Sy nett Area Wiy25 Baty PoP Ht Need Beara Ry Rees tS Cha , ee Wo ie H i oe cat? ees 3 5 55) 4! y 3 P wey uty HSie iahet : fe Resa Bes "1 7 tay ore ai tas ; p Beale Ge uf haute ; bia } avian i eit Beta he 4 aa Ulnn oi aa bap : its yeas ¢ Tater A + saat OeNe re saat vO" a aN “ se Heinen nee sis erase wad r, Neate pt tnt ; Re Baresi lta Ha iH eas 13°55 uh he Mitel ‘ * cai * * ie fs see vom wi PORE Y b ae ne < et) K " reMey tw! ‘ 2 ab Neto) See Ae Seisannebat Dano n ns ts OER St areata atin nh e a 2 by ; : ese’ ? . ; J a6 ¥ a : ae ’ Pref | 2 e Pens Ae 3 es hha: est: k Slt Ahem nlaises ie RS Sere Bre st os ee oY 3 one 2 ; se rite Nottie nanan tata me i Mathiciaey Shea pees tb ary ; : i : a bites patency nn aie ead aun re See caayice re 2 yy Ng t * 1" . ate axe Teco psted yan S ios PO eRe Git Helene as : ay Ailes af Bulletin of the British Museum (Natural History) Botany series Voll4 1985-6 British Museum (Natural History) London 1986 Dates of publication of the parts No 1 : . : ; ‘ ‘ P : ; ; 29 August 1985 No2_. : : : : ; ; : ‘ ; . 19 December 1985 No 3 : . : j : 5 : : 2 ; ; 30 January 1986 No4 : : i : : F ; ; : . 27 February 1986 ISSN 0068-2292 Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset No 1 No 2 No 3 No 4 Contents Botany Volume 14 Page Cytological observations on Indian subcontinent and Chinese Dryopteris and pect aula de ee M. Gibby : : : . 1 A redisposition of the species referred to the ascomycete genus Microthelia D. L. Hawksworth . : : : . : ‘ : , : 43 A classification of the genus Dryopteris (Pteridophyta: Dryopteridaceae) C. R. Fraser-Jenkins : : ; P . ‘ ‘ orto Evolutionary cladistics of marattialean ferns C.R. Hill & J. M. Camus . : : : ; ; : ; . “219 + s t Ca Ley ‘gus 7 ern ie 2 wt - i" > tor hk. tht if ned Bulletin of the British Museum (Natural Wee) Cytological observations on Indian subcontinent and Chinese Dryopteris and Polystichum (Pteridophyta: Dryopteridaceae) Mary Gibby Botany series Vol 14 No 1 29 August 1985 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the Museum, both by the scientific staff of the Museum and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself, - available separately, and individually priced. Volumes contain about 300 pages and several volumes may appear within a calendar year. Subscriptions may be placed for one or more of the series on either an Annual or Per Volume basis. Prices vary according to the contents of the individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History), Cromwell Road, London SW7 5BD, England. World List abbreviation: Bull. Br. Mus. nat. Hist. (Bot.) © Trustees of the British Museum (Natural History), 1985 The Botany series is edited in the Museum’s Department of Botany Keeper of Botany: Mr J. F. M. Cannon Editor of Bulletin: Mr J. R. Laundon Assistant Editor: Dr A. J. Harrington Editor’s Assistant: Miss M. J. Short ISBN 0 565 08007 5 ISSN 0068-2292 Botany series Vol 14 No 1 pp 1-42 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 29 August 1985 Chinese 5 tna and Polystichum Dryopteridaceae) Mary Gibby Department of Botany, British Museum (Natural History), Cromwell Road, SW7 5BD BRITISH MUSEUM Contents (NATURAL HISTORY) onten , 3,0 AUG I'750 IIE Sag otha trices Site snis Uonevxeahsacvede ssa cecestat staecpavecstsdes ronnadbeabaedeekvawnsesesensess ei rk a OS A ENN AEDT RE Oreo eR IV DM REG Sane ASR tes ere ae Ne eee L.. PRESENTED Ek EEE SO DAE SERRE PT RERR ONT PTO Te ETT ORR Tr Ee er eT ear : IBRARY NOR MEIC RIUONONND gd 05s cc evraich x coca ds scadsiivevwns weds sober e wor as Tehava ve 7eb tee Soaks oder sesene! GENERA a aie BaP II RE cap ure Ser ttinu Sou sade nes ok sane Cowseau dass 4h 5 hes ¥¥ a5) Sein $49 Sauk tee Sued SvaeddiveveiGseee teks 2 PRR MINES MY OF MAS sa de seve cp nnik Ce eceu JAN Sac aE cen anceai ioe mage cea nonin toc aasdetauegedeet 9 III rs kaon ssi ssc cacetw deta gs ¥edc Sok wong sen cediGa Foun a noes euece taal eedeay hues 9 RRM UNCED! esses cg cise coco coven vg so Sin shen usee «Sa nans cxWare vane banaeVeions oven sdeaunas oe tes wae 12 eM CINTININLS 5, So eros ees xcact cen cmenalburss bias veces a seseceh ansuaiae boas sddeusaspyinsech@esaaves 13 IRs 65 co oi cecn aa veesetvadssaercesangGauevses oattt apy srabet Se. "6 0 cas 2. °0d a SS og? ° S0@ oe”? @o os SEN hin Yo? We" Se @ wi? ‘on «40 D Oye ~ o Of ae STS Woe, NY) x date Te at Sem at 4? A 4a a” WP ay (yes Se * Tx i v2 . >” VE \‘¥ ee ~~ “S SUN eo ae Silk ee m ar 4 \ fad xX Az o rio oy ayy “ y acl ayy ~ & | wan of \'¢ eh a gee ERR i wy =. ' ate € ze —/ LT fall y F4-- eit) ax i| Se \/ * , - aa? /~ a 1 a ey Se A me tas <3 ee a ie fol rf Jee nm ote, Pg Cee re "Tt ne