiki; C. wusukungii Miyamoto & Ohba). Intermediate between the last and next species and sharing the same phytochemistry of its white flavonoid powder (Fraser-Jenkins & Wollenweber in prep.). W. to E. Indo-Himalaya; N. Western Ghats; FRASER-JENKINS & DULAWAT: INDIAN CHEILANTHOID FERNS Faw. Myanmar; Tibet; S.W. and S. China; Taiwan; Thailand; the Philippines. Confused in China and reported as the next species. A. albomarginata (C.B.Clarke) Panigrahi (syn.: A. dalhousieae (Hook.) Ching, nom. utrique rejic., non sensu auct. Ind.). Widespread at somewhat higher altitudes from the far W. to N.E. Himalaya; Orissa; N. Western Ghats; Tibet; S.W. China; Taiwan; Thailand; Vietnam. The fronds are deltate and instead of hair-like scales have bicolorous scales extending up the stipe, rachis and costae. The Summer (monsoon) fronds are taller and more developed and almost or quite without farina, unlike the smaller, basal Winter fronds. A. chrysophylla (Hook.) Ching (syn.: A. humatifolia X.C.Zhang & L.Shi; A. flavopygmaea $.R.Ghosh). The only species with bright sulphur-yellow farina beneath in the Indian subcontinent. The W. Indo-Himalaya (Simla); C. Nepal to N.E. India: Myanmar; Tibet; S.W. China; Thailand. A. formosana (Hayata) Tagawa (syn.: C. brevifrons (Khullar) Khullar). One of the most widespread species, frequently reported from China under the name 4. anceps in error. W. Africa (Guinea); far W. to the E. Indo-Himalaya; Rajasthan (C.S. Dulawat, det. CRFJ); Uttar Pradesh; Bihar (Parasnath); Orissa; N.W. Ghats; Arunachal Pradesh; Tibet; S.W. and S. China; Taiwan; Myanmar; Thailand; the Philippines. It is fairly close to A. anceps, but has a narrower and usually smaller frond, which is characteristically bullulate-wrinkled above, and the slightly narrower, bicolorous stipe-scales extend up the rachis as well, but not (except rarely an odd one) onto the costae. The rachis usually bears scattered glands. A. anceps (Blanf.) Panigrahi (syn.: A. pseudofarinosa Ching & S.K.Wu; A. interrupta Saiki; ?4. javanensis Saiki, non C. javanensis (Willd.) T.Moore). Widespread at lower altitudes in India etc., far W. to N.E. Himalaya; Rajasthan; N.W. Ghats; C. India: S. India; Sri Lanka; Tibet; S.W. China (rare); Taiwan; Myanmar; Thailand; ?Java; ?Timor. The lamina is deltate-lanceolate, but not as triangular and wide-based as in C. bicolor (below), and has a brighter white farina; the slightly wide, bicolorous stipe-scales do not normally extend above the top of the stipe. A. dealbata Fée (syn.: C. dealbata D.Don, non Pursh; 4. doniana S.K.Wu ex Ching, nom. superfl.; C. doniana Fras.-Jenk. & Khullar; A. sikkimensis S.R.Ghosh). E. part of the W. Himalaya (Uttarakhand); Nepal to N.E. India; Tibet; S.W. China; Myanmar; ??Thailand. This species is closely related to C. anceps, but has a longer, often very large frond and wider, less lobed segments, with narrower indusia, and a very bright white farina. The stipe-scales, though vaguely bicolorous, do not have such an obvious dark central stripe and tend to have a yellower basal region and slightly darker apical region. It was illustrated from Nagarjun, Kathmandu, Nepal, on the front dust-jacket by Fraser-Jenkins (1997). A. bullosa (Kunze) Ching (syn.: ?4. indica Fée; A. flaccida (Bedd.) B.K.Nayar & S.Kaur; C. flaccida (Bedd.) Mehra & Bir, non sensu Mehra & Bir [= A. bicolor]). Sri Lanka and S. India. A very large species with long fronds, a bullulate upper surface, well lobed pinnules, thick stipes and concolorous red stipe-base scales. A small-sized, more creamy-yellowish powdered segregate of A. deal/bata from S. India (especially the Shevaroy Hills) is 4. wollenweberi Fras.-Jenk. (2008), superficially similar to the Chinese, Taiwan and Japanese A. krameri (Franch. & Sav.) Ching. A. stenochlamys Ching ex S.K.Wu. E. part of the W. Himalaya (Uttarakhand); Nepal; Bhutan; Manipur; Tibet, S.W. China. This rare, high-altitude taxon is closely related to A. grisea and is perhaps of slightly doubtful status, requiring further study, though ano FERN GAZ. 18(5): 216-229. 2009 possibly a good species. Fraser-Jenkins previously confused its type and used the name C. stenochlamys for it in error. A. grisea (Blanf.) Panigrahi, non sensu Panigrahi (syn.: A. platychlamys Ching; C. platychlamys (Ching) Fras.-Jenk.). A high-altitude Himalayan species with concolorous red stipe-scales; widely misrecorded from C. or S. India and Orissa by Panigrahi (1965) and Dixit (1996) in error for A. formosana among other species. Far W. to E. Indo- Himalaya; Tibet, S.W. China; Taiwan. A, bicolor (Roxb.) Fras.-Jenk., comb. nov., basionym: Pteris bicolor Roxb. in Griff., Calcutta J. Nat. Hist. 4: 507 (1844) (syn.: Cheilanthes farinosa var. tenera C.B.Clarke & Baker; C. bicolor (Roxb.) Griff. ex Fras.-Jenk.; A. bicolor (Roxb.) Punetha & Kholia, comb. inval., sin. basionym; A. kathmanduensis Ching & S.K.Wu; misapplied name: Cheilanthes farinosa sensu auct. Ind. plur., Blanf. et al., non (Forssk.) Kaulf.). A very common and widespread, rather low-altitude species in India. W. Africa (Nigeria, Jos); far W. to N.E. Indo-Himalaya; Rajasthan; Bihar (Parasnath); C. India; Orissa; S. India; Bangladesh; Myanmar; ?Thailand; Laos. A similar taxon, but more delicate and with russet, concolorous stipe-base scales (like those of A. subdimorpha) occurs in Sumatra. There is some apparent transition, or some intermediate taxon, between the otherwise highly distinct 4. bicolor and A. anceps (and also somewhat towards A. subdimorpha) in S.E. Bangladesh (Chittagong Hills), S. India, Myanmar, S.E. China and ?New Guinea (Papua). But it is not yet clear why this should appear to be so. A, subdimorpha (C.B.Clarke & Baker) Fras.-Jenk., comb. nov., basionym.: Cheilanthes farinosa (Forssk.) Kaulf. var. suwbdimorpha C.B.Clarke & Baker, J. Linn. Soc., Lond. 24: 411 (1888) (syn.: Cheilanthes subdimorpha (C.B.Clarke & Baker) Hieron.; A. Jongipes Ching & S.K.Wu, nom. inval., sin. typ. (which is not A. bicolor, as tentatively thought by Fraser-Jenkins 1997, in the absence of the type); C. /ongipes (Ching & S.K.Wu) Dixit & Bal Krishna, comb. inval.; A. pentagona Saiki; pseudoargentea S.K.Wu). From C. Nepal eastwards to N.E. India; Arunachal Pradesh; Manipur; abundant in Meghalaya; ?Bangladesh; Myanmar; Thailand; S.W. to S.E. China; Vietnam. This rather little known species is generally similar to C. bicolor, but has a longer, thicker stipe, shorter and more coarsely lobed lamina and concolorous, russet stipe-scales. 4. Negripteris Pic.Serm. is closely similar to Aleuritopteris but differs in having only rather few (1-4) sporangia per sporangium (also shown by some Aleuritopteris), which are rather deeply embedded in the laminar powder (“subsessile”) and an apparently primitive, large sporangium, without a definite stomium area and a very broad annulus with all its walls thickened (indurated), including the outer one, appearing like a solid, dark cap to the sporangium. During dehiscence, the sporangium splits laterally and the whole annulus or top half of the sporangium falls off to release the spores. This may be a retained adaptive feature connected with a very dry climate, allowing protection of the developing sporangia against desiccation. However, on the strength of this hypothetically being supposed to be a very primitive characteristic Pichi Sermolli not only raised a new genus, but even a new family for the single specimen he had seen (which was undoubtedly and obviously mistaken), despite pointing out its evident similarity to A/euritopteris. In all its other features it is not separable from Aleuritopteris in its concolorous, lanceolate, pale-reddish stipe-scales, frond-shape, white farina beneath and shallow, interrupted pseudo-indusia formed by the leaf- margin. Weatherby (1948) pointed out tendencies within what is now Chrysochosma FRASER-JENKINS & DULAWAT: INDIAN CHEILANTHOID FERNS 224 from N. America, towards a similar annulus and embedded sporangium, and pointed out the likeness of N. scioana to some “Notholaena” species (i.e. Chrysochosma spp.). While not in any way invalidating the genera A/euritopteris and Notholaena, it must be said that a few species in either genus can hardly be distinguished morphologically from the other genus. The likelihood is therefore that Negripteris is either an Aleuritopteris or a Chrysochosma, or may perhaps be nearer to an ancient ancestor of both. However it is maintained here as a somewhat dubious genus, resting on its peculiar sporangia, pending further study. 4. rosulata (C.Chr.) Ching (syn.: 4. pygmaea Ching), from Tibet and S.W. China, has similar scales and lamina (with the lobes rather broadly joined at their bases and fusing at the apex as in Negripteris) and appears to be very close to it, but without the characteristic sporangia. A rare S.W. Chinese species, A. sichouensis Ching & S.K.Wu, is again similar and by definition only would most probably be placed in Chrysochosma. N. scioana (Chiov.) Pic.Serm. (syn.: Mohria scioana Chiov.; Negripteris tricholepifera Pic.Serm.). An Afro-Arabian species, formerly known from N.E. Africa and South Arabia only; Ethiopia, Sudan, Somalia, Socotra (also seen there by the first author in 1967), N. Kenya, Yemen, Saudi Arabia. To this must now be added a remarkable discovery of it from W. India in the semi-arid hills of Rajasthan. Last year, Dr. C.S. Dulawat (the second author) sent the first author material of his Rajasthan Aleuritopteris for identification and most surprisingly there was some fine, unidentified material of typical Negripteris scioana (Chiov.) Pic.Serm. among them that he had collected from the Kumbhalgarh and Sitamata Sanctuaries, Aravalli Hills and Chittorgarh, Rajasthan. This discovery is of considerable phytogeographical interest as it was otherwise only known to be in N-E. Africa and S. Arabia and fits in with a small handful of other Afro-Arabian connections in Western India, mainly in Rajasthan. The photograph purported to be Cheilanthes albomarginata C.B.Clarke sensu lato published by Chaudhary & Khichi (2007) is actually of the tip of the frond of Negripteris from Sitamata. Specimens collected are: 1. Rajasthan, Sitamata Wild Life Sanctuary, Ambaretti, crevices of rocks, 500-800 m. C.S. Dulawat CSD/SM/05-147, 14 Oct. 2005, Herb. Botany Dept., M.L.S. University, Udaipur. 2. Rajasthan, Sitamata Wild Life Sanctuary, Ambaretti, crevices of rocks and bank of nalah [stream], c. 500-800m. C.S. Dulawat CSD/SM/06-197, 5 Oct. 2006, Herb. Botany Dept., M.L.S. University, Udaipur. 3. Rajasthan, Khumbhalgarh Wild Life Sanctuary, Ranakpur Ghat, crevices of rocks, 550-800 m. C.S. Dulawat & B.L. Chaudhary CSD/SM/05-183, 23 Oct. 2005, BSD. 5. Doryopteris J.Sm. Characterised by the long, uninterrupted sori and often palmate fronds, but though widely and usefully accepted, is difficult to define exclusively, especially in relation to the Aleuritopteris argentea group, which also contains some efarinose species. Two species occur in the Indian subcontinent: D. concolor (Langsd. & Fisch.) Kuhn (syn.: Doryopteris geraniifolia (Raddi) Klotzsch). S. and C. America; Africa; Sri Lanka; S. India (N. to Orissa); S. China; throughout S.E. Asia; Australia; Oceania. D. kirkii (Hook.) Alston from Africa has occasionally been reported from India in error for D. concolor, but is anyway a somewhat doubtful taxon. The taxonomic position of this fern has long been in some doubt as it is very similar to an efarinose member of the A/euritopteris argentea group 225 FERN GAZ. 18(5): 216-229. 2009 and has often been placed in Cheilanthes. Pending further study, including molecular DNA work, it is maintained here in Doryvopteris, where it appears most likely to belong and has more usually been placed. D. ludens (Wall. ex Hook.) J.Sm. A distinctive species with tall black stipes and black midribs and the horizontal frond varying from cordate to considerably and deeply palmate-pinnately lobed into long, narrow segments, the lowest lobe basiscopically lobed again and the fertile fronds taller and more narrowly lobed than the sterile. It occurs in Orissa (Dixit 1996) and from N.E. India eastwards; Assam; Manipur: Nagaland; Tripura; Mizoram; Bangladesh; Myanmar; S.W. and S. China; Thailand; and S.E. Asia. Hope (1901) reported it from Chitral, N.W. Pakistan, in error on the basis of a specimen given to him by General Gatacre, ostensibly collected during the Chitral Relief Military Expedition, at Kaffir Rock on the road S. of Ziarat, along with Lygodium microphyllum (Link) R.Br. The specimens of both species, along with drawings and a sketch-map provided by Gatacre are in DD (!) and are correct. But anyone who knows the Pakistan fern-flora (Fraser-Jenkins 1992, 1993) and who has been there, where Kaffir Rock still exists beside the main road into Chitral from the Lowarai Pass, would know that neither species could possibly occur there or anywhere within thousands of miles. The specimens were almost certainly collected during Gatacre’s duty in Myanmar shortly previous to the Chitral expedition and seem most likely to have been more in the way of a “military prank”, than due to confusion. It is surprising that such an experienced expert as Hope could possibly have swallowed it! 6. Pellaea Link, nom. cons. Although various species, including those subsequently placed in Mildella by Hall & Lellinger (1967) and now in Cheilanthes, have been put into Pellaea in the past because of their glabrous, efarinose segments with long sori, the genus is now confined to those species with imparipinnate fronds, the apical segment being similar to a lateral one. The species may be glabrous or hirsute. Ghosh (1985) claimed to have “done” the taxonomy of Indian Pel/aea and listed 3 species, but the 5 species occurring in India are as follows: P. falcata (R.Br.) Fée (syn.: P. seticaulis (Hook.) S.R.Ghosh). This species has simple, unlobed, elongated, nearly glabrous pinnae with a only few hair-like scales beneath, and a hairy and scaly stipe and rachis, similar in both Australasia and S. India. It is absent from Malaysia (given by Ghosh), Hooker’s type-locality of “Penang” being in error for Lady Dalhousie’s collection from Sri Lanka (from where it was reported by Sledge 1982), which was omitted by Ghosh (1985). P. longipilosa Bonap. (syn.: P. malabarica B.K.Nayar & Geev.). Similar to P. falcata, but with narrow, tripartite lower and mid pinnae. Africa; S. India; confined to a few localities in Kerala. Though redescribed as if a new, endemic species, Fraser-Jenkins has reidentified it as being the African species, P. /ongipilosa, which is thus one of a rather small group of tropical African elements in S. India. P. boivinii Hook. A rather small, bipinnate species with from 2 to 3 pairs of elongated ovate, articulated-stalked pinnules on the lower pinnae and all the axes densely covered in very short, dark, blackish-brown hairs. S. and E. Africa; Madagascar; Mascarenes; Sri Lanka (Sledge 1982, omitted by Ghosh 1985); and S. India. Another tropical African element in S. India. P. viridis (Forssk.) Prantl. Another bipinnate species, with slightly larger, thinner and non-articulate-stalked, often biauriculate-based segments and glabrous axes. Adventive in S. India and Sri Lanka (Sledge 1982). Also naturalised in Australia and Oceania. S. FRASER-JENKINS & DULAWAT: INDIAN CHEILANTHOID FERNS 226 and E. Africa; Madagascar; and Yemen (type). P. calomelanos (Sw.) Link non Pityrogramma calomelanos (L.) Link. (syn.: P. hastata (Thunb.) Prantl, non (L.) Thunb.). A rare and very seldom-collected species occurring in a few scattered localities in very dry rocky areas. The stalked leaf-segments are cordate-hastate and without scales, borne on glossy black costae. Africa; Pakistan (Swat; Hazara); Himachal Pradesh; Uttarakhand; W. Nepal: S.W. China. An Afro- Arabian element reaching the W. Himalaya. 7. Pityrogramma Link. The “silver ferns” and “gold ferns” of cultivation where they often spread by self-sporing. They are exindusiate and the sori spread along the veins and cover the whole surface beneath the segments. Four New World species are adventive in the Indian subcontinent: P. calomelanos (L.) Link. The farina beneath the leaf, which is usually rather thin and weak, is white. S. and C. America. Adventive throughout the world in warmer climates and so abundant in even remote places throughout nearly all of the Indian subcontinent that it is sometimes difficult to remember that it is in fact an alien species not recorded before the later 19th Century. P. austroamericana Domin (syn.: Pitvrogramma calomelanos var. austroamericana (Domin) Farw.; P. calomelanos (L.) Kaulf. var. aureoflava (Hook.) Weath. ex F.M.Bailey; misapplied name: P. chrysophyila (Sw.) Link). The farina beneath the leaf is sulphur yellow. S. and C. America. A common adventive in Sri Lanka and S. India. In addition to the bright yellow powder, which is often lost in old specimens by treatment with alcohol (containing pesticides), it also has a shorter lamina with rather shorter and often less lobed segments. For some reason it has not reached N. India to date. It was first recognised as a separate species by Domin (1928, 1929) and detailed further in his later publications. The genus was monographed by Tryon (1962), and information was thence extracted by Panigrahi (1975) and from the determinations at Kew. P. dealbata (C.Presl) R.Tryon. White farina, a small, thin lamina with well lobed and toothed segments. C. America. Adventive in Sri Lanka, but probably not well established. P. sulphurea (Sw.) Maxon. Pale lemon-yellow, rarely white farina and the frond narrowed towards the base, with the ultimate segments cuneate-based and fanned out or flabellate at their strongly toothed apices. C. America. Adventive in Samoa and Sri Lanka, but probably not well established in the latter. 8. Parahemionitis Panigrahi. Following Mickel’s (1974) establishing that the Indian species is not a true Hemionitis, and Tryon, Tryon & Kramer’s (1990) clear statement that “H. arifolia” is not readily included in any genus, Panigrahi (1993) took it upon himself to utilise that information and make several attempts to interject a new generic name of his own for it (see Fraser-Jenkins 1997: 187-188), finally succeeding in validating one. Parahemionitis is therefore accepted here as the name for the genus and its single species-complex. P. cordifolia (Roxb.) Fras.-Jenk. (misapplied name: Hemionitis arifolia (Burm.f.) T.Moore). Distinctive cordate-rounded leaves. N.E. India (Bihar; Assam: Manipur; Nagaland; Tripura; Mizoram); Sri Lanka; S. India; Bangladesh; Myanmar: S. China; Vietnam; Malesia; Philippines. The widespread Indian triploid cytotype may often be proliferous with small plantlets developing in the axil of the leaf during the wet season, Bet FERN GAZ. 18(5): 216-229. 2009 but dropping off during the dry season. A Chinese diploid, which is more delicate, is never proliferous (Huang, Manickam & Chiou 2007). 9. Hemionitis L. (syn.: Gymnopteris Bernh.; Gymnogramme Desv.). The detailed work carried out by Mickel and his colleagues (Mickel 1974, 1988, Gianassi & Mickel 1979, Ranker 1989) on this genus has been usefully summarised in Pichi Sermolli & Bizzarri’s (2005: 66-68) final magnum opus. One species in the Indian subcontinent: H. tomentosa (Lam.) Raddi (syn.: oo tomentosa (Lam.) Underw.) S. America. Adventive and very common in Sri Lanka CONCLUSION The discovery of the African Negripteris in India is not only of considerable phytogeographical interest, but also suggests that further collection and study by Indian pteridologists with a specialist knowledge, able to recognise all the species they see, may reveal a number of other species hitherto overlooked in the subcontinent. The discovery of Notholaena borealisinensis, Cheilanthes hancockii, C. bhutanica, C. tibetica and Aleuritopteris subargentea, all previously unknown in the Indian subcontinent are further examples. ACKNOWLEDGEMENTS The author thanks Dr. Tandin Wangdi of the National Biodiversity Conservation Centre, Serbithang, Thimphu, Bhutan, who, along with Dr. Rebecca Pradhan, kindly accompanied him during collection excursions in W. Bhutan in October 2005 and is a joint authority of some of the resulting new taxa, above. REFERENCES ALSTON, A.H.G. 1936. Fern Notes -1. J. Bot. 74: 172-174. BADRE, F. & REICHSTEIN, T. 1983. The two cytotypes of Notholaena lanuginosa (Sinopteridaceae, Pteridophyta). Willdenowia 13: 361-367. BEDDOME, R.H. 1864. The ferns of Southern India and Ceylon: 65, ¢. 189. Gantz Bros., Madras. CHAUDHARY, B.L. & KHICHI, Y.S. 2007 [2006”]. Ferns of Kumbhalgarh Wild Life Sanctuary in Rajasthan, India. Indian Fern J. 23(1-2): 83-91. CHAUHAN, A:S. (1996) Pteridophytes, in HAJRA, P.K. (ed.) A Contribution to the Flora of Namdapha, Arunachal Pradesh: 67-98. Botanical Survey of India, Calcutta. CHRISTENSEN, C. 1905. Index Filicum: [i-] xli [-lix, 1-744]. H. Hagerup, Hafnia [Kobenhavn]. DIXIT, R.D. 1984. A census of the Indian Pteridophytes: 1-177. Botanical Survey of India, Howrah, Calcutta. DIXIT, R.D. 1996. Pteridophytes, in Saxena, H.O. & Brahmam, M. (eds.) The Flora of Orissa 4: 2536-2654. Orissa Forest Development Corp. Ltd., Bubaneshwar. DOMIN, K. 1928. Generis Pityrogramma (Link) species ac sectiones in clavem analyticam re or Spisy Vyd. Parod. Fak. Karlovy Univ. (Publ. Fac. Sci. Charles Univ.) 88: 1-10. DOMIN, K. 1929. ie ferns from Tropical America and the West Indies. Kew Bull. Misc. Inf. 1929: 215-222. F RASER-J ENKINS, C.R. 1992 [“1991”]. The ferns and allies of the far west Himalaya. Pakistan Syst. 5(1-2): 85-120. FRASER-JENKINS & DULAWAT: INDIAN CHEILANTHOID FERNS 228 FRASER-JENKINS, C.R. 1993 [1992]. The ferns and allies of es far west Himalaya - Some Additions and Corrections. Bot. Helvetica 102(2): 143-157. FRASER-JENKINS, C.R. 1997. New species syndrome in Indian Pteridology and the ferns of Nepal: 1-403 and errata. International Book Distributors, Dehra Dun. FRASER-JENKINS, C.R. 2008. Taxonomic Revision of Three Hundred Indian Subcontinental Pteridophytes With a Revised Census-List, a new picture of fern-taxonomy and nomenclature in the Indian subcontinent. 1-679. Bishen Singh Mahendra Pal Singh, Dehra Dun. FUCHS, H.P. 1961. The genus Cheilanthes Swartz and its European species. Brit Fern Gaz. 9(2): 38-48. GIANASSI, D.E. & MICKEL, J.T. 1979. Systematic implications of flavonoid pigments in the fern genus Hemionitis (Adiantaceae). Brittonia 31(3): 405-412. HALL, C.C. & LELLINGER, D.B. 1967. A revision of the fern genus Mildella. Amer. Fern J. 57(3): 113-134. HOPE, C.W.W. 1901. The ferns of north-western India e¢c., J. Bombay Nat. Hist. Soc. 13(3): 456-458 HUANG, Y.-M., MANICKAM, V.S. & CHIOU, W.L. 2007. Cryptic morphology and reproductive biology of two cytotypes of Hemionitis arifolia L. (Pteridaceae, Pteridophyta). Abstract in AMOROSO, V.B. (ed.) 4th Symposium on Asian Pteridology and Garden Show, 13-17 November 2007: 78. Central Mindanao University, Musuan, Bukidnon, Philippines. KHULLAR, S.P., SHARMA, S.S. & VERMA, S.C. 1988. Pteridophyte report, in BIR, S.S. (ed.) SOCGI Plant Chromosome Number reports 6. J. Cytol. Genet. 23: 38-52. MICKEL, J.T. 1974. A Redefinition of the Genus Hemionitis. Amer. Fern J. 64(1): 3- Lz. MICKEL, J.T. 1988 [1987”]. A new fern from Western Mexico and its bearing on the taxonomy of the Cheilanthoid ferns. Amer. Fern J. 77(4): 109-114. NARDI, E. & REICHSTEIN, T. (1986) Proposal to reject Polypodium pteridioides Reichard and all combinations based on it (Sinopteridaceae: Pteridophyta), Taxon PANIGRAHI, G. 1975. The genus Pityrogramma (Hemionitidaceae) in Asia. Kew Bull. 30: 657-667. PANIGRAHI, G. 1993 [“1992”]. Parahemionitis, a New Genus of Pteridaceae. Amer. Fern J. 83(3): 90-92. PICHI-SERMOLLI, R.[E.G.] 1946. Negripteridaceae e Negripteris, Nuova famiglia e nuovo genere delle filicales. — Giorn. Bot. Ital., n.s., 53(1-2): 129-169, 4. 14-16. PICHI-SERMOLLI, R.E.[G] 1950. The Geographical Distribution of Negripteris scioana. Amer. Fern J. 40(1): 59-69. PICHI SERMOLLI, R.E.G. 1981. Report of the Committee for Pteridophyta. Taxon 30(1): 161-163. PICHI SERMOLLI, R.E.G. 1983. Fragmenta Pteridologiae 8. Webbia 37(1): 111-140. PICHI SERMOLLI, R.E.G. 1985. The fern genus Cosentinia Todaro. Webbia 39(1): 179-189. PICHI SERMOLLI, R.E.G. 1989. Again on the typification of the generic name Nothlaena R.Brown. Webbia 43(2): 301-310. PICHI SERMOLLI, R.E.G. & BIZZARRI, M.P. 2005. A revision of Raddi’s pteridological collection from Brazil (1817-1818). Webbia 60(1): 1-393. RANKER, T.A. 1989. Spore morphology and genetic delimitation of New World Pas FERN GAZ. 18(5): 216-229. 2009 Hemionitis, Gymnopteris and Bommeria (Adiantaceae). Amer. J. Bot. 76(2): 297- 306. SINGH, S. & PANIGRAHI, G. 2005. Ferns and fern-Allies of Arunachal Pradesh 1: 1- 426; 2: 427-881. Bishen Singh Mahendra Pal Singh, Dehra Dun. SLEDGE, W.A. 1982. An annotated check-list of the Pteridophyta of Ceylon. Bot. J. Linn. Soc. 84: 1-30. SMITH, J. 1875. Historia Filicum: 1-249. London. TRYON, R.M. 1956. A revision of the American species of Notholaena. Contrib. Gray Herb. 179: 1-106. TRYON, R.M. 1962. Taxonomic Fern Notes 2. Pityrogramma (including Trismeria) and Anogramma. Contrib. Gray Herb. 189: 52-76. TRYON, R.M. 1964. Ferns of Peru - Polypodiaceae (Dennstaedtiaceae to ae Contrib. Gray Herb. 194: 1-253. TRYON, R.M. 1986. Some new names and combinations in [the] Pteridaceae. Amer. Fern J. 76: 184-186. TRYON, R.M. & TRYON, A.F. 1980. Proposal to conserve Notholaena with the type Pteris trichomannoides, Taxon 29: 160-161. TRYON, R.M., TRYON, A.F. & KRAMER, K.U. 1990. Pteridaceae in Kramer, K.U. & Green, P.S. (eds.) The Families and Genera of Vascular Plants (ed. KUBITZKI, K.) 1, Pteridophytes and Gymnosperms: [1-] 230-256 [-404]. Springer Verlag, Berlin, Heidelberg. VERMA, S.C. 1961. Cheilanthes, in Mehra, P.N., Chromosome numbers in Himalayan ferns. Res. Bull. Panjab. Univ., n.s., 12: 134-169. WEATHERBY, C.A. 1948. A Proposed New Genus and Family of Ferns. Amer. Fern J. 38(2): 58-61. YATSKIEVYCH, G. & SMITH, A.R. 2003. Typification of Notholaena R. Br. (Pteridaceae). Taxon 52(2): 331-336. FERN GAZ. 18(5):230-263. 