ag oS ayty » » ee ‘ = Pe, were *, 8, eye oe a & ov'S e te aes. a Ye kg ew bg a 6° 2 3ia 31b . oat se Ant ee ay ‘ ping’ w Pa ne 5 mit @ Wie “i 0 sil @ 0% | ee , y * 4 Ji ® oS “ D> ’ . Y Gen a é Q Y P eed! “le sg es + ~4 - wt toa Se) Spe Vie’ pX%o Seer S “at ais Sk, 33a BD Figs 3la, b Dryopteris juxtaposita, TR 4590: photograph and drawing of meiosis from an 8-celled sporangium showing 123 bivalents (in black). Figs 32a, b Dryopteris juxtaposita, TR 4590: photograph and drawing of meiosis from a 16-celled sporangium showing 8 trivalents (striped), 33 bivalents (in black), and 33 univalents (outlined). Figs 33a, b Dryopteris juxtaposita, CRFJ 9894: photograph and drawing of meiosis from a 16-celled sporangium showing c.123 univalents (outlined). a CYTOLOGICAL OBSERVATIONS ON DRYOPTERIS & POLYSTICHUM pay 3556 L ¢ Figs 34a,b Dryopteris nigropaleacea, CRFJ 6925: photograph and drawing of root tip mitosis showing 82 chromosomes. Figs 35a, b Dryopteris nigropaleacea, CRFJ 6933: photograph and drawing of meiosis showing 41 bivalents (in black). Figs 36a,b Dryopteris nigropaleacea, CRFJ 6990: photograph and drawing of root tip mitosis showing 82 chromosomes. 28 MARY GIBBY 8 Cee 4-5 Oh ate sf tn ELEY ane mete tN ) of ities } %y LS Seo) ee Soy ay &% wm a bared : i / Rae BPE ee NS ee ORV, APEC R SL AyEHIRY &. pera Sie a nein, JALAL V4 ag ER OR Liss d~ Pd TRS Fees Wily 2 Se oe “Sa ptvend ee ee j 37a 37b 38b ) Ww, \Zs x —l S. PERN, 77 pe ) \ =) iid ~ NN. er BSN Ay WZ A 39b Figs 37a, b_ Dryopteris stewartii, CRFJ 6332: photograph and drawing of root tip mitosis showing 123 chromosomes. Figs 38a, b Dryopteris stewartii, CRFJ 6804: photograph and drawing of meiosis from an 8-celled sporangium showing c. 123 bivalents (in black). Figs 39a, b Dryopteris stewartii, CRFJ 6970: photograph and drawing of root tip mitosis showing 123 chromosomes. 40b 41b Figs 40a, b Dryopteris fructuosa, CRFJ 9996: photograph and drawing of root tip mitosis showing 123 chromosomes. Ane ‘ Figs 41a, b;42a,b Dryopteris basisora, CRFJ 9986: photograph and drawing of root tip mitosis showing 123 chromosomes. MARY GIBBY 43b 44b bI7, ea AG 1% WI 12 ‘ 4 7] 45b Figs 43a, b Dryopteris pteridiiformis, CRFJ 10028: photograph and drawing of root tip mitosis showing 164 chromosomes. Figs 44a,b Dryopteris subimpressa, CRFJ 7217: photograph and drawing of root tip mitosis showing c.82 chromosomes. Figs 45a,b Dryopteris caroli-hopei, CRFJ 7062: photograph and drawing of root tip mitosis showing 82 chromosomes. CYTOLOGICAL OBSERVATIONS ON DRYOPTERIS & POLYSTICHUM 31 = els 46b 47b + 48b Figs 46a, b Dryopteris caroli-hopei, CRFJ 7159: photograph and drawing of root tip mitosis showing 82 chromosomes. Figs 47a, b_ Dryopteris porosa, CRFJ 9887: photograph and drawing of root tip mitosis showing c.82 chromosomes. Figs 48a, b Dryopteris subtriangularis, CRFJ 9897: photograph and drawing of meiosis from an 8-celled sporangium showing 123 bivalents (in black). S32 MARY GIBBY 49b 50D Figs 49a, b Dryopteris gymnosora, CRFJ 9939: photograph and drawing of root tip mitosis showing 123 chromosomes. Figs 50a,b Dryopteris integriloba, CRFJ 10065: photograph and drawing of root tip mitosis showing c. 120 chromosomes. Figs51a,b Dryopteris integriloba, CRFJ 10067: photograph and drawing of root tip mitosis showing c. 123 chromosomes. CYTOLOGICAL OBSERVATIONS ON DRYOPTERIS & POLYSTICHUM 33 \ SZ Vidas ee \\ \ Wid pes PN A ff \ oo = —_ aR == a8 rd | \ oe 1 Al o> ~ t= CA \» ~ 52b Pe Fi ~N Ve 2 7 as Ss ow A —_ - 7 pea -'|- ae ND eee NG Nas ag ot Yy et 4 SI oe “— ay BS A SS oe we of s\) ~~ eS) 4 Se — \ ee pe Pe Fig on Noa WA NYS AG Mid | ae | I4_ VASA Oy eke Figs 52a,b Dryopteris championii, CRFJ 10056: photograph and drawing of root tip mitosis showing 82 chromosomes. Figs 53a, b Dryopteris varia, CRFJ 10054: photograph and drawing of root tip mitosis showing 123 chromosomes. Figs 54a,b Dryopteris viridescens, CRFJ 9934: photograph and drawing of root tip mitosis showing c. 164 chromosomes. 34 _MARY GIBBY 55b 56b Figs 55a,b Dryopteris viridescens, CRFJ 10068: photograph and drawing of root tip mitosis showing 164 chromosomes. Figs 56a,b Dryopteris x flemingii, CRFJ 6968: photograph and drawing of root tip mitosis showing 164 chromosomes. CYTOLOGICAL OBSERVATIONS ON DRYOPTERIS & POLYSTICHUM 6) co fal ©) & ° 58b a ry —*% s : i v2 a nae , 59a &o 59b ¥... : .. , feo 7 . ; Figs 57-59 Dryopteris x wechteriana, CRFJ 6948: photograph and drawing of meiosis showing: 57a, b: 4 bivalents (in black) and 74 univalents (outlined); 58a, b: 2 bivalents (in black) and 78 univalents (outlined); 59a, b: 8 bivalents (in black) and 66 univalents. 