2009 230 CHECKLIST OF THE FERNS AND LYCOPHYTES OF ACRE STATE, BRAZIL J. PRADO! & R. C. MORAN2Z Mnstituto de Botanica, Secdo de Curadoria do Herbario, C.P. 3005, CEP 01061-970, Sao Paulo, SP, Brazil E -mail: jprado.01@uol.com.br 2The New York Botanical Garden, Bronx, NY 10456-5126, U.S.A. Key words: checklist, Brazil, Acre State, Amazonian forest, ferns, lycophytes One hundred and seventy eight species and five varieties in 24 families and 60 genera of ferns and lycophytes are recorded for state of Acre, Brazil. Data about habit, habitat, material examined, and distribution of each taxon are also presented. Acre contains about one-third of the species of ferns and lycophytes estimated to occur in the Brazilian Amazon region. INTRODUCTION The Project of the Flora of Acre State, Brazil, is a collaborative endeavour between the Federal University of Acre (UFAC) and The New York Botanical Garden (NY). For some groups of plants, specialists were invited to contribute, such as for ferns and lycophytes. This checklist is a result of the collaborative project. The first expeditions to Acre and the Brazilian Amazon took place in 1900-1901 when Ernest Heinrich Georg Ule collected along the rivers Tejo and Jurua-Mirim. Locally, this area is called “Regiéo do Alto Rio Jurua.” In 1911-1912, a second expedition was conducted by Ule in the region of the Acre River. The first Brazilian botanist to explore the state was Jodo Geraldo Kuhlmann, who travelled with Marechal Rondon through Amazonia in 1923. Later, in 1933, Boris Alexander Krukoff collected in the region, especially around the Macaua River, in the Basin of Purus River. During the 1960s and 1970s, several botantists collected in Acre State, the most prominent being Joio Murca Pires, Ghillean T. Prance, Enrique Forero, Paul J. Maas, and Paulo G. Windisch. Other important collections from Acre were made for the Project RADAMBRASIL during the 1970s by Luis Coélho, Cid Ferreira, Bruce Nelson, and Thomas Croat. They were among the members of three expeditions to Acre (Silveira, 2003). In 1979 the Herbarium of the Federal Universty of Acre (HPZ) was established. Its main goal was to have a reference collection for the flora of Acre. In 1990 the herbarium signed an international agreement of cooperation with The New York Botanical Garden (NY) to promote studies of the Acre flora. During the last twenty years Douglas Daly (NY), Marcos Silveira (UFAC), and several other collaborators from many institutions have greatly increased the number of collections in Acre. According to Daly (personal communication), at the beginning of the project “Floristics and Economic Botany of Acre, Brazil’ the Herbarium of the Federal University of Acre (HPZ) had 2,000 specimens. It now has more than 30,000 specimens, including non-vascular and vascular plants. These collections provided many new records of ferns (e.g., Adiantum decoratum 231 FERN GAZ. 18(5): 230-263. 2009 Maxon & Weath., 4. poeppigianum (Kuhn) Hieron, A. scalare R. M. Tryon, Bolbitis oligarchica (Baker) Hennip., Tectaria draconoptera (D. C. Eaton) Copel., Thelypteris membranacea (Mett.) R. M. Tryon; Labiak & Prado, 2007) and new species (e.g., Adiantum windischii J. Prado and Triplophyllum boliviense J. Prado & R. C. Moran; Prado, 2005; Prado & Moran, 2008). Acre State is located in northwestern Brazil in the southwestern part of the Brazilian Amazon. It covers an area about 153,149 km~ and harbours great variation in topography, soils, and climates. This variation promotes a high diversity of vegetation types, floristic affinities, and life forms. Several factors divide the state into two regions: the southeastern region is drained by the Purus River and by the Basin of the Rivers Acre/Purus. Here there is a pronounced dry season, and several species are shared with Central Brazil and with other extra Amazonian vegetation types (Prado & Gibbs, 1993). The second region is northwestern Acre, drained by the Jurua River. This area is continuously wet, lacking a dry season. Along its border with Peru are the mountainous Moa Range and Divisor Range (locally called the Serra do Moa and Serra do Divisor, respectively). These ranges shelter several Andean species and genera such as Cyathea bipinnatifida (Baker) Domin (Cyatheaceae), Preris haenkeana C. Presl, (Pteridaceae), Solanopteris bifrons (Hook.) Copel. (Polypodiaceae); flowering plant examples include Croton lechleri Miill Arg. (Euphorbiaceae), Ladenbergia lambertiana (A. Braun ex Mart.) Klotzsch, L. oblongifolia (Rubiaceae), and Monolena primuliflora Hook. f. (Melastomataceae)). This region has open vegetation called “Campinas” and “Caatingas Amazénicas” on whitish sandy soils. The flora of this region has affinities with those of the Rivers Rio Madeira and Negro (Silveira, 2003). preliminary checklist of the Acre Flora (http://www.nybg.org/bsci/acre/ I/checklist.html), which includes all vascular plants and fungi, lists about 3,273 species (including infraspecific taxa), of which 156 are either ferns or lycophytes, these belonging to 50 genera. In general, ferns and lycophytes from Acre are poorly known and papers containing some information about them are few and sparse (Tryon & Conant, 1975; Windisch, 1979; Prado, 2005, and Labiak & Prado, 2007). The present study revealed 178 species, 60 genera, and five varieties in 24 families (Table 1). According to our estimates, Amazonian Brazil has ca. 550 species of ferns and lycophytes; thus, Acre contains about 32% of that total number. LIST OF SPECIES The present list is based on material from the following herbaria: CEN, HB, HPZ, HRCB, INPA, K, MG, MO, NY, R, RB, SP, UC. In 2001 the senior author collected about 250 specimens in Acre. In total, about 820 specimens have been examined for the present checklist. The list is organised in alphabetic order by families, genera, species, and varieties. The arrangement of familes and genera follows Smith ef al. (2006). For each species and variety the full name of the taxon is presented with the reference of the original publication, as well as additional information such as: habit, habitat, material examined, and geographic distribution. Voucher information for each species and variety is organised in alphabetic order by municipality; material collected by same collector for the same locality appears in ascending numerical order. Author abbreviations follow Pichi-Sermolli (1996). In general, synonyms are omitted, except the basionym of the accepted name. PRADO & MORAN: CHECKLIST OF THE FERNS OF ACRE STATE, BRAZIL 232 Table 1. Synopsis of taxa in Acre State, Brazil. Families Number of genera Number of spp. Number of varieties Anemiaceae Aspleniaceae Blechnaceae Z 3 Cyatheaceae Z 4 Dennstaedtiaceae 2 2 Dryopteridaceae 10 2} | Gleicheniaceae Z 2 Hymenophyllaceae 4 17 Lindsaeaceae l is 3 Lomariopsidaceae 3 8 Lycopodiaceae 2 2 Lygodiaceae l 2 Marattiaceae l 3 Metaxyaceae l l Polypodiaceae 9 28 Psilotaceae | Pteridaceae 8 30 Saccolomataceae l A Salviniaceae l Schizaeaceae l l Selaginellaceae l 7 Tectariaceae Z 6 Thelypteridaceae l 12 l Woodsiaceae S a Totals: 24 60 178 5 233 FERN GAZ. 18(5): 230-263. 2009 ANEMIACEAE Anemia phyllitidis (L.) Sw., Syn. Fil. 155. 1806. Osmunda phyllitidis L., Sp. Pl. 1064. 1753. Habit/Habitat: Herb; terrestrial on riverside vegetation and on Terra Firme forest. Material examined: BRAZIL. ACRE: Assis Brasil, D. C. Daly et al. 9779 (HPZ, NY); Brasiléia, D. C. Daly et al. 9895 (HPZ, NY). Distribution: Mexico, Greater Antilles, except Puerto Rico, Mesoamerica, tropical and subtropical South America. ASPLENIACEAE Asplenium angustum Sw., Vet. Ak. Handl. 38: 66, tab. 4, fig. 1. 1817. Habit/Habitat: Herb; epiphyte in forest on Terra Firme. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, G JT. Prance et al. 12637 (NY); Mancio Lima, D. C. Daly et al. 8953 (HPZ, NY). Distribution: N South America. Asplenium auritum Sw., J. Bot. (Schrader) 1800(2): 52. 1801. Habit/Habitat: Herb; epiphyte in forest on Terra Firme. Material examined: BRAZIL. ACRE: Manoel Urbano, D. C. Daly et al. 11466 (NY): Mancio Lima, G T. Prance et al. 11977 (NY); Marechal Thaumaturgo, D.C. Daly 10531 (NY); Porto Valter, P J. M. Maas et al. P12930 (MG, NY), P. J. M. Maas et al. P13180 (NY); Santa Rosa, D. C. Daly et al. 11305 (NY); Sena Madureira, M. Silveira et al. 617 (HPZ, NY); Tarauaca, G 7. Prance 7247 (MG, NY), G. T. Prance et al. 7382 (NY), M. Silveira et al. 921 (NY). Distribution: S Mexico, Antilles, Mesoamerica, N South America. Asplenium cirrhatum Rich. ex Willd., Sp. Pl. 5: 321. 1801. Habit/Habitat: Herb; epiphyte on flooded vegetation. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, G T. Prance et al. 12202 (NY); Mancio Lima, M. Silveira et al. 1245 (NY). Distribution: Mexico, Antilles, Mesoamerica and tropical South America. Asplenium cuneatum Lam., Encycl. 2: 309. 1786. Habit/Habitat: Herb; epiphyte on Terra Firme. Material examined: BRAZIL. ACRE: Porto Valter, P J. M. Maas et al. P13103 (NY). Distribution: Antilles, Mesoamerica and tropical South America, Polynesia, Africa. Asplenium delitescens (Maxon) L. D. Gomez, Brenesia 8: 52. 1976. Diplazium delitescens Maxon, Contr. U.S. Natl. Herb. 10: 497. 1908. Habit/Habitat: Herb; terrestrial grows riverside vegetation and on Terra Firme forest. Material examined: BRAZIL. ACRE: Assis Brasil, D. C. Daly et al. 9811 (HPZ, NY); Marechal Thaumaturgo, D. C. Daly et al. 7337 (HPZ, NY). Distribution: S Mexico, Antilles, Mesoamerica, N South America. Asplenium hallii Hook., Sp. Fil. 3: 202. 1860. Habit/Habitat: Herb; epiphyte on Terra Firme. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1166 (HPZ, SP), J. Prado et al. 1306 (HPZ, SP), J. Prado et al. 1379 (HPZ, SP), L. R. Marinho 251 PRADO & MORAN: CHECKLIST OF THE FERNS OF ACRE STATE, BRAZIL 234 (NY); P. J. M. Maas et al.P12794 (NY); Mancio Lima, G T. Prance et al. 2850 (MG, NY); G 7. Prance et al. 12246A (MG, NY); J. Prado et al. 1225 (HPZ, SP) Distribution: N South America. Asplenium juglandifolium Lam., Encycl. 2: 307. 1786. Habit/Habitat: Herb; epiphyte on flooded vegetation near river margins. Material examined: BRAZIL. ACRE: Buyjari, D. C. Daly et al. 8464 (HPZ, NY), D. C. Daly et al. 9330 (NY); Cruzeiro do Sul, G T. Prance et al. 2786 (MG, NY): Mancio Lima, J. Prado et al. 1224 (HPZ, SP), J. Prado et al. 1318 (HPZ, SP); Marechal Thaumaturgo, D. C. Daly et al. 7764 (NY); Porto Acre, A. R. S. Oliveira et al. 761 (NY); Rodrigues Alves, J. Prado et al. 1262 (HPZ, SP). Distribution: Mexico, Jamaica, Mesoamerica, tropical South America. Asplenium ortegae Murakami & R. C. Moran, Ann. Missouri Bot. Gard. 80: 23. 1993. Habit/Habitat: Herb; terrestrial . Material examined: BRAZIL. ACRE: Cruzeiro do Sul, D. C. Daly et al. 11707 (NY). Distribution: N South America. Asplenium pearcei Baker, Syn. Fil. (ed. 2) 483. 1874. Habit/Habitat: Herb; epiphyte on trees in Terra Firme forests. Material examined: BRAZIL. ACRE: Brasiléia, D. C. Daly et al. 11906 (NY); Cruzeiro do Sul, J. Prado et al. 1350 (HPZ); Feij6, P. Delprete et al. 8543 (NY); Mancio Lima, G.T. Prance et al.12168 (NY); Manoel Urbano, D. C. Daly et al. 11226 (NY); Marechal Thaumaturgo, D. C. Daly et al. 7336 (HPZ, NY); Porto Valter, G 7. Prance et al. P13269 (NY), P. J. M. Maas et al.P13295 (NY), S. R. Lowrie et al. 525 (NY); Santa Rosa, D. C. Daly et al. 9948 (NY); Tarauaca, C. Herinhaus et al. 397 (NY); Without locality, J. Jangoux et al. 85-096 (MG, NY). Distribution: N South America. Asplenium pedicularifolium St. Hilaire, Voy. Distr. ea 1: 380. 1833. abit/Habitat: Herb; epiphyte on forest on Terra Firm Material examined: BRAZIL. ACRE: Mancio Lima, D, Cc Daly et al. 8926 (HPZ, NY), G. T. Prance et al. 12246 (NY). Distribution: Guianas and Brazil. Asplenium serratum L., Sp. Pl. 2: 1079. 1753. Habit/Habitat: Herb; epiphyte on forest on Terra Firme and on flooded vegetation near river margins. Material examined: BRAZIL. ACRE: Assis Brasil, D. C. Daly et al. 9808 (HPZ, NY); Brasiléia, D. C. Daly et al. 6771 (HPZ, NY); Bujari, D. C. Daly et al. 9328 (NY), M. de Pardo et al. 80 (NY); Cruzeiro do Sul, D. C. Daly et al. 11717 (NY), G T. Prance et al. 2785 (NY), P. J. M. Maas et al. P12863 (MG, NY), J. Prado et al. 1187 (HPZ, SP), J. Prado et al. 1197 (HPZ), J. Prado et al. 1317 (HPZ, SP), J. Prado et al. 1340 (HPZ, SP), J. Prado et al. 1382 (HPZ, SP); Mancio Lima, J. Prado et al. 1135 (HPZ., SP); Manoel Urbano, D. C. Daly et al. 9136 (NY), D. C. Daly et al. 11241 (NY); Placido de Castro, C. Figueiredo & I. Riveiro 592 (NY), L. G Lohmann & E. C. de Oliveira 492 (NY); Rodrigues Alves, J. Prado et al. 1237 (HPZ, SP), J. Prado et al. 1255 (HPZ, SP), J. Prado et al. 1291 (HPZ, SP); Santa Rosa, D. C. Daly et al. Za FERN GAZ. 18(5): 230-263. 2009 10053(HPZ, NY), D. C. Daly et al. 10984 (NY), D. C. Daly et al. 11106 (NY); Sena Madureira, D. C. Daly et al. 7841 (HPZ, NY). Distribution: S Florida, Mexico, Antilles, Mesoamerica, tropical and subtropical South America. Asplenium stuebelianum Hieron., Hedwigia 47: 222. 1908. Habit/Habitat: Herb; epiphyte or terrestrial, in open forest with bamboo. Material examined: BRAZIL. ACRE: Feijé, D. C. Daly et al. 8500 (NY); Mancio Lima, D. C. Daly et al. 8952 (HPZ, NY); Marechal Thaumaturgo, D. C. Daly et al. 10524 (NY), D. C. Daly et al. 10552 (NY); Rio Branco, D. C. Daly et al. 6871A (HPZ, NY). Distribution: Tropical South America. BLECHNACEAE Blechnum occidentale L., Sp. Pl. 2: 1077. 1753. Habit/Habitat: Herb; terrestrial in forest on Terra Firme. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, P J. M. Maas et al. P13242 NY). Distribution: S EUA, Mexico, Antilles, Mesoamerica, tropical and subtropical South America. Salpichlaena hookeriana (Kuntze) Alston, Bull. Misc. Inform. Kew 1932: 312. 1932. Spicanta hookeriana Kuntze, Revis. Gen. Pl. 2: 81. 1891. Habit/Habitat: Herb; climbing on flooded vegetation near stream margin Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 11 “ (HPZ, SP); Mancio Lima, J. Prado et al. 1229 (HPZ, SP). Distribution: N South America. Salpichlaena volubilis (Kaulf.) J. Sm., J. Bot. — 4: 168. 1841. Blechnum volubile Kaulf., Enum. Filic. 159. 18 Habit/Habitat: Herb; climbing on flooded niin near stream margin Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. nae, SP). Distribution: Antilles, Mesoamerica, and tropical South America. CYATHEACEAE Alsophila cuspidata (Kunze) D. S. Conant, J. Arnold Arbor. 64: 371. 1983. Cyathea cuspidata Kunze, Linnaea 9: 101. 1834 Habit/Habitat: Tree-fern; forest on Terra Firme. Material examined: BRAZIL. ACRE: Brasiléia, D. C. Daly et al. 9881 (NY), D. C. Daly et al. 11886 (NY); Marechal Thaumaturgo, D. C. Daly et al. 7427 (HPZ, NY). Distribution: Mesoamerica and tropical South America. Cyathea microdonta (Desv.) Domin, Pteridophyta 263. 1929. Polypodium microdontum Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 319. 1811. Habit/Habitat: Tree-fern; forest near river margins. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1190 (HPZ, SP); Manoel Urbano, D. C. Daly et al. 11558 (NY). Distribution: S Mexico, Antilles, Mesoamerica and N South America. PRADO & MORAN: CHECKLIST OF THE FERNS OF ACRE STATE, BRAZIL 236 Cyathea pilosissima (Baker) Domin, Pteridophyta 262. 1929. Polypodium pilosissimum Baker, Syn. Fil. ed. 2: 457. 1874. Habit/Habitat: Tree-fern; in disturbed forest on Terra Firme. Material examined: BRAZIL. ACRE: Mancio Lima, D. C. Daly et al. 11603 (NY); Rodrigues Alves, J. Prado et al. 1231 (HPZ, SP). Distribution: Mesoamerica and N South America. Cyathea pungens (Willd.) Domin, Pteridophyta 263. 1929. Polypodium pungens Willd., Sp. Pl. 5: 206. 1810. Habit/Habitat: Tree-fern; flooded vegetation near stream margins. Material examined: BRAZIL. ACRE: Brasiléia, D. C. Daly et al. 6823 (HPZ, NY), L. Ferreira & L. C. Ming 117 (HPZ, NY); Mancio Lima, D. C. Daly et al. 1252 (NY); Manoel Urbano, D. C. Daly et al. 9087 (NY); Marechal Thaumaturgo, D. C. Daly et al. 10239 (NY); Santa Rosa, D. C. Daly et al. 10165 (NY), D. C. Daly et al. 11175 (NY). Distribution: Antilles and N South America. _ DENNSTAEDTIACEAE Dennstaedtia bipinnata (Cav.) Maxon, Proc. Biol. Soc. Wash. 51: 39. 1938. Dicksonia bipinnata Cav., Descr. Pl. 154. 1802 Habit/Habitat: Tree-fern; flooded vegetation near stream margins. Material examined: BRAZIL. ACRE: Manoel Urbano, D. C. Daly et al. 11418 (NY). Distribution: S Mexico, Antilles, Mesoamerica and N South America. Pteridium caudatum (L.) Maxon, Proc. U.S. Natl. Mus. 23: 631. 1901. Pieris caudata L., Sp. Pl. 2: 1075. 1753. Habit/Habitat: Herb; terrestrial in cleared land be-side tracks. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1178 (HPZ, SP); Mancio Lima, G T. Prance et al. 2864 (NY). Distribution: S Florida, Mexico, Antilles, Mesoamerica and N South America. DRYOPTERIDACEAE Bolbitis lindigii (Mett.) Ching, Index Filic., Suppl. 3: 48. 1934. Chrysodium lindigii Mett., Ann. Sci. Nat., Bot., sér. 5, 2: 205. 1864. Habit/Habitat: Herb; terrestrial, in open forest on undulating terrain, except for baixios (waterlogged low-lying areas). Material examined: BRAZIL. ACRE: Mancio Lima, G T. Prance et al. 120694 (NY): Marechal Thaumaturgo, D. C. Daly et al. 10252 (NY). Distribution: Mesoamerica and N South America. Bolbitis nicotianifolia (Sw.) Alston, Bull. Misc. Inform. Kew 1932: 310. 1932. Acrostichum nicotianifolium Sw., Syn. Fil. 13: 199. 1806. Habit/Habitat: Herb; terrestrial, in open forest on undulating terrain, except for baixios (waterlogged low-lying areas). Material examined: BRAZIL. ACRE: Bujari, D. C. Daly et al. 9477 (NY), W. R. Anderson 12122 (NY); Cruzeiro do Sul, J. Prado et al. 1349 (HPZ); Manoel Urbano. D. C. Daly et al. 9128 (NY), D. C. Daly et al. 11413 (NY); Porto Valter, P J. M. Maas et al. P13292 (NY); Sena Madureira, D. C. Daly et al. 8072 (NY). Distribution: Antilles, Mesoamerica and N South America. 237 FERN GAZ. 18(5): 230-263. 2009 Bolbitis oligarchica (Baker) Henipp., Amer. Fern J. 65: 30. 1975. Acrostichum oligarchicum Baker, Syn. Fil. 418. 1868. Habit/Habitat: Herb; terrestrial, in forest. Material examined: BRAZIL. ACRE: Mancio Lima, M. Silveira et al. 1361 (NY). Distribution: Mesoamerica and N South America. Ctenitis refulgens (Klotzsch ex Mett.) Vareschi, Fl. Venezuela 1: 404. 1969. Phegopteris refulgens Klotzsch ex Mett., Ann. Sci. Nat., Bot., sér. 5, 2: 240. 1864 Habit/Habitat: Herb; terrestrial, in forest. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1343 (HPZ, SP). Distribution: Mexico, Mesoamerica and N South America. Cyclodium guianense (Klotzsch) L. D. Gomez, Phytologia 60: 371. 1986. Aspidium guianense Klotzsch, Linnaea 20: 364. 1847 Habit/Habitat: Herb; terrestrial, in Terra Firme forest. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1342 (HPZ, SP): Without locality, J. Jangoux et al. 85-034 (NY). Distribution: N South America. Cyclodium meniscioides (Willd.) C. Presl, Tent. Pterid. 85. 1836. Aspidium meniscioides Willd., Sp. Pl. 5: 218. 1810. Infraspecific: var. meniscioides Habit/Habitat: Herb; terrestrial, in Terra Firme forest. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1330 (HPZ, SP); Mancio Lima, J. Prado et al. 1226 (HPZ, SP); J. Prado et al. 1228 (HPZ, SP). Distribution: N South America. Didymochlaena truncatula (Sw.) J. Sm., J. Bot. (Hooker) 4: 196. 1841-1842 [1841]. Aspidium truncatulum Sw., J. Bot. (Schrader) 1800(2): 36. 1801. Habit/Habitat: Herb; terrestrial in primary forest, unudulating terrain, in places dissected by many small streams. Material examined: BRAZIL. ACRE: Brasiléia, D. C. Daly et al. 9880 (HPZ, NY), C. Daly et al. 11893 (NY); Cruzeiro do Sul, G T. Prance et al. 12087 (MG, NY), G Prance et al. 12377 (NY); Mancio Lima, D. G. Campbell et al. 8913 (NY). Distribution: S Mexico, Antilles, Mesoamerica and N South America. Tropical Asia and Africa. D. Z, Dryopteris patula (Sw.) Underw., Native Ferns ed. 4 117. 1893. Aspidium patulum Sw., Kong. Vetensk. Acad. Handl. 1817(1): 64. 1817. Habit/Habitat: Herb; terrestrial in moist forest on Terra Firme, canopy discontinuous. Material examined: BRAZIL. ACRE: Placido de Castro, C. Figueiredo & I. Riveiro 560 (NY); Sena Madureira, D. C. Daly et al. 8117 (NY). Distribution: S Mexico, Antilles, Mesoamerica and tropical South America. Elaphoglossum discolor (Kuhn) C. Chr., Index Filic. 306. 1905. Acrostichum discolor Kuhn, Linnaea 36: 53. 1869. Habit/Habitat: Herb; terrestrial or epipetric, in Campina. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, D. C. Daly et al. 10607 (NY). PRADO & MORAN: CHECKLIST OF THE FERNS OF ACRE STATE, BRAZIL 238 J. Prado et al. 1159 (HPZ, SP). J. Prado et al. 1170 (HPZ, SP), J. Prado et al. 1188 (HPZ, SP), J. Prado et al. 1284 (HPZ, SP), J. Prado et al. 1322 (HPZ, SP), J. Prado et al. 1324 (HPZ, SP), J. Prado et al. 1329 (HPZ), J. Prado et al. 1388 (HPZ, SP); Mancio Lima, J. Prado et al. 1223 (HPZ, SP). Distribution: N South America. Elaphoglossum flaccidum (Fée) T. Moore, Index Filic. 356. 1862. Acrostichum flaccidum Fée, Mem. Foug. 2: 35, tab. 7, fig. 2. 1845. Habit/Habitat: Herb; epiphyte, in Terra Firme forest. Material examined: BRAZIL. ACRE: Rodrigues Alves, J. Prado et al. 1266 (HPZ, SP); Xapuri, Silva et al.186 (CEN, SP). Distribution: Antilles and N South America. Elaphoglossum glabellum J. Sm., London J. Bot. 1: 197. 1842. Habit/Habitat: Herb; epiphyte, in Terra Firme forest. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1297 (HPZ, SP). Distribution: S Florida, Mexico, Antilles, Mesoamerica, and N South America. Anand luridum (Fée) H. Christ, Neue Denkschr. Allg. Schweiz. Ges. Gesammten Naturwiss. 36(1): 33. 1899. parte luridum Fée, Mem. Foug. 2: 35, tab. 19, fig. 1. 1845. Habit/Habitat: Herb; epiphyte, in Terra Firme forest. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, /. Prado et al. 1152 (HPZ, SP), J. Prado et al. 1174 (HPZ, SP), J. Prado et al. 1198 (HPZ, SP), J. Prado et al. 1310 (HPZ, SP), J. Prado et al. 1327 (HPZ, SP); Mancio Lima, J. Prado et al. 1219 (HPZ, SP), J. Prado et al. 1220 (HPZ, SP); Rodrigues Alves, J. Prado et al. 1238 (HPZ, SP), J. Prado et al. 1241 (HPZ, SP), J. Prado et al. 1260 (HPZ, SP), J. Prado et al. 1281 (OEs, SP). Distribution: Antilles, Mesoamerica and N South America. Elaphoglossum plumosum (Fée) T. Moore, Index Filic. 364. 1862. Acrostichum plumosum Fée, Mem. Foug. 54, tab. 20, fig. 1. 1845. Habit/Habitat: Herb; epiphyte, in Terra Firme forest. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1154 (HPZ, SP), J. Prado et al. 1285 (HPZ); Mancio Lima, J. Prado et al. 1221 (HPZ, SP), J. Prado et al. 1390 (HPZ, SP). Distribution: N South America. Elaphoglossum raywaense (Jenm.) Aslton, Bol. Soc. Brot., ser. 2, 32: 24. 1958. Acrostichum raywaense Jenm., Ferns Brit. W. Ind. 341. 1909 Habit/Habitat: Herb; epiphyte, in Terra Firme forest. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, /. Prado et al. 1314 (HPZ. SP): Mancio Lima, D. C. Daly et al. 8948 (HPZ, NY). Distribution: N South America. Elaphoglossum styriacum Mickel, Brittonia 39: 326, fig. 4I-K. 1987. Habit/Habitat: Herb; epiphyte, in Terra Firme forest. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, /. Prado et al. 1381 (HPZ, SP). 2a9 FERN GAZ. 18(5): 230-263. 2009 Distribution: N South America. Elaphoglossum tantalinum Mickel, Brittonia 39: 326, fig. 4I-K. 1987. Habit/Habitat: Herb; epiphyte, in Terra Firme forest. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1179 (HPZ, SP), J. Prado et al. 1321 (HPZ, SP), J. Prado et al. 1384 (HPZ, SP). Distribution: N South America. Lastreopsis effusa (Sw.) Tindale, Victoria Naturalist 73: 184. 1957. Polypodium effusum Sw., Prodr. 1788. Habit/Habitat: Herb; terrestrial, Koieateal slope between Baixio and higher terrace. Material examined: BRAZIL. ACRE: Manoel Urbano, D. C. Daly et al. 11421 (NY). Distribution: S Mexico, Antilles, Mesoamerica, and tropical South America. Lomagramma guianensis (Aubl.) Ching, Amer. Fern J. 22: 17. 1932. Polypodium guianense Aubl., Hist. Pl. Guiane 962. 1775. Habit/Habitat: Herb; hemiepiphyte, in Terra Firme forest. Material examined: BRAZIL. ACRE: Brasiléia, D. C. Daly et al. 6757 (HPZ, NY), D. C. Daly et al. 6786 (NY), D. C. Daly et al. 11882 (NY); M. Silveira et al. 1662 (NY); Cruzeiro do Sul, J. Prado et al. 1292 (HPZ, SP); Mancio Lima, D. C. Daly et al. 11623 (NY); Marechal Thaumaturgo, D. C. Daly et al. 7430 (HPZ, NY); Rodrigues Alves, J. Prado et al. 1252 (HPZ, SP), J. Prado et al. 1278 (HPZ, SP); Xapuri, L. G Lohmann & E. C. de Oliveira 590 (NY). Distribution: Greater Antilles (except Jamaica) and tropical South America. Polybotrya caudata Kunze, Linnaea 9: 23. 1834. Habit/Habitat: Herb; hemiepiphyte, in Terra Firme forest. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1295 (HPZ, SP): Mancio Lima, J. Prado et al. 1143 (HPZ, SP), J. Prado et al. 1145 (HPZ, SP) G T. Prance et al. 12069 (NY), M. Silveira et al. 1662 (NY). Distribution: S Mexico, Mesoamerica and N South America. Polybotrya pubens Martius, Icon. Pl. Crypt. 87, tab. 25. 1834. Habit/Habitat: Herb; hemiepiphyte, in Terra Firme forest. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1205 (HPZ, SP). J. Prado et al. 1207 (HPZ, SP), J. Prado et al. 1394 (HPZ, SP) Distribution: N South America. Rumohra adiantiformis (G. Forst) Ching, Sinensia 5: 70. 