36 MARY GIBBY 60b wal mi fe A eG) —) ) | Z Z) ra ‘A \ ee Ah Zp bf. te. , y, \ pro N Md, 47 Gd ars Ja ag te 4 i _ ins au PS (\* at 2 2 “ —t /-* ca i je ee ge) xy 1.3 2 oe Naty 62a 62b Figs 60a,b Dryopteris x zygo-parentalis, CRFJ 8577: photograph and drawing of root tip mitosis showing c. 200 chromosomes. Figs 6la,b Polystichum acutidens, CRFJ 9945: photograph and drawing of root tip mitosis showing c.164 chromosomes. Figs 62a,b Polystichum acutidens, CRFJ 9947: photograph and drawing of root tip mitosis showing 164 chromosomes. CYTOLOGICAL OBSERVATIONS ON DRYOPTERIS & POLYSTICHUM 37 63 b Figs 63a, b Polystichum discretum, CRFJ 6689: photograph and drawing of root tip mitosis showing 82 chromosomes. Figs 64a, b Polystichum piceo-paleaceum, CRFJ 7179: photograph and drawing of root tip mitosis showing 164 chromosomes. Figs 65a, b Polystichum lentum, CRFJ 7153: photograph and drawing of root tip mitosis showing 82 chromosomes. 38 MARY GIBBY w 66b ou, 67b / se mee 2 u “IN 68b ~ ~ >, | S , > Six / vo Ll = igh ¢ = Vays ad a Ay | _ & on ner J ws lee 7 i a & a Maths 7 ! j a gs 7\. “ SS / I \ = Wis £ : 7 ghey Z ie ok ‘ = = / — - s | S v\l - f-\ _ \ Ads oo By Oe, ae “7 ” 4 Jew ff oe ows “cl sty . 69a Ske 69b Figs 66a, b; 67a, b_ Polystichum lentum, CRFJ 7155: photograph and drawing of meiosis showing 41 bivalents (in black). Figs 68a, b ? Polystichum pseudo-makinoi, CRFJ 9929: photograph and drawing of root tip mitosis showing 82 chromosomes. Figs 69a, b Polystichum xiphophyllum, CRFJ 9921: photograph and drawing of root tip mitosis showing 123 chromosomes. CYTOLOGICAL OBSERVATIONS ON DRYOPTERIS & POLYSTICHUM AY aN f. 7 ’ t- < ae r a S Ww AN ARV _—— \ ) 4 NS = = ae < ar J 7!) a \ Z 7 ZI sit (x fT ine ." a. : XN ees - Bo ae TANS > ee aes ' ee \ee ‘ 3 wt ~ wT * \ “A ac * 5 tae A re a - oo ; Papas 2 a ee ie &, ¢ ae ’0.0 12:0 39 Figs 70a, b Polystichum nepalense, CRFJ 6792: photograph and drawing of meiosis showing 41 bivalents (in black). Figs 71a, b Polystichum tsus-simense, CRFJ 9940: photograph and drawing of root tip mitosis showing 123 chromosomes. Figs 72a, b Polystichum tsus-simense, CRFJ 9999: photograph and drawing of root tip mitosis showing 123 chromosomes. 40 MARY GIBBY Nt ie \ al as = Gar) ag Ree aN ‘ pe : di + oe 7 c Xe / Be we Q) Se aa See us lh |! ie vJ - lane J Tay gS ~ / Y ) 3b Figs 73a,b Polystichum yunnanense, CRFJ 9998: photograph and drawing of root tip mitosis showing 164 chromosomes. CYTOLOGICAL OBSERVATIONS ON DRYOPTERIS & POLYSTICHUM 41 74b 73.0 76b Figs 74a, b Ctenitis mariformis, CRFJ 9870: photograph and drawing of root tip mitosis showing 82 chromosomes. Figs 75a, b Hypodematium crenatum agg., CRFJ 10015: photograph and drawing of root tip mitosis showing 164 chromosomes. Figs 76a, b Aleuritopteris albo-marginata, CRFJ 6959: photograph and drawing of root tip mitosis showing 58 chromosomes. : ; 78b Figs 77a, b Aleuritopteris albo-marginata, CRFJ 6959: photograph and drawing of root tip mitosis showing 58 chromosomes. Figs 78a, b Neocheiropteris palmato-pedata, CRFJ 10017: photograph and drawing of root tip mitosis showing c.70 chromosomes. British Museum (Natural History) Ferns of Jamaica A guide to the Pteridophytes G. R. Proctor This flora records and describes the 579 species and 30 varieties of ferns occurring in Jamaica. The succinct species descriptions include relevant synonymy and incorpo- rate distributional data both within and outside Jamaica. Special emphasis is given to the subtle distinctions between closely related species and all genera are illustrated. Keys to the genera and species facilitate a wider use of the flora in the West Indies and northern South America. The author, one time Senior Botanist in charge of the Herbarium of the Science Museum, Kingston, Jamaica, is an outstanding field botanist and his expertise is reflected in the practicality of the flora and especially in the habitat and ecological information. This volume represents an important addition to our knowledge of the flora of the West Indies. 1985, 631pp, 135 line illustrations, 22 maps. Hardback. 