1934. Polypodium adiantiforme G. Forst, Prodr. 82. 1786 Habit/Habitat: Herb; terrestrial, in Terra Firme forest. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, /. Prado et al. 1157 (HPZ. SP). J. Prado et al. 1181 (HPZ, SP), J. Prado et al. 1287 (HPZ, SP). Distribution: Bermuda, Antilles, N South America. Africa, Madagascar, New Zealand and Australia. GLEICHENIACEAE Gleichenella pectinata (Willd.) Ching, Sunyatsenia 5: 276. 1940. PRADO & MORAN: CHECKLIST OF THE FERNS OF ACRE STATE, BRAZIL 240 Mertensia pectinata Willd., Kongl. Vetensk. Acad. Nya Handl. 25: 168, tab. 4. 1804. Habit/Habitat: Herb; terrestrial along forest margins. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1189 (HPZ, SP): Mancio Lima, G 7: Prance et al. 12119 (MG, NY), M. Silveira et al. 1348 (HPZ, NY). Distribution: S Mexico, Antilles, Mesoamerica and tropical South America. Sticherus remotus (Kaul.) Chrysler, Amer. 7 Bot. 31: 483. 1944. Mertensia remota Kaulf., Enum. Filic. 39. 1824 Habit/Habitat: Herb; terrestrial along forest margins. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1300 (HPZ, SP). Distribution: Mesoamerica and N South America. HYMENOPHYLLACEAE Didymoglossum krausii (Hook. & Grev.) C. Pres], Hymenophyllaceae 115. 1843. Trichomanes krausii Hook. & Grev., Icon. Filic. 2: pl. 149. 1830 Habit/Habitat: Herb; epiphyte in forest. Material examined: BRAZIL. ACRE: Manoel Urbano, D. C. Daly et al. 9115 (HPZ, Distribution: Florida, Mexico, Antilles, Mesoamerica, tropical South America. Didymoglossum ovale E. Fourn., Acta Bot. Neerl. 11: 296. 1962. Habit/Habitat: Herb; epiphyte along river margins. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1288 (HPZ): Mancio Lima, J. Prado et al. 1230 (HPZ, SP). Distribution: S Mexico, Antilles, Mesoamerica and N South America. Didymoglossum punctatum (Poir.) Desv., Mém. Soc. Linn. Paris. 6: 330. 1827. Trichomanes punctatum Poir., Encycl. 8: 64. 1808 Habit/Habitat: Herb; epiphyte. Material examined: BRAZIL. ACRE: Marechal Thaumaturgo, D. C. Daly et al. 10330 NY ( ; Distribution: Antilles, Mesoamerica and N South America. Hymenophyllum angustum Bosch, Ned. Kruidk. Arch. — 183. 1861. Habit/Habitat: Herb; epiphyte in waterlogged low-lying area Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. rman etal. 1153 (HPZ, SP). Distribution: Amazonian Brazil. Hymenophyllum polyanthos (Sw.) Sw., J. Bot. (Schrader) 1800(2): 102. 1801. Trichomanes polvyanthos Sw., Prodr. 137. 1788. Habit/Habitat: Herb; epiphyte in forest. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1149 (HPZ, SP), J. Prado et al. 1389 (HPZ, SP), P. J. M. Maas et al. 12722 (NY); Mancio Lima, J. Prado et al. 1222 (HPZ, SP). Distribution: Mexico, Mesoamerica and tropical South America. Trichomanes ankersii C. Parker ex Hook. & Grev., Icon. Filic. 2(11): tab. 201. 1831. Habit/Habitat: Herb; hemiepiphyte, in Terra Firme forest. 241 FERN GAZ. 18(5): 230-263. 2009 Material examined: BRAZIL. ACRE: Cruzeiro do Sul, G 7 Prance et al. 11854 (MG NY), J. Prado et al. 1150 (HPZ, SP), J. Prado et al. 1158 (HPZ, SP), J. Prado et al. 1168 (HPZ, SP), J. Prado et al. 1214 (HPZ, SP), J. Prado et al. 1312 (HPZ, SP), 2. Prado et al. 1387 (HPZ, SP); Rodrigues Alves, J. Prado et al. 1236 (HPZ, SP), J. Prado et al. 12357 (HPZ, SP). Distribution: Mesoamerica and N South America. Trichomanes bicorne Hook., Icon. Plan. 9: tab. 892. 1854. Habit/Habitat: Herb; terrestrial, in Terra Firme forest. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, D. C. Daly et al. 10613 (NY), J. Prado et al. 1147 (HPZ, SP), J. Prado et al. 1176 (HPZ, SP), J. Prado et al. 1268 (HPZ, SP), J. Prado et al. 1323 (HPZ, SP), J. Prado et al. 1271 (HPZ, SP). Distribution: N South America. Trichomanes crispum L., Sp. Pl. 2: 1097. 1753. Habit/Habitat: Herb; terrestrial, in Terra Firme forest. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1313 (HPZ, SP), J. Prado et al. 1380 (HPZ, SP). Distribution: S Mexico, Antilles, Mesoamerica, and N South America Trichomanes elegans Rich., Actes Soc. Hist. Nat. Paris 1: 114. 1792. Habit/Habitat: Herb; terrestrial, in forest near stream margins. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, G 7 Prance et al. 12225 (MG, NY), J. Prado et al. 1212 (HPZ, SP); Mancio Lima, M. Silveira et al. 1247 (NY), M. Silveira et al. 1250 (NY); Rodrigues Alves, J. Prado et al. 1364 (HPZ, SP). Distribution: Antilles, Mesoamerica, tropical and subtropical South America. Trichomanes hostmannianum (Klotzsch) Kunze, Bot. Zeitung (Berlin) 5: 352. 1847. Neurophyllum hostmannianum Klotzsch, Linnaea 18: 532. 1844 Habit/Habitat: Herb; terrestrial, in forest near stream margins on wet soils. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1194 (HPZ, SP), L. R. Marinho 217 (NY); Rodrigues Alves, J. Prado et al. 1272 (HPZ, SP). Distribution: N South America. Trichomanes macilentum vy. d. Bosch, Ned. Kruidk. Arch. 5(2); 146. 1861. Habit/Habitat: Herb; terrestrial in waterlogged low-lying areas. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1385 (HPZ, SP). Distribution: N South America. Trichomanes martiusii C. Pres|, Hymenophyllaceae 15, 36. 1843. Habit/Habitat: Herb; terrestrial in forest on Terra Forme. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1146 (HPZ, SP), J. Prado et al. 1171 (HPZ, SP), J. Prado et al. 1269 (HPZ, SP), J. Prado et al. 1326 (HPZ, SP), NV. A. Rosa 657 (NY); Mancio Lima, J. Prado et al. 1217 (HPZ, SP). Distribution: N South America. Trichomanes pilosum Raddi, Opusc Sci. 3: 296. 1819. Habit/Habitat: Herb; terrestrial in forest on Terra Firme. PRADO & MORAN: CHECKLIST OF THE FERNS OF ACRE STATE, BRAZIL 242 Material examined: BRAZIL. ACRE: Cruzeiro do Sul, /. Prado et al. 1182 (HPZ, SP). Distribution: Bolivia, Paraguay and Brazil (south of the Amazon Basin). Trichomanes pinnatum Hedw., s./., Fil. Gen. Sp. tab. 4, fig. 1. 1799. Habit/Habitat: Herb; terrestrial in forest on Terra Firme, on clay and sand soils. Material examined: BRAZIL. ACRE: Assis Brasil, D. C. Daly et al. 9608 (HPZ, NY); Cruzeiro do Sul, J. Prado et al. 1183 (HPZ), J. Prado et al. 1311 (HPZ, SP), D. C. Daly et al. 11640 (NY); Mancio Lima, D. C. Daly et al. 11589 (HPZ, NY), G T. Prance et al. 12178 (MG, NY); Porto Valter, D. C. Daly et al. 7576 (HPZ, NY), D. C. Daly et al. 11739 (NY), P. J. M. Maas et al.13010 (MG, NY); Rodrigues Alves, J. Prado et al. 1244 (HPZ); Sena Madureira, G 7. Prance et al. 7730 (NY), G T. Prance et al. 7948 (NY). Distribution: S Mexico, Antilles, Mesoamerica and N South America. Obs.: the material studied for this taxon cited above probably belong to more than two or three species, but more studies are necessary to clarify this situation. Trichomanes tanaicum Hook. ex J. W. Sturm., Fl. Bras. 1(2): 260. 1859. Habit/Habitat: Herb; hemiepiphyte, in Terra Firme forest. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1209 (HPZ); Rodrigues Alves, J. Prado et al. 1254 (HPZ, SP), J. Prado et al. 1258 (HPZ). Distribution: N South America. Trichomanes trollii Bergdolt, Flora 127: 256, 264, fig. 3. 1933. Habit/Habitat: Herb; terrestrial, in forest on Terra Firme. Material examined: BRAZIL. ACRE: Rodrigues Alves, J. Prado et al. 1366 (HPZ, SP). Distribution: N South America. Vandenboschia collariata (Bosch) Ebihara & K. Iwats., Blumea 51: 242. 2006. Trichomanes collariatum Bosch, Ned. Kruidk. Arch. 4: 368. 1859 [1858]. Habit/Habitat: Herb; epiphyte in forest on Terra Firme. Material examined: BRAZIL. ACRE: Without locality, J. Jangoux et al. 85-053 (MG, NY), J. Jangoux et al. 850-73 (MG, NY). Distribution: Mexico, Mesoamerica and N South America. LINDSAEACEAE Lindsaea divaricata Klotzsch, Linnaea 18: 547. 1844. Habit/Habitat: Herb; terrestrial in forest on Terra Firme, on sandy soils. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 115] (HPZ, SP), J. Prado et al. 1177 (HPZ, SP), J. Prado et al. 1270 (HPZ, SP), T. B. Croat & A. Rosas Jr. 62671 (NY); Rodrigues Alves, J. Prado et al. 1359 (HPZ, SP). Distribution: Mesoamerica and tropical South America. Lindsaea hemiglossa Kramer, Acta Bot. Neerl. 6(2): 257. 1957. Habit/Habitat: Herb: terrestrial in forest on Terra Firme, on slopes. Material examined: BRAZIL. ACRE: Mancio Lima, G T. Prance et al. 12142 (NY). Distribution: N South America. 243 FERN GAZ. 18(5): 230-263. 2009 Lindsaea lancea (L.) Bedd., Suppl. Ferns S. Ind. 6. 1876. Adiantum lanceum L., Sp. Pl. ed. 2 1557. 1763. Infraspecific: var. Jancea Habit/Habitat: Herb; terrestrial in forest on Terra Firme. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1309 (HPZ, SP). Distribution: S Mexico, Antilles, Mesoamerica and tropical South America. Infraspecific: var. falcata (Dryand.) Rosenst., Hedwigia 46: 79. 1906. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1148 (HPZ, SP), J. Prado et al. 1167 (HPZ, SP), J. Prado et al. 1308 (HPZ, SP), J. Prado et al. 1309 (HPZ, SF). Distribution: Mexico, Mesoamerica and N South America. Lindsaea schomburgkii Klotzsch, Linnaea 18: 545. 1844. Habit/Habitat: Herb; terrestrial in forest on Terra Firme. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1172 (HPZ, SP). Distribution: N South America. Lindsaea sp Habit/Habitat: Herb; terrestrial in forest on Terra Firme, on clay soils. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1348 (HPZ, SP). Distribution: Peru and Brazil. Lindsaea stricta (Sw.) Dryand. Trans. Linn. Soc. London 3: 42. 1797. Adiantum strictum Sw., Prodr.: 135. 1788. Infraspecific: var. stricta Habit/Habitat: Herb; terrestrial, in Campina. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, D. C. Daly et al. 10633 (NY), J. Prado et al. 1335 (HPZ, SP). Distribution: Widespread in the Neotropics. Lindsaea ulei Hieron., Hedwigia 44: 365. 1905. Habit/Habitat: Herb; terrestrial in forest on Terra Firme, on sandy soils. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1206 (HPZ, SP), J. Prado et al. 1356 (HPZ, SP); Rodrigues Alves, J. Prado et al. 1232 (HPZ, SP), J. Prado et al. 1243 (HPZ, SP), J. Prado et al. 1367 (HPZ), J. Prado et al. 1369 (HPZ, SP). Distribution: N South America. LOMARIOPSIDACEAE Cyclopeltis semicordata (Sw.) J. Sm., Bot. Mag. 72: 36. 1846. Polypodium semicordatum Sw., Prodr. 132. 1788. Habit/Habitat: Herb; terrestrial in forest on Terra Firme, on sandy soils. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, D. C. Daly et al. 11721 (NY); Manoel Urbano, D. C. Daly et al. 11427 (NY), M. Silveira et al. 1574 (NY); Marechal Thaumaturgo, D. C. Daly et al. 7342 (HPZ, NY), D. C. Daly et al. 10473 (NY), D. C. Daly et al. 10810 (NY); Porto Acre, C. A. Cid & B. Nelson 2877 (NY), P. J. M. Maas et al. 13247 (MG, NY); Quixada, C. A Cid & A. Rosas 2961 (NY); Reserva Extrativista PRADO & MORAN: CHECKLIST OF THE FERNS OF ACRE STATE, BRAZIL 244 Chico Mendes, Colocacaéo Simitumba, A. R. S. Oliveira et al. 131 (HPZ, NY); Santa Rosa, D. C. Daly et al. 11367 (NY); Santa Rosa, D. C. Daly et al. 10978 (NY); Sena Madureira, D. C. Daly et al. 7857 (HPZ, NY); Rio Branco, D. C. Daly et al. 6881 (HPZ, NY); Tarauaca, G. T. Prance et al. 7516 (NY), D. C. Daly et al. 8667 (HPZ, NY), D. C. Daly et al. 8799 (HPZ, NY); Xapuri, D. C. Daly et al. 8420 (NY); Without locality, /. Jangoux et al. 85-074 (MG, NY). Distribution: S Mexico, Antilles, Mesoamerica and N South America. Lomariopsis japurensis (Mart.) J. Sm., Hist. Fil. 140. 1875. Acrostichum japurense Mart., Icon. Pl. Crypt. 86, tab. 24. 1834. abit/Habitat: Herb; hemiepiphyte, in Terra Firme forest. Material examined: BRAZIL. ACRE: Assis Brasil, D. C. Daly et al. 9817 (HPZ, NY); Cruzeiro do Sul, P. J. M. Maas et al. P12886 (NY); Mancio Lima, G. T: Prance et al. 12055 (NY); Marechal Thaumaturgo, L. G. Lohmann et al. 480 (NY); Santa Rosa, D. C. Daly et al. 9945 (NY) Distribution: Mesoamerica and N South America. Lomariopsis nigropaleata Holttum, Bull. Misc. Inform. Kew 1939: 618. 1940. Habit/Habitat: Herb; hemiepiphyte, in Terra Firme forest. Material examined: BRAZIL. ACRE: Brasiléia, S. R. Lowrie et al. 726 (NY); Cruzeiro do Sul, J. Prado et al. 1202 (HPZ, SP), J. Prado et al. 1211 (HPZ, SP), J. Prado et al. 1352 (HPZ, SP); Mancio Lima, D. C. Daly et al. 11568 (NY); Marechal Thaumaturgo, L. C. Ming & L. A. Ferreira 324 (NY). Distribution: N South America. Lomariopsis prieuriana Fée, Mem. Foug. 2: 66, pl. 25, fig. 1. 1845. Habit/Habitat: Herb; hemiepiphyte, in Terra Firme forest. Material examined: BRAZIL. ACRE: Mancio Lima, J. Prado et al. 1137 (HPZ, SP); Rodrigues Alves, J. Prado et al. 1363 (HPZ, SP). Distribution: Mesoamerica and N South America. Nephrolepis biserrata (Sw.) Schott, Gen. Fil. pl. 3. 1834. Aspidium biserratum Sw., J. Bot. (Schrader) 1800(2): 32. 1801. Habit/Habitat: Herb; epiphyte in forest on Terra Firme near stream marg Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. io (HPZ, SP), J. Prado et al. 1180 (HPZ, SP), J. Prado et al. 1199 (HPZ, SP); Mancio Lima, J. Prado et al. 1216 (HPZ, SP). Distribution: Florida, Mexico, Antilles, Mesoamerica, tropical South America. Nephrolepis brownii (Desv.) Hovenkamp & Miyam., Blumea 50: 293. 2005. Nephrodium brownii Desv., Mem. Soc. Linn. Paris 6: 252. 1827. Habit/Habitat: Herb; epiphyte in forest on Terra Firme near stream margin Material examined: BRAZIL. ACRE: Cruzeiro do Sul, P. Delprete et al. "ee (NY); Mancio Lima, D. C. Daly et al. 11395 (NY). Distribution: Florida, S Mexico, Antilles, Mesoamerica, tropical South America, Paleotropics. 245 FERN GAZ. 18(5): 230-263. 2009 Nephrolepis pendula (Raddi) J. Sm., J. Bot. (Hooker) 4: 197. 1841-1842 [1841]. Aspidium pendulum Raddi Opusc. Sci. 3: 289. 1819. Habit/Habitat: Herb; epiphyte on Terra Firme. Material examined: BRAZIL. ACRE: Tarauaca, GT. Prance et al. 7296 (NY). Distribution: S Mexico, Mesoamerica and N South America. Nephrolepis rivularis (Vahl) Mett. ex Krug, Bot. Jahrb. Syst. 24: 122. 1897. Polypodium rivulare Vahl, Eclog. Amer. 3: 51. 1807. Habit/Habitat: Herb; epiphyte on Varzea forest. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1286 (HPZ, SP), J. Prado et al. 1325 (HPZ, SP); Mancio Lima, G T. Prance et al. 12179 (MG, NY). Distribution: S Mexico, Antilles, Mesoamerica and N South America. LYCOPODIACEAE Huperzia dichotoma (Jacq.) Trevisan, Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Lycopodium dichotomum Jacq., Enum. Stirp. Vindob. 314. 1762 Habit/Habitat: Herb; epiphyte in forest on Terra Firme. Material examined: BRAZIL. ACRE: Without locality, M. Silveira s.n. (HPZ). Distribution: Florida, Mexico, Antilles, Mesoamerica and N South America. Lycopodiella cernua (L.) Pic.Serm., Webbia 23: 166. 1968. Lycopodium cernuum L., Sp. Pl. 2: 1103. 1753 Habit/Habitat: Herb; terrestrial on sandy soils in open places. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1160 (HPZ, SP), J. Prado et al. 1191 (HPZ, SP), J. Prado et al. 1331 (HPZ), A. Rosas et al. 313 (NY), D. C. Daly et al. 10599 (NY), D. C. Daly et al. 10683 (NY), G. T. Prance et al. 12525 (MG, NY), L. R. Marinho 64 (NY), L. R. Marinho 212 (NY), P. Delprete et al. 8063 (NY). Distribution: Pantropical. LYGODIACEAE Lygodium venustum Sw., J. Bot. (Schrader) 1801(2): 303. 1803. Habit/Habitat: Herb; vine-like, in forest on Terra Firme. Material examined: BRAZIL. ACRE: Manoel Urbano, M. Silveira et al. 1543 (HPZ, NY); Rio Branco, L. Coélho & A. Rosas 1925 (NY); Tarauaca, C. Ehringhaus et al. 346 NY). Distribution: Mexico, Antilles, Mesoamerica and tropical South America. Lygodium volubile Sw., J. Bot. (Schrader) 1801(2): 303. 1803. Habit/Habitat: Herb; vine-like, in forest on Terra Firme. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1371 (HPZ, SP). Distribution: Mexico, Antilles, Mesoamerica and tropical South America. MARATTIACEAE Danaea leprieurii Kunze, Mém. Acad. Roy. Sci. (Turin) 5(1790-1791): 429 tab. 9. 1793 Habit/Habitat: Herb; terrestrial in forest on Terra Firme, near stream margi Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1204 (HPZ, SP). PRADO & MORAN: CHECKLIST OF THE FERNS OF ACRE STATE, BRAZIL 246 Distribution: S Mexico, Antilles, Mesoamerica, and N South America. Danaea nodosa (L.) Sm., Mem. Acad. Roy. Sci. (Turin) 5(1790-1791): 420, tab. 9, fig. 11. 1793. Achrostichum nodosum L., Sp. Pl. 2: 1070. 1753. Habit/Habitat: Herb; terrestrial in forest on Terra Firme. Material examined: BRAZIL. ACRE: Brasiléia, D. C. Daly et al. 6787 (HPZ, NY), D. C. Daly et al. 9884 (HPZ, NY); Feijo, P. Delprete et al. 8301A & B (NY); Sena Madureira, D. C. Daly et al. 8137 (HPZ, NY); Without locality, J. Jangoux et al. 85- 071 (NY). Distribution: S Mexico, Antilles, Mesoamerica and N South America. Danaea trifoliata Reichenb. ex Kunze, Analecta Pteridogr. 4, tab. 2. 1837. Habit/Habitat: Herb; terrestrial in forest on Terra Firme, near stream margins. Material examined: BRAZIL. ACRE: Rodrigues Alves, J. Prado et al. 1365 (HPZ, SP). Distribution: Guianas and Brazil. METAXYACEAE Metaxya rostrata (Kunth) C. Presl, Tent. Pterid. 60, tab. 1, fig. 5. 1836. Aspidium rostratum Kuhn, Nov. Gen. Sp. (quarto ed.) 1: 12. 1815 [1816]. Habit/Habitat: Herb; terrestrial in forest on Terra Firme, near stream margins. Material examined: BRAZIL. ACRE: Cruzeiro do Sul, D. C. Daly et al. 11705 (NY), J. Prado et al. 1186 (HPZ, SP), J. Prado et al. 1301 (HPZ, SP), J. Prado et al. 1320 (HPZ, SP); Mancio Lima, D. C. Daly et al. 11624 (NY), M. Silveira et al. 1365 (NY): Marechal Thaumaturgo, D. C. Daly et al. 10526 (NY); Porto Valter, D. C. Daly et al. 7577 (NY), P. J. M. Maas et al. P13150 (NY); Without locality, J. Jangoux et al. 85- 043 (NY), J. Jangoux et al. 85-108 (NY). Distribution: S Mexico, Lesser Antilles, Mesoamerica and N South America. POLY PODIACEAE Campyloneurum abruptum (Lindm.) B. Léon, Fieldiana, Bot., n.s. 32: 172. 1993. Polypodium repens var. abruptum Lindm., Ark. Bot. 1: 245. 1903. Habit/Habitat: Herb; epiphyte, Varzea forest. Material examined: BRAZIL. ACRE: Manoel Urbano, D. C. Daly et al. 11225 (NY), M. Silveira et al. 1582 (NY). Distribution: N South America. Campyloneurum angustifolium (Sw.) Fée, Mem. Foug. 5: 257. 1852. Polypodium angustifolium Sw., Prodr. 130. 1788. Habit/Habitat: Herb; epiphyte in forest on Terra Firme. Material examined: BRAZIL. ACRE: Bujari, M. de Pardo et al. 93 (NY); Cruzeiro do Sul, G T. Prance et al. 12617 (NY), L. R. Marinho 163 (NY); Feijo, P. Delprete et al. 8311 (NY); Marechal Thaumaturgo, D. C. Daly et al. 10936 (NY); Porto Valter, P. J. M. Maas et al. 12931 (NY); Tarauaca, D. C. Daly et al. 8628 (HPZ, NY), G T. Prance et al. 7384 (NY), G T. Prance et al. 7249 (NY), M. Silveira et al. 1146 (NY). Distribution: S Mexico, Lesser Antilles, Mesoamerica and tropical South America. —_ 247 FERN GAZ. 18(5): 230-263. 2009 Campyloneurum aphanophlebium (Kunze) T. Moore, Index Filic. 223. 1861. Polypodium aphanophlebium Kunze, Bot. Zeitung (Berlin) 3(17): 288. 1845. Habit/Habitat: Herb; epiphyte, forest in broad, meandering stream vally in steeply hilly terrain. Material examined: BRAZIL. ACRE: Brasiléia, G. P. da Silva et al. 173 (NY); Mancio Lima, D. C. Daly et al. 11601 (NY); Marechal Thaumaturgo, D. C. Daly et al. 7496 (HPZ, NY), D. C. Daly et al. 10255 (NY); Porto Valter, P J. M. Maas et al. P1318! (NY); Rodrigues Alves, J. Prado et al. 1261 (HPZ, SP); Santa Rosa, D. C. Daly et al. 10173 (NY); Tarauaca, D. C. Daly et al. 8630 (HPZ, NY), G T. Prance et al. 7267 Distribution: Mesoamerica and N South America. Campyloneurum fuscosquamatum Lellinger, Amer. Fern J. 78: 21, fig. 4, 10. 1988. Habit/Habitat: Herb; epiphyte on Terra Firme forest and Varzea forest. Material examined: BRAZIL. ACRE: Brasiléia, D. C. Daly et al. 6802 (HPZ, NY), D. C. Daly et al. 11894 (NY); Bujari, W. R. Anderson 12123 (NY); Feijo, P. Delprete et al. 8580 (NY); Mancio Lima, G T. Prance et al. 12052 (MG, NY), G T. Prance et al. 12052A (NY); Quixada, E. Forero et al. 6388 (MG, NY); Santa Rosa, D. C. Daly et al. 10151 (NY), D. C. Daly et al. 10994 (NY); Sena Madureira, D. C. Daly et al. 7856 (HPZ, NY), G T. Prance et al. 7694 (NY); Tarauaca, D. C. Daly et al. 8620 (HPZ, NY), D.C. Daly et al. 8760 (HPZ, NY); Without locality, J. Jangoux et al. 85-039 (MG, NY). Distribution: N South America. Campyloneurum ophiocaulon (Klotzsch) Fée, Mem. Foug. 258. 1852. Polvpodium 0.0, a 4-parameter logistic curve was a This has a lower asymptote corresponding to the value of RWC or F. Fy, in dry material and an upper asymptote at the maximum (fully recovered) sides ot hi. Where F\/F,,, in dry material was ~0 or unknown, a 3-parameter logistic sincting at 0 was fitted, ies either case, the IC-50 value (point of inflection) of the curve was taken as a measure of the time for ‘half-recovery’. The relation of RETR to irradiance is generally well fitted by negative-exponential curve of the form y = (4 — e **), where = RETR and x = PPFD and A and & are constants, at least at low irradiances (<400 mmol m2 sl), Deviations may arise through electron flow to sinks other than photosynthetic carbon fixation (bryophytes often show non-saturating electron flow of this kind (Marschall & Proctor, 2004)), or through photoinhibition depressing electron flow at high irradiances. A useful test of goodness-of-fit of the exponential curve is that the product Ak should approximate to the dark-adapted value of F,/F,,, (c. 7.5—-8.5). NPQ is generally well fitted by a logistic curve starting at 0. The parameter /—g p is also often well fitted by a logistic curve, but the fit is very sensitive to the top few data points, which can lead to misleading estimates of the upper asymptote and IC-50 value; suspect values have been omitted from Table 4. Some comment may be made on measures of water content and water availability. The water content of desiccation-tolerant plants is often given as a percentage of dry weight. This is useful for comparisons within a population of a single species, but not for comparisons between different populations or different species because of differences in the proportion of cell-wall materials and cell contents. Water content relative to that at full turgor (RWC) is a much more widely useful measurement, and has been emphasised here. Relative humidity (RH) of the air is only useful if the temperature is known (in this paper c. 20°C), and its biological relevance is limited. Evaporation rates depend on saturation deficit, the difference between the absolute humidity of the air and the saturated absolute humidity at the same temperature (which can be looked up in tables); it is usually expressed in terms of partial pressure of water vapour (kPa). Water potential (YY) which defines the tendency for water to move from one place to another, is usually expressed in pressure units (MPa); for air, it can be calculated from relative humidity and temperature. For the physical and mathematical relations between these quantities see, e.g., Slatyer (1967), Nobel (1974), Jones (1992). RESULTS Table 1 lists some water-content data for the ferns investigated, expressed as a percentage of oven-dry weight, and as relative water content. Water content at full turgor on a dry-weight basis varies widely even within the same species; at full turgor RWC for all species is 1.0 by definition. A number of measurements were made on ‘air dry’ material. The figures in the table show clearly that ‘air dry’ can embrace a substantial range in actual water content measured as RWC; cold dry weather outdoors will tend to give low absolute (and relative) humidity indoors, and warm or humid weather the opposite. Fern material equilibrated in a desiccator at c. 74% RH had an RWC around 0.14—0.16; material equlibrated at 43% RH showed RWC mostly below 267 FERN GAZ. 18(5): 264-282. 2009 0.10. Of the air dried material, Asplenium trichomanes (1) and Polypodium cambricum (1) were probably dried to an equivalent of c.60—-70% RH at 20 °C, and the other samples at c.30-40% RH. Three small saxicolous Asplenium species The recovery of relative water content and the fluorescence parameter F\/F',, on re- wetting after a few days’ desiccation in Asplenium ruta-muraria L.and A. trichomanes L. is shown in Figures la and 1b and Table 2. A. trichomanes that had been dried to 14% RWC recovered rather faster than material dried to a RWC of 7.5%; the half-recovery times of F\/F,,, in the fitted curves of Figure 1b are respectively 1.7 and 3.9 hours, but the asymptotes of the two curves are not significantly different. A much rarer species than either of these in Britain, A. septentrionale L. was reported as highly desiccation tolerant by Kappen. Leaves collected from a population in North Wales in September 2005, were allowed to air dry. The chlorophyll fluorometer was not available from then until January 2006, by which time the leaves had been dry for four months. However, on re-wetting (12 Jan.) they recovered their normal fresh appearance and appeared fully turgid the following day. Ten leaves re-wetted after 3 days moist followed by 2 days air dry gave F\/F,,, 0.754 + 0.035 after 24 h rehydration (Figure Ic). Of these three Asplenium species, A. ruta-muraria showed the highest light-saturation level, followed by A. septentrionale with A. trichomanes the lowest; all three showed at least some indication of non-saturating electron flow at high irradiances (Figure | 1a). Two further species of Asplenium Asplenium adiantum-nigrum L. sometimes accompanies the other small saxicolous Asplenium species in crevices of rocks and walls, but usually avoids the most sun- exposed and water-stressed situations. Leaves of this species from a mortared pebble wall facing the sea at Budleigh Salterton were promisingly firm textured, and looked possible candidates for desiccation tolerance. However, re-wetted after a few days’ desiccation at 43% RH they showed only slight temporary signs of recovery of F\/Fy, for the first few hours, and by the following day it was clear they would not recover further (Table 2.) Plants from another population, on a steep dry roadside bank near Kennford, Devon, behaved quite differently. Figure 2 shows the rapid and progressive loss of weight of leaf segments dried at 74% and 43% RH, with fitted exponential curves; the asymptotes approximated to the figures in Table 1; the leaves dried quickly, losing half their water within 6.2 h at 74% RH (-41 MPa) and 4.7 h at 43% RH (—114 MPa). The recovery of F\/F, on re-wetting after the two desiccation regimes is shown in Figure 3. The data and the fitted curves suggest that intensity of drying has a marked effect, but neither the asymptotes nor the half-recovery times are significantly different, and the slope factor just fails to reach significance at the 5% level. Asplenium obovatum Viv. dried rather more slowly than A. adiantum- nigrum; the half-drying times seen in Figure 4 are around I1 h at 74% RH and 9 h at 43% RH Recovery of F\/F,,, on re-wetting was both less complete and more variable than in the Kennford A. adiantum-nigrum. After drying at 74% RH, the dark green leaves of the previous season showed only a transient rise in F\/F,,, on re-wetting; further recovery failed completely. However, in material dried at 43% RH, the older leaves recovered consistently and well. The bright green new-season’s leaves recovered well from drying at 74% RH, but very erratically from the 43% RH treatment (Figure 5). NS / ft PROCTOR: DESICCATION TOLERANCE IN SOME BRITISH FER HU %Eh I JWUdIUOS JAVA HY %Pl I 1UDdUOS JOB AIP JIB, JUdJUOD JIB $10°0 + ZS0'0 SPT LSI IOOF CHO] LEl FE br . - Se F90¢ wnjoaliajul 9000 FTLOO] GTLFSSI | 9TOFIDIO] LS FTEE - - OL FEIT (CZ) wnoiquis unipodaAjog : 7 . 