0 565 00895 1 £50.00 Titles to be published in Volume 14 Cytological observations on Indian subcontinent and Chinese Dryopteris and Polystichum (Pteridophyta: Dryopteridaceae) By Mary Gibby A redisposition of the species referred to the ascomycete genus Microthelia By David L. Hawksworth A classification of the genus Dryopteris (Pteridophyta: Dryopteridaceae) By Christopher R. Fraser-Jenkins Evolutionary cladistics of marattialean ferns By Christopher R. Hill & Josephine M. Camus Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk Printed in Great Britain by Henry Ling Ltd, Dorchester yr kD Nee Fe a Bulletin of the British Museum (Natural History) _A redisposition of the species referred to the ascomycete genus Microthelia David L. Hawksworth Botany series Vol 14 No 2 19 December 1985 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the Museum, both by the scientific staff of the Museum and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself, available separately, and individually priced. Volumes contain about 300 pages and several volumes may appear within a calendar year. Subscriptions may be placed for one or more of the series on either an Annual or Per Volume basis. Prices vary according to the contents of the individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History), Cromwell Road, London SW7 5BD, England. World List abbreviation: Bull. Br. Mus. nat. Hist. (Bot.) © Trustees of the British Museum (Natural History), 1985 The Botany series is edited in the Museum’s Department of Botany Keeper of Botany: Mr J. F. M. Cannon Editor of Bulletin: Mr J. R. Laundon Assistant Editor: Dr A. J. Harrington Editor’s Assistant: Miss M. J. Short ISBN 0 565 08008 3 ISSN 0068-2292 Botany series Vol 14 No 2 pp 43-181 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 19 December 1985 A redisposition of the species referred to the* ascomycete genus Microthelia David L. Hawksworth Commonwealth Mycological Institute, Ferry Lane, Kew, Richmond, Surrey TW9 SAF 17 Mi Contents ie rs EMME AEG LE Sai S425 SpA ub Sieh W Vian sih ck4s05ctes Lev ancescadTecssdesscavesssxdaeceaccsereed » 4 SN ccs ois yp \indk c nian cos sacoadedskisisaninsssooveseonyevabisions sovseeendici } . 44PRESE NSE Bd od hoi vfyh ok ciSxvusas skeen vesscisceesssenetooscsesieidoidsvestades 1. GEAIERA! hg ere aaa tea ccd y isu secasnaeed sex ks PeFE Mein RI 2 35 cs 9) Shing Sassi voc veaVs'caigacasaen Givi Gatudessinky caverexeibeseatacastes 47 aia oa Lied tn Ws ban V visas ¢adces oiisnd das b04 Gbnje vodenwnsictiasesiedsesbes 47 aL 3k. ca5 60) 9040 3550404 0263 adae o ov edxdis bvcsaee se voaee ha Wekcunabuvausdsvenensedeue 47 EN so ce oc a Abdi ys Dac a8s saacduscniaqlerssadseesederereveeveeccsssinicebinewsses 47 IE 85 Le dco cso g cak cost Goeetxissaae0009.054esesich¥aas es cbueecekovinesCéedudenvaiens’ 50 I ROE Soc co5 fcc Ric shskepeXh bed voce cuscseabobe tds adecbnievedudernsscaeeceres 52 GET 68 regs Shion st csak cy so db sed inas ocdshdaghersccvetsuesdsbagussscasasareuneraas nee 62 ee AB ian nce okanss pe27) 1900s cosbskas oheulwdvtoncbodesevddeuswadesdbadeacbesieccassiis 62 nod icy 505 eon a Causa dn sas Soc aua Zeek Li eeSGacs hao co ee Rasen 64 nes AS cs ns C0 ele sleeve AmeaA edb ap he Cosen ea ue ece cuKe ete te aaeee tow ceateh 64 NE aah pcnscekenncvas oseawens Sottssa dad lade daveeh vddseeheivadsera lever teneen: 65 Be sal b racy Colas 5 heats ce are eileen slat anon thec eevias bpaevook Fake sideaceprcsecaaiouee 67 eG 05 0555.4 0k 456.64 ohh nis We ec ann indenie deve Wsxnas¥y tdavcrnee res soewoladsddoedtess 120 I Puce coass 554 ss4asyacuiaecdssrsassiesaawcessasesuvgqunescceeerssecanoccsonayes 121 EME olf 3 a chen koveveay cdcansstcoai shard vodevescusenca Ceooweer mater oheneekciasens 121 NL Ios danse cues shndnad cake sav eas U809R Soe nkaninsd eakuuabaaniess ven A easoeianetsess 130 IIL 55 Ala 5. resin aisialesixpin tins ssisceties oevntabovordhteesoesusa vadedressesiacknaesets 174 EIEN a5 98s Svcs pa pbcate ae sav 0 Ns duvceacaaacened huntsanetasbeds duces dai es cadene> 174 IRA TASS os ie (3.5 Sais vad ada ou do¥n vw bai'es sonessencwes dos sdebands Uetecdn ct ied vencdesieaesene 177 Synopsis ~ Following a discussion of the usage of the name Microthelia K6rber, generic concepts to be applied to the lichen-forming and non-lichenized species referred to it are reviewed. A key to 15 genera is provided and notes on 35 of the generic names used for bitunicate pyrenomycetes with brown 1-septate ascospores are presented. Detailed treatments are provided for the genera Mycomicrothelia Keissler (Arthopyreniaceae; 26 species) and Peridiothelia D. Hawksw. gen. nov. (Phaeosphaeriaceae; 3 species). Two new species of Mycomicrothelia (M. atlanticaD. Hawksw. & Coppins, M. confusaD. Hawksw.) are described and the following new combinations made: M. anonacea (Mill. Arg.), M. apposita (Nyl.), M. captiosa (Krempelh.), M. confluens (Mill. Arg.), M. conothelena (Nyl.), M. dothideaspora (Cooke & Harkn.), M. fumulosa (Zahlbr.), M. miculiformis (Nyl. ex Mill. Arg.), M. minutula (Zahlbr.), M. modesta (Mill. Arg.), M. nonensis (Stirton), M. obovata (Stirton), M. pachnea (K6rber), M. socialis (Zahlbr.), M. subfallens (Mill. Arg.), M. thelena (Ach.), and M. willeyana (Mill. Arg.) D. Hawksw. combs nov. The predominantly tropical genus includes a mixture of doubtfully or facultatively lichenized and non-lichenized species; the latter are mostly host-specific. poems includes three non-lichenized European species: P. fuliguncta (Norman), P. grandiuscula (Anzi), and P. oleae (Kérber) D. Hawksw. combs nov. The identities of 144 epithets described in or referred to Microthelia are reviewed as excluded species and the following new taxa introduced: Arthopyrenia clavaeformis (Stirton), A. subgregans (Mill. Arg.), Didymosphaeria calyciospora (Massal.), Massarina ribesiella (Nyl. ex Vainio), Melanomma oxysporum (Zahlbr.), Muellerella vesicularia (Lindsay), Pleurotrema uniserialis (Zahlbr.), Polycoccum cartilagino- sum (Arnold), P. innatum (Mill. Arg.), P. umbilicariae (Lindsay), P. vermicularium (Lindsay), P. versisporum (Bagl. & Car.), Polypyrenula gen. nov. (Pyrenulaceae), P. sexlocularis (Mill. Arg.) D. Bull. Br. Mus. nat. Hist. (Bot.) 14 (2): 43-181 Issued 19 December 1985 43 44 DAVID L. HAWKSWORTH Hawksw. combs nov. Descriptions and illustrations (photographs and spore outlines) are provided for all accepted species of Mycomicrothelia and Peridiothelia and also of many of the taxa referred to other genera. This paper concludes the first phase of a re-investigation of all bitunicate pyrenomycetes with paraphysate ascomata and brown one-septate ascospores. Introduction The systematics of the fissitunicate paraphysate pyrenomycetes with brown 1-septate ascospores has been badly neglecged in recent decades, with the result that taxa continue to be referred to entities based on outdated generic concepts, not indicative of either their systematic rela- tionships or biology. With perhaps about 70 generic names and in excess of 1000 species epithets already in the literature this neglect is understandable, but unless inroads into this morass of names are made the existing confusions can only be compounded. This contribution aims to make some progress in improving this situation by providing a revision of all the taxa referred to a particular generic name in the group, Microthelia Kérber. This name was selected as preliminary studies showed that it had come to be used in a variety of senses, even by contemporary authors, and was becoming a repository for a bewildering array of both lichen-forming and non-lichenized fungi. A critical study of these taxa also necessitates a re-appraisal of the criteria used for the separation of genera and species; this should prove of value, through extrapolation, in allied groups. It would have been ideal to treat all taxa referred to Didymosphaeria Fuckel at the same time as those placed in Microthelia, but so many names are involved that the task could not be completed for many years. However, some epithets in Didymosphaeria may be expected to belong to genera or even species accepted in the present revision, rather than to Didymo- sphaeria itself (see p. 54). This paper can consequently also be viewed as a first contribution to a re-appraisal of taxa referred to Didymosphaeria. Further, epithets placed under both these generic names by different authors are considered. There has been a tendency for students of pleosporaceous fungi to develop broad generic concepts in the temperate zone and then later attempt to accommodate tropical taxa within their limits. This approach has already led to considerable difficulties in some groups, for example the Leptosphaeria- and Pleospora-like fungi. The approach adopted here, a re-appraisal of the described world species prior to the formulation of generic concepts, should tend to reduce the risk of a similar situation arising in the Didymosphaeria- and Microthelia-like taxa, although the under-collecting of both lichen-forming and non-lichenized pyrenomycetes in the tropics means that the danger cannot be eliminated. The number of epithets which have been referred to Microthelia or genera which have been considered as synonyms of it is 200. I have presented the results of my studies of three ‘segregates’ elsewhere, as they proved to be remote from the bulk of the taxa referred to Microthelia. These are Astrosphaeriella (Hawksworth, 1981a), Kirschsteiniothelia (Hawks- worth, 1985a), and Lichenothelia (Hawksworth, 1981b). This paper is primarily concerned with general aspects of the group, the genera Mycomicrothelia and Peridiothelia, and the 144 names used in or referred to Microthelia which belong elsewhere. These names are treated in the section on excluded species, which also indexes relevant epithets discussed in Hawksworth (1981a, 1981b, 1985a). History of usage The generic name Microthelia (Gk wyod6s, micro — small; dnd%, thele — wart or nipple) was first introduced by Kérber (1855: 372) to accommodate perithecial taxa producing brown one- septate ascospores. He included four species in the genus, M. micula Flotow ex K6rber, M. atomaria (Ach.) Kérber, M. propinqua Koérber, and M. pygmaea (K6rber) Koérber. The typification and nomenclature of these is considered under ‘Excluded species’ in this treatment but it is pertinent to mention their identities here: (1) M. micula = Anisomeridium biforme (Borrer) R. C. Harris (see p. 157), M. atomaria = Leptorhaphis atomaria (Ach.) Szat. (p. 135), SPECIES REFERRED TO MICROTHELIA 45 (3) M. propinqua = Endococcus propinquus (K6rber) D. Hawksw. (p. 160), and (4) M. pygmaea = Muellerella pygmaea (K6rber) D. Hawksw. (p. 160). The principal objective of KOorber in introducing the name Microthelia appears to have been to replace Tichothecium Flotow (see p. 59), which he considered too similar to the name of the hyphomycete genus Trichothecium Link (Link, 1809: 18); as the type species of Flotow’s name is T. nigrescens (Pers.) Flotow (i.e. Verrucaria nigrescens Pers.; see Hawksworth, 1979a) this interpretation renders Microthelia an obligate later synonym of Verrucaria Schrader nom. cons. Unfortunately the nomenclatural position here is complicated by K6érber (1855: 341) having retained V. nigrescens in Verrucaria as a synonym of V. fuscoatra (Wallr.) K6rber. However, most authors since Fries (1861: 111) have unquestioningly cited M. micula as the type species of Kérber’s generic name, misapplying it to the taxon named as Peridiothelia fuliguncta (Norman) D. Hawksw. in this revision, and not appreciating that the epithet micula was intended as a replacement for Borrer’s ‘biformis’ (see p. 157). Harris (1973: 26) first pointed out that ‘micula’ could be treated as superfluous but retained both genus and species names as an interim measure. To select M. propinqua or M. pygmaea could threaten Endococcus Nyl. or Muellerella Hepp ex Miill. Arg. respectively. The nomenclatural situation was considered further by Hawksworth & Sherwood (1981: 339-340) who concluded that it was most appropri- ate to use the conservation procedures and proposed Anisomeridium (Mill. Arg.) M. Choisy for conservation over Microthelia, with M. micula nom. illegit. as the lectotype of the later name. Whatever the outcome of this proposal, K6rber’s name will continue to be unavailable for any of the genera recognized in this present revision. The heterogeneity of K6érber’s concept of Microthelia was quickly recognized by his contem- poraries. Massalongo (1856: 57-8) restricted the genus to three species which he considered (incorrectly) to be lichenized: ‘M. biformis’, M. atomaria, and the newly described M. macularis Hampe ex Massal. (i.e. Mycomicrothelia macularis (Hampe ex Massal.) Keissler; p. 92), referring the lichenicolous species to Tichothecium. The name was not accepted by Nylander (1858: 60) in his synopsis of the pyrenocarpous lichens, M. micula being placed as synonym of Verrucaria cinerella Flotow (i.e. Mycomicrothelia melanospora (Hepp) D. Hawksw.; p. 95). Trevisan (1860: 10) attributed the genus to ‘K6rb. (exclus. spec. parasit. — 1856)’ following Massalongo, and excluding the lichenicolous taxa. Trevisan accepted seven corticolous (includ- ing both M. micula and M. atomaria) and one saxicolous species. K6rber (1865: 396-399), however, maintained a broad concept of the genus admitting six species additional to his 1855 treatment; these included a wide range of fungi with + perithecial ascomata and brown one-septate ascospores including lichen-forming, non-lichenized, and lichenicolous species. He harshly rejected Massalongo’s (1856) interpretation, which he viewed as illogical in splitting species out on a biological basis. A remarkable usage of Microthelia is seen in the works of Lindsay (e.g. 1869: 519) who provisionally adopted the name as a ‘pseudo-genus’ for a wide range of lichenicolous fungi ‘confessedly most diverse in character’. Taxa with simple, 1-septate, or 3-septate, brown or colourless spores were included, drawn from a wide range of groups, even including some Coelomycetes. Fuckel (1870: 140-142) introduced a new genus, Didymosphaeria Fuckel, for fungi occurring on lichens or phanerogams which recalled Pleospora but had brown one-septate ascospores. Rehm (1879) recognized that this posed problems and endeavoured to restrict Didymosphaeria to non-lichenized and Microthelia to lichen-forming species; six species were referred to the latter