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(Z) WNASIU-UINJUDIPD “y 7 7 ~ = LiO'0 + £800 67 +967 9€ F PPE (|) wnssiu-unjupipp “y 7 . - - P1000 + SLO'0 SO0F6L Il = 80I (Z) saupwmoyoia “y : 7 ’ ‘ LEOOFOPIO| IFFCO bp + 61 ([) saupwoyoiy “y 7 ° e = €10°0 + 890°0 Pr+F602 Ol + 60F pipiAnu-pind wuniuajdsp OMA Md% IMA Md% OMA MA% Moiozim — (ed t1I-) (Rd Ib-) 14 , ‘PlOJUUDY WO SI (Z) WinssIU-wunjUDIPY “py SUOWaYyeS Ysloypng wor, st (1) wnssIU-UunjuDIPD “PY “Kparyoadsas [OT “URL PUR QOOT 99q Ul “YSIa;pnyy wo yoq o19M wnraqups unipodajog pure saupwoyoiy uniualdsp UO SJUSLUDINSBIU JO SJ9S OA} SUL (OMY) 19 UOS JayeM IANLIAI se puK TYBIIM Alp % SB (DOT IB Bd PII— 2) HY %EP pure (9,07 18 Wd Or“) HY YPL 1 Ae TIA wuNLginba ur pue Ap de, pue (5,01 18 IYSIOM Arp-udAo % se) JOS.IN} [[N] 1 SAR Ula} JO JUDJUOD JAVA *T IQUE, 269 FERN GAZ. 18(5): 264-282. 2009 The rusty-back fern, Asplenium ceterach The curling of the dry leaves in Asplenium ceterach initially prevented measurement of F\/F yy in the early stages of rehydration, so the data in Figure 6 are from leaves dried under light pressure to keep them flat. The rise in F\/F,,, was rather slower to get going in A. ceterach, but climbed more steeply once it had started. Light saturation was reached at around half noon summer sunlight, but showed some depression of electron flow at higher irradiances. An experiment with Polystichum aculeatum (L.) Roth Pinnae of Polystichum aculeatum dried rather quickly, giving half-drying times of 12.9 h at 74% RH and 8.4 h at 43% RH, reaching practical equilibrium with the atmosphere in the desiccator in the course of a week. The corresponding equilibrium water contents (asymptotes of the curves) were 0.109 and 0.069 RWC. Remoistened after 9 days, they showed small but erratic increases of F\/F,,, for an hour or two, but these were not maintained and by the following day it was clear that all the pinnae were moribund. The polypodies, Polypodium cambricum L. and P. interyectum Shivas These two species were very tolerant of desiccation, and differ in important respects from the ferns just considered. They dried very much more slowly (Figure 7); the half- drying times for P. interjectum at 74% RH (-41 MPa) and 43% RH (—114 MPa} were respectively 2.33 days (56 h) and 1.4 days (34 h). These drying times are almost an order of magnitude longer than those seen in Asplenium adiantum-nigrum, around five times longer than in A. obovatum. As a consequence, the asymptotes of the exponential curves, 0.074 and 0.037 RWC respectively, are substantially below the figures in Table 1 from weighings after a week’s drying. Clearly these latter figures do not reflect equilibrium with the air. The recovery curves of F\/F,,, in Figures 8 and 9 thus start from relative water contents above equilibrium with the ambient air — but still well below levels at which normal metabolism is possible, or survivable by typical vascular-plant mesophyll tissues. The curves differ significantly only in the initial level of F\/F;,,. Little sign of recovery is apparent in the first hour after re-wetting, either in the physical appearance of the leaves (Figure 10) or in F\/F,,,, but after that recovery is rapid. The half-recovery times are around 2h, and in the same range as in the other species examined (Table 3). Photosynthetic electron flow saturated at around 40% of noon summer sunlight, but was markedly depressed at high irradiances so that the effective saturation irradiance must be below the PPFDgs5o, figure in Table 4 (Figure | 1b). DISCUSSION The results confirm that the leaves of some ferns are remarkably tolerant to desiccation, recovering well from drying for a week or more to a water content of 15—20% of their dry weight, and a tissue relative water content (RWC) of 4-7%. The mesophyll of typical vascular plants is generally damaged beyond recovery if RWC falls below about 30% (or a water potential below about —7 MPa), so these ferns are far more tolerant of desiccation than most drought-tolerant vascular plants (Larcher 1995). Their tolerance is comparable with such mosses as TJortula (Syntrichia) ruralis or Racomitrium lanuginosum. However it is also clear there are degrees of desiccation tolerance, and that tolerance may vary within species. Of our Asplenium species, A. septentrionale is PROCTOR: DESICCATION TOLERANCE IN SOME BRITISH FERNS 270 Asplenium ruta-muraria 1 (8) Ss i u® a ae Time /h Asplenium trichomanes oO Ss xX E Ww a he Qa FSF, (2) = RWC (1) Time /h Asplenium septentrionale uf dL ti tiith ie ese itil ti iiiul 0.01 0.1 1 10 100 Time /h Figure 1. Recovery on remoistening of (a) F\/F m and RWC in air-dry leaves of Asplenium ruta-muraria. Fitted logistic curves. Mean = s.d., n = 3, (b) F\/F,,, in two samples of air-dry leaves of A. trichomanes (sample (1) from 0.140 RWC, (2) from 0.075 RWC), and recovery of RWC for sample (1). Means (7 = 3), error bars omitted for clarity (log scale), (c) F\/F,,, in A. septentrionale after four months air dry, with fitted 4-parameter logistic curve. Mean + s.d., 7 = FERN GAZ. 18(5): 264-282. 2009 ari uonespdyas y pz doye “Ay d1ojog rosin} ypny ye “O77 6100 + £180 010°0 C80 7 L100 ¥ 1c8°0 wnpooliomuy “ Lv0'0 * 6920 €£0'0 + C£8 0 ' C100 ¥ 898°0 (z) wnotaquivs wnipodajog ° : 601°0 + €0L'0 - ; (|) wnotqups unipodAjog £c0'0 + 6820 yO0'0 F C18 0 : 8100 F 9180 yoo.sa}a2 “Vy ell0 FLIP O +CC1'0 F OCS 0 - L100 + 008°0 winyvaogo *V SL0'0 + 9b9'0 £00 + StL 0 - LeO'0 F S8L'°0 (z) wnsStu-unjuvipp “py . 6S0°0 + SL0'0 : (|) eunssiu-wnjupdipp ‘py - 4 6S0°0 ¥ LOL'0 - (Z) SauDWOY IIA “YW : 0c0'0 ¥ 9620 7 (|) SauDWOYILY “Y * ‘ S00 F 9SL'0 7 piuipinu-pjnd wuniuadsp : . Aip- (edW PII) HY %EPr (@dW It-) HY %bL Preys uoNRodISOp sa1seds ‘uOnRIPAYas Jaye Y pZ puke uoNRddISap asojaq sos.iny pny we “oy/Ly °Z aTqRE, PROCTOR: DESICCATION TOLERANCE IN SOME BRITISH FERNS 272 Asplenium adiantum-nigrum O 74% RH A 43% RH a ae oe 0 20 40 60 80 100 120 140 160 Desiccation time /h Figure 2. Drying data for leaves of Asplenium adiantum-nigrum in desiccators at ~74% and 43% RH (-41 and —114 MPa); the fitted exponential curves give half-drying times of respectively 6.2 h and 4.7 h. Means + s.d., 1 = 6. Asplenium adiantum-nigrum 1 til elas 0.8- — _ fa , 06- i 0.4- ff ras 02- O 74% RH A = 4S 43% RH Qo 0.1 1 10 100 Recovery time /h Figure 3. Recovery of F,/F,,, on re-wetting leaves of Asplenium adiantum-nigrum after 6 days drying at 74% RH (—41 MPa; circles and full line) and 43% RH (-114 MPa; triangles and broken line; error bars omitted for clarity), with fitted 4-parameter logistic curves. Means + s.d., 7 = 6. 273 FERN GAZ. 18(5): 264-282. 2009 probably the most tolerant, followed by A. ruta-muraria and A. trichomanes. In A. adiantum-nigrum it appears that some populations are tolerant, but others are not, or perhaps that tolerance varies with season, as Kappen found. A. obovatum seems (from one sample) to be less tolerant than A. adiantum-nigrum, and to show some differences between young and old leaves. Two leaves of A. marinum L. from near Kynance Cove, Cornwall in early June 2001 did not recover after drying, but this species should be tested from more sites. Kappen (1964) found the leaves of summer-green ferns such as Dryopteris filix-mas are similar to flowering plant leaves in their sensitivity to drying, with little variation through the season. Species which remained green through the winter were in general more tolerant in the winter and spring months, but the limiting leaf saturation-deficit given by Kappen for Polystichum lobatum (aculeatum) is little different from his corresponding figure for Dryopteris filix-mas. The experiment reported here gave no indication of desiccation tolerance in P. aculeatum. A high degree of desiccation tolerance is seen in Polypodium cambricum and P. interjectum; the ‘Polypodium vulgare’ that Kappen found the most desiccation tolerant of all his ferns was probably P. vulgare L. sensu stricto. The Asplenium species dried quickly, with a half-drying time of a few hours, and there is clearly little stomatal control over water loss; six leaves of A. ceterach gave half-drying times ranging from 2.5 to 9.0 h (depending on size), with a mean of 4.8 + 2.3 h. Drying of Polvstichum aculeatum was only a little slower. The Polypodium species dried much more slowly, with half-drying times in P. interjectum of a day or two, suggesting that the stomata close under water stress. This may be related to the habitats of the plants. The Asplenium species primarily occupy open, brightly lit and often sunny rock-crevice habitats. In bright sunshine in summer, a small Asplenium is sitting immovably in the steep temperature gradient close to the ground surface. In this situation a turgid, actively metabolising plant needs the cooling effect of transpiration to keep its leaves from reaching lethal temperatures (c.45—50°C), and leaves of these small ferns are correspondingly dependent on a constant supply of water from the root system to maintain turgor. Once air dry, desiccation-tolerant plants can withstand much higher temperatures. Plants tall and open enough for the air to circulate freely around their leaves do not face this overheating hazard, and Polypodium leaves, held free of the substratum on flexible petioles would be expected to be more closely coupled to the temperature of the air. Perhaps Asplenium ceterach, with its dry leaves curved away from the substratum and protected by their thick scaly covering, comes somewhere between A. ruta-muraria and Polypodium cambricum in this respect. In their photosynthetic responses, the small Asp/enium species have something in common the desiccation-tolerant mosses in the same habitats. In this they differ from A. ceterach and P. cambricum, but these latter two are still not, photosynthetically, unequivocal sun plants. The ferns rooted in rock crevices can probably all draw on substantial reserves of water at depth, which dry up only in hot dry spells in summer. Polypodium is probably drawing mainly on a more superficial reserve of water, which both dries up and is replenished more frequently, and this must underlie its success as an epiphyte (Zotz & Hietz, 2001). Recovery took place at similar rates in all the ferns examined, with half- recovery times around 2-4 hours, in contrast to the wide range of recovery rates in bryophytes (Table 3). hese desiccation-tolerant ferns are interesting in an ecological and evolutionary context. Proctor & Tuba (2002) postulated two strategies for plant life on land, PROCTOR: DESICCATION TOLERANCE IN SOME BRITISH FERNS 274 Asplenium obovatum i 1 oy = _L- 08— New growth : oO = 74% RH (-41 MPa) = | aw 06- | = G 7 / ® = 04- | Bi | \ 0.2 . ‘i. tae a NS: 0 ye ae 14 4 ~O- Mature leaves hii ~7- New growth g . 43% RH (-114 MPa) ww 06> , = = 4 o = §4- 0.254 0 a ee eae eee ee o @ 5 6 id 8 9 10 11 12 Date /May 2006 Figure 4. The course of RWC during drying and re-wetting with (a) desiccation at 74% RH (—-41 MPa) and (b) at 43% RH (—114 MPa). Note in (a) the lower final RWC and less complete recovery of new leaves, and lack of significant difference between old and new growth in (b); half-drying times in (a) are about 11 h, and in (b) abour 9 h. a be FERN GAZ. 18(5): 264-282. 2009 Asplenium obovatum 1 ~@ 74% RH, old -A- 74% RH, new -O- 43% RH, old ee | \ uw Og] \ de re 7 \ -~7- 43% RH, new c = ee / S seed iw = y Vv o2- ‘ 9 J ae y 0 Faas v nV, a “Pe | a ae ia | 345 6 7 8 9 10 11 12 13 14 15 Date /May 2006 Figure 5. The course of F\/F,,, over the duration of the experiment. Note more rapid decline in F\/F,,, in the new growth, parallelling its faster drying, and the varied recovery of old and new growth following the two desiccation treatments. Asplenium ceterach 1 ja CE aa (a 6 Se ee 0.8 + ‘ 0.6 = w e Re i 0.4 - 0.2- OQ Fv/Fm 4 gs RWC 0 Ly ryt of 1 10 100 Time /h Figure 6. Recovery of F\/F,, and RWC (from different samples) in Asplenium ceterach, (data set (2) of Table 3) Mean + s.d., 7 = 2 (F\/F,,), 1 = 3 (RWC). Fitted 4- parameter logistic curves. Note the very wide error bars in the lower part of the curve, reflecting the variation between replicates in the ‘latent’ time before recovery of F. vem begins. PROCTOR: DESICCATION TOLERANCE IN SOME BRITISH FERNS Polypodium interjectum ae Lua \ O 74% RH oa 4 A 43% RH i es ‘se a a nada aie as ae a ae 0 2 4 6 8 10 Days elapsed Figure 7. Drying data for RWC in Polypodium interjectum at 74 and 43% RH (41 and ~114 MPa), with fitted exponential curves. Polypodium cambricum Litt js She a he tp 0.8 — . 4 L . 0.6 - on Ww | B uw 0.4 - = 9, 02 “ O 74% RH | a @ 43% RH b 0 l 0.1 1 10 100 Time /h Figure 8. Recovery of F. Fy, following re-wetting of Polypodium cambricum atter 8 days’ drying at 74% and 43% RH (-41 and —114 MPa). The only significant difference in the parameters of the fitted curves is in their lower limit. ati FERN GAZ. 18(5): 264-282. 2009 Polypodium interectum ee pip iin ae ee SE fe O 74% RH @ 43% RH FF o =) Oo o i) a o fo) iy 1 Pre, Pe] 0.1 1 10 100 Time /h Figure 9. Recovery of F\/F,,, following re-wetting of Polypodium interjectum after 8 days’ drying at 74% and 43% RH (-41 and —114 MPa). As in P. cambricum, the only significant difference between the fitted curves is in their lower limit. Polypodium intenectum 3s DF aD D ; ; ee. - 0.8 17) 2 is en . rs) = 3 3) - 3 = _— c 52, Oo 5 © o oO ead 3 i= phe 0.6 = os ¢ @EsuS g = | 3 mo ¢ = re S E 5. =o Fe fla = —- Z>wos Ww re) = HS 0.4 = na . @© OL ; = ec © s— ie = [7 z+ cc =a Ze QO => 2 3 6 0.2 38 a= Le p= 35 a (re V : T fe Be SG | gees | PTTITy T i if T 0.01 0.1 1 Time (h) Figure 10. The 74% RH recovery curve of Figure 10 annotated with the appearance of the leaves at intervals during re-wetting; the comments relate to the successive data points. The mean value of F\/F,,, in dry leaves is arbitrarily plotted at 0.01 h. PROCTOR: DESICCATION TOLERANCE IN SOME BRITISH FERNS RETR RETR Asplenium tichomanes 250 — 4 (a) - san] en 200 5 Pd a Pa a = 150 — o- | we 100 4 “| 50 0 rrr. ia | ea oe 0 500 1000 1500 2000 Polypodium cambricum a i a a a i ad a a a al te Da a 0 500 1000 1500 2000 PPFD /umol m? s’ 278 Figure 11. Light response of photosynthetic electron flow in (a) Asplenium trichomanes, showing (very variable) non-saturating electron flow at high irradiance, and (b) Polypodium cambricum, showing (consistent) depression of electron flow at high irradiance. FERN GAZ. 18(5): 264-282. 2009 a ~ N S00 + C00 [10 + O80 Gur ist Leo oe 1 (Wd bl I-) wnoatiaiut c0'0 + 810 $00 ¥ 180 ey OF £F co 810 + £0? 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Tt Ses LOOT URL ‘Ysla[pnyD ‘enosnw-pjins wniuajdsp EPS | L8°0 IIS 89'()| Ob + 1Z8 O0OT 99d “Ysla[pnyD ‘pivpinwu-pjns wniuajdsp 0S-OI asuvi-A 0S-Ol asuvs-X | )_s -_wt [ow I/ SJUDWUILUOD pue sa1sads db - | OdN %$ OC ddd ‘sjutod BJP MAJ ISIYSIY SY} Ul 10.19 0] DAIISUDS AIDA SI SOAIND SY] JO WY YI Seep dy} 01 WZ AJOJI"FsHeS APUIIOIYNS B IABS SAINO ONSIDO] 9 UDYM ATUO dh-] 10} UIAIS udaq aAvy siojoWeIeg “sprepuEis jUL]d-se[NoseA JeoidAy Aq sanyea CqN Ysly Moys saioods [py “aAoqe UDAIS sonjRA ,uoNLNes % 6, BY} Yas wopyas sonjoeid ul ABU wnoLiquins unipodadjog pue Yyoviajad uniuajdsp yey) OS ‘dOURIPR.AI YSTY Je MOL] UO.NIETA INAYyUASOOYd JO uolssoidap dUIOs ysva] ye MOYS saIddds JOYIO dy JO ISO, “(HOOT “JOIOIg WZ [[eyosseP\) sarAydoA1q yULII]O}-uUONROdISap UI PoArasqo A[UOWUOD MOL UOI}I]O SuNvinyes-uou sy} MOYS [[e aypuoLmuatdas “py puke saubWOYI “"p ‘pliDanu-vin« wniuazdsp ‘saduRIpR.L JdYs1Y TV “SAIND ay} Jo Led sIy} OF WJ poos v DAIS saloads ay] []B *[-S Z-LW [OULU QOP MOJO S9dURIPRLII JO] sjUIOd-eep dy} 0} pany saaino jenusuodxo -SANLBIU WO payejnoyes oie (IS F UBIUT) SAN|VA UONBINIS 9% CG OY] SUID} JULID[O} UOIBOOISAp awWOS JO ssaj}oUIeIed asUOdsaI-1YST] “fp ATqRL, 281 FERN GAZ. 18(5): 264-282. 2009 exemplified by the homoiohydric mainstream vascular plants on the one hand, and the poikilohydric, desiccation-tolerant bryophytes on the other. Alongside the contrast between homoiohydry and poikilohydry there is another contrast, between endohydry and ectohydry. In vascular plants the physiologically important free water is in the xylem; they are endohydric, and the plant body is isolated from its surroundings by a waterproof and water-repellent epidermis. In most bryophytes, the physiologically important water is in capillary spaces outside the plant; they are ectohydric. The desiccation tolerant fern sporophytes considered in this paper withstand drying (and are to that extent poikilohydric), but are endohydric. This is also true of most other vascular ‘resurrection plants’. All of these plants function as normal vascular plants when water is freely available, but they have added to their repertoire of responses the fall-back option of desiccation tolerance (already inherent in their spores) when the water supply fails. We are in a region of ecological niche-space where several adaptive strategies converge, and none is optimal. Desiccation-tolerant ferns share their habitat with fully poikilohydric (and ectohydric) mosses and liverworts, with small succulents, and (seasonally) with winter annuals. Fern gametophytes, on the other hand are unequivocally poikilohydric and ectohydric, and take their place alongside bryophytes in a diversity of habitats where some of them are as desiccation tolerant as their bryophyte neighbours (Watkins ef al., 2007). REFERENCES GILDNER, B.S. & LARSON, D.W. 1992. Seasonal changes in photosynthesis in the desiccation-tolerant fern Polypodium virginiacum. Oecologia 89: 383-389. HARTEL, O. 1940a. Physiologische Studien an Hymenophyllaceen. I. Zellphysiologische Untersuchungen. Protoplasma 34: 117-147. HARTEL, O. 1940b. Physiologische Studien an Hymenophyllaceen. II. Wasserhaushalt und Resistenz. Protoplasma 34: 489-514. HOLLOWAY, J.E. 1923a. Studies in the New Zealand Hymenophyllaceae. Part 1. The distribution of the species in Westland, and their growth-forms. Transactions of the New Zealand Institution 54: 577-618. HOLLOWAY, J.E. 1923b. Studies in the New Zealand Hymenophyllaceae. Part 2. The distribution of the species throughout the New Zealand biological region. Transactions of the New Zealand Institution 55: 67-94 ILJIN, W.S. 1931. Austrocknungsresistenz des Farnes Notochlaena Marantae R.Br. Protoplasma 13: 322-330. JONES, H.G. 1992. Plants and microclimate. 2nd edn. Cambridge University Press, Cambridge. KAPPEN, L. 1964. Untersuchungen iiber den Jahreslauf der Frost-, Hitze- und Austrocknungsresistenz von Sporophyten einheimischer Polypodiaceen (Filicinae). Flora 155: 123-166. KESSLER, M. & SIORAK, Y. 2007. Desiccation and rehyration experiments on the leaves of 43 pteridophyte species. American Fern Journal 97: 175-185. LARCHER, W. 1995. Physiological plant ecology, 3rd edn. Springer-Verlag. New York MARSCHALL, M. & PROCTOR, M.C.F. 2004. Are bryophytes shade plants? Photosynthetic light responses and proportions of chlorophyll a, chlorophyll 5 and total carotenoids. Annals of Botany 95: 593-603. MAXWELL, K. & JOHNSON, GN. 2000. Chlorophyll fluorescence — a practical guide. Journal of Experimental Botany 51: 659-668. PROCTOR: DESICCATION TOLERANCE IN SOME BRITISH FERNS 282 MUSLIN, L.E.H. & HOMANN, P.H. 1992. Light as a hazard for the desiccation- resistant ‘resurrection’ fern Polypodium polypodioides. Plant, Cell & Environment 15: 81-89, NOBEL, P.S. 1974. An introduction to biophysical plant physiology. W.H. Freeman, San Francisco. OPPENHEIMER, H.R. & HALEVY, A. 1962. Anabiosis of Ceterach officinarum Lam. et DC. Bulletin of the Research Council of Israel D3 11: 127-147. POTTS, R. & PENFOUND, W.T. 1948. Water relations of polypody fern, Polypodium polypodioides (L.) A.S, Hitchcock. Ecology 29: 43-53. PROCTOR, M.C.F. 2003. Comparative ecophysiological measurements on the light responses, water relations and desiccation tolerance of the filmy ferns Hymenophyllum wilsonii Hook. and H. tunbrigense (L.) Smith. Annals of Botany 91: 717-727 PROCTOR, M.C.F. & PENCE, V.C. 2002. Vegetative tissues: bryophytes, vascular resurrection plants and vegetative propagules. In: Black, M. & Pritchard, H.W. (Eds) Desiccation and survival in plants: drying without dying, pp. 207-237. CABI Publishing, Wallingford, UK. PROCTOR, M.C.F. & TUBA, Z. 2002. Poikilohydry and homoiohydry: antithesis or spectrum of possibilities? New Phytologist, 156: 327-49. PROCTOR, M.C.F., LIGRONE, R. & DUCKETT, J.G. 2007. Desiccation tolerance in the moss Polvtrichum formosum: physiological and fine-structural changes during desiccation and recovery. Annals of Botany 99: 75—93. REYNOLDS, T.L. & BEWLEY, J.D. 1993. Abscisic acid enhances the ability of the desiccation-tolerant fern Polypodium virginiacum to withstand drying, Journal of Experimental Botany 44: 1771-1779. RICHARDS, P.W. & EVANS, GB. 1972. Biological Flora of the British Isles. Hymenophyllum. Journal of Ecology 60: 245-268. SCHREIBER, U., BILGER, W. & NEUBAUER, C. 1995. Chlorophyll fluorescence as a nonintrusive indicator for rapid assessment of in vivo photosynthesis. In: SCHULZE, E.-D. & CALDWELL, M.M. (Eds) Ecophysiology of photosynthesis., pp. 49-70. Springer-Verlag Berlin,. SCHWAB, K.B., SCHREIBER, U. & HEBER, U. 1989. Response of photosynthesis and respiration of resurrection plants to desiccation and rehydration. Planta 177: —227. SHREVE, F. 1911. Studies on Jamaican Hymenophyllaceae. Botanical Gazette 51: 184-209, SLATYER, R.O. 1967. Plant-water relationships. Academic Press, New York & London. STUART, T.S. 1968. Revival of respiration and photosynthesis in dried leaves of Polypodium polypodioides Planta 83: 185-206. WATKINS, J.E., MACK, M.C., SINCLAIR, T.R. & MULKEY, S.S. 2007. Ecological and evolutionary consequences of desiccation tolerance in tropical fern gametophytes. New Phytologist 176: 708-717. ZOTZ, G & HIETZ, P. 2001. The physiological ecology of vascular epiphytes: current knowledge, open questions. Journal of Experimental Botany 52: 2067-2078. 283 FERN GAZ. 18(5). 2009 INSTRUCTIONS FOR AUTHORS PAPERS should not usually exceed 20 printed oor and are generally expected to be considerably shorter. Review articles, as well as reports of original research, are encouraged. Short notes are acceptable e.g. new ae The senior author should supply a fax and email address to facilitate correspondence. MANUSCRIPTS should be submitted in English (British) in electronic format (preferably) or hard copy (two copies), in 10-point Times New Roman font and double spaced. 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Species Filicum, 5. Dulau & Co., Lon MORTON, C.V. 1947. The American species of Visits section Sphaeroconium. Contr. U.S. Natl. Herb. 29(3): 139-201. STEVENSON, D.W. & LOCONTE, H. 1996. Ordinal and familial relationships of pteridophyte genera. In: CAMUS, J.M., GIBBY, M. & JOHNS, R.J. (Eds) Pteridology in perspective, pp. 435-467. Royal Botanic Gardens, Kew. JOURNAL ABBREVIATIONS: should follow Botanico Periodicum Huntianum & Supplements. Alterations from the original text at proof stage will be charged for unless they are minor points of detail. Twenty-five offprints will be supplied free to the senior author. Missouri Botanical Ga pani cae soc | (i/lili oy President: R.W. Sykes, Cesmaie House, Crosthwaite, Kendal, Cumbria LA8 8BP — E-mail: President@eBPS.org.uk _ Vice-Presidents: P.J. Acock, A.R. Busby, R.J. Cooke, A.F. Dyer, Miss J.M. Ide, M.S. Porter _ General Secretary: Dr Y.C. 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Website: ebsite: www.eBPS.org uk ee ee = A, bt hip Me p +4. 2.8 re ly b ; : 7 ( SOS fc ae each year) are Full Personal Members £20, Personal Members ing The Fern Gazett £16, Student ae Members £10, Subscribing In Institutions £33. Family ene ay aa an oon oe — n forther THE FERN GAZETTE VOLUME 18 PART 5 2009 . CONTENTS A summary of Indian cheilanthoid ferns and the discovery of Negripteris ielopenaes Afro-Arabian fern genus new to India CR. Fraser-Jenkins & C.S. Dulawat 216-229 ne 230-263 wees rey . : o ome British ferns i : 264-282 : : ee . riOES es