group, but none of these are now considered as definitely lichenized. Stein (1879: 331), however, had a different view separating Tichothecium from Microthelia on the basis of habitat, restricting the latter to species on bark. Niessl (1881) rejected Rehm’s interpretation also, drawing attention to differences in ascomatal structure and comparing Microthelia to Massar- iopsis Niessl (i.e. Amphisphaeria Ces. & de Not.; see Miller & von Arx, 1973: 120). However, a more superficial view was adopted by Saccardo (1882: 709) who treated Microthelia as a synonym of Didymosphaeria. The first author to endeavour to deal with tropical Microthelia-like taxa in detail was Miller Argoviensis (1885: 416-7). He recognized two groups in material from Cuba he attributed to the 46 DAVID L. HAWKSWORTH genus, one with ‘Perithecia dimidiata’ and the other ‘Perithecia globosa aut subglobosa’. Later he extended the concept of the genus to include a species with (3—)5 septate spores, M. sexlocularis Mill. Arg. (Miller Argoviensis, 1888: 38); Clements (1909: 173) placed the latter in anew monotypic genus, Polythelis Clem. (see p. 165). Jatta (1911), in providing a treatment of the Italian species, recognized four sections in the genus based on the numbers of spores in the asci and whether the cells in the spores were equal or unequal in size. Vainio (1921a), however, formally recognized the two groups of Miiller Argoviensis (1885), and introduced the sectional name Holothelia Vainio for that with entire ascomata and Hemithelia Vainio for the one with dimidiate ascomata. In his careful re-evaluation of the Finnish lichenized pyrenomycetes, Vainio (1921b: 144) used ‘Microthelia (Koerb.) Vain.’ for the single lichenized maritime species M. scopularia (Nyl.) Blomb. & Forss., though noting that M. metzleri Lahm was also close to this species. In character with his treatment of other groups, Vainio did not retain non-lichenized species in a genus he interpreted as lichenized. Thus, he took up Didymosphaeria Fuckel for the consistently non-lichenized corticolous species (‘D. micula (Flotow) Vainio’, D. analeptoides (Bagl. & Car.) Vainio, D. ribesiella (Nyl.) Vainio, D. wallrothii (Hepp) Sacc. & Trott., and a few non-Finnish species treated cursorily). This interpretation was rejected.by Zahlbruckner (1926a: 75), however, who continued to follow a broad concept of the genus — at the same time misinterpret- ing Vainio’s sectional names. Keissler (1936: 21-57) utilized Vainio’s sectional classification and, in the first attempt at a critical re-appraisal of the central European species, recognized eight species in the genus, viz. ‘M. micula’ (with f. grandiuscula (Anzi) Keissler), M. pachnea K6rber, ‘M. atomaria’ (with f. calyciospora (Massal.) Keissler), M. macularis Massal. (with f. ribesiella (Nyl.) Keissler), M. oleae K6rber, M. betulina Lahm, M. aterrima (Anzi) Zahlbr. (with f. atramentea (Norman) Keissler, var. dermatinoides Servit), and M: marmorata Hepp ex K6rber (with f. cartilaginosa (Arnold) Keissler, f. minor (Kernst.) Keissler). Keissler simultaneously proposed the new genus Mycomicrothelia Keissler for two non-lichenized species with dimidiate ascomata accepted in his treatment, ‘M. atomaria’ and M. macularis. In introducing this generic name he separated it from Didymosphaeria on the basis of the dimidiate rather than entire ascomata. In addition, Keissler provided notes on the identities of 38 excluded or imperfectly known species, the types of some of which he had studied. The next significant contribution to our knowledge of the Didymosphaeria-like fungi was that of Scheinpflug (1958) who recognized four groups within the genus. He did not discuss Microthelia itself, but placed M. micula as a synonym of a species he referred to as Astrophaeriel- la pinicola (Rehm) Scheinpflug (i.e. Kirschsteiniothelia aethiops (Berk. & Curtis) D. Hawksw.; see p. 50, Hawksworth, 1985a)). Miiller & von Arx (1962: 282-284) realized that this situation was not satisfactory, and re-instated Microthelia, placing Astrosphaeriella Sydow & H. Sydow, Jahnula Kirschst., and Kirschsteiniella Petrak as synonyms. Didymosphaeria was used for species with entire ascomata, Microthelia for ones which were hemispherical and superficial, while Mycomicrothelia was employed for immersed dimidiate ascomata. While this was the first time that the justification of separating Keissler’s genus Mycomicrothelia on structural rather than biological grounds had been appreciated, the broad view of Microthelia adopted placed together a disparate assemblage of six species (five newly combined into the genus) which subsequent more detailed studies have shown must be dispersed between four genera referred to different families of Dothideales. Poelt (1969: 394-397) did not follow Miller & von Arx (1962) but retained Keissler’s (1936) broad concept of the genus. He provided keys to 13 species accepted as occurring in central Europe, arranged according to Vainio’s subgenera. The importance of the reaction of the centrum with iodine in separating the dimidiate taxa was stressed, and species groups broadly equivalent to Peridiothelia and Mycomicrothelia as accepted in this revision were distinguished. Subsequently those working with lichens have tended to follow Poelt (e.g. Harris, 1973), while other mycologists have adhered to the schemes of either Scheinpflug (e.g. Dennis, 1968), or Miiller & von Arx (e.g. Kohlmeyer & Kohlmeyer, 1979: 429-431). It is also pertinent to draw attention here to the use of Microthelia for the teleomorph (perfect SPECIES REFERRED TO MICROTHELIA 47 state) of the dematiaceous hyphomycete Dendryphiopsis atra (Corda) S. Hughes by Corlett & Hughes (Hughes, 1978: 360-362); this fungus is now referred to Kirschsteiniothelia (Hawks- worth, 1985a). In order to clarify this confused situation my own investigations on the taxa referred to Microthelia were initiated in 1979 (Hawksworth, 1981a, 1981b, 1984, 1985a; Hawksworth & Sherwood, 1981). The present contribution completes this first phase of my investigations on the Didymosphaeria-like fungi. Material and methods Material In the course of this study, collections have been received on loan or studied in the following institutional herbaria: BG, BM, CAN(L), DAOM, E, G, H, IMI, K, L, LPS, M, NY, O, PAD, PC, S, TNS, TRH, TUR, UPS, VER, W, WELT, WRSL, and ZT (abbreviations follow Holmgren et al., 1981), and the private herbarium including H. Fabre’s material at L’;Harmas. Methods Macroscopic features were studied with a Nikon stereomicroscope fitted with a measuring eyepiece graticule and zoom lens (to x 60). Microscopic characters were investigated with an Olympus BH or BH-2 microscope equipped with Nomarski Differential Interference Contrast (DIC) optics, a drawing tube, and camera. Magnifications up to X 1562 were used for measurement with an eyepiece graticule, and drawings were routinely made at xX 3300. Macroscopic photographs were taken on a Zeiss Tessovar system with zoom lenses, and those of sections and microscopic features in the Olympus microscopes used. For each specimen 1-2 of the larger ascomata were removed and vertical sections cut at a thickness of 10 wm with an LBH freezing microtome. These sections were mounted in lactophenol-cotton blue (LCB), warmed, and sealed with Glyceel when cold for the examina- tion of ascomatal structre; most sections preserved in the course of this study have been preserved in IMI. In addition ascomata were cut open with the aid of a surgical scalpel blade, and rough hand-cut sections mounted in 10% potassium hydroxide or Lugol’s iodine solution. Where no reaction with Lugol’s solution was obtained, specimens were pre-treated with potassium hydroxide and a mount also made in Melzer’s iodine solution to check negatives. Measurements of ascospores were made on spores not included within asci (although many of these would have been released in the mounting process). Wherever the material permitted, at least 10 ascospores were measured and drawn using the drawing tube fitted to the microscope. Where pycnidia were being investigated, erythrosin in ammonia was used as a mountant to show details of the conidiogenous cells. Scanning electron micrographs were taken on the ISI-60 SEM at CMI using air-dried specimens mounted on stubs with double-sided Sellotape and coated with gold. Generic concepts Taxonomy As will be evident from the above review of the history of the usage of the name of Microthelia (pp. 44-47), generic concepts in the fungi to which this name has been applied have not previously been critically revised. Features such as ascospore colour and septation which had been stressed as generic criteria by most mycologists and lichenologists in the period 1880-1940 led to heterogeneous assemblages of often rather distantly related species being included under a Single generic name (e.g. Karschia K6rber, see Hafellner, 1979), and closely related species being dispersed amongst several generic names (e.g. Micarea Fr., see Coppins, 1983). The situation in Microthelia was found to be similar to that formerly existing in Karschia, but is probably not atypical of most pyrenocarpous genera, including lichen-forming species which are predominantly tropical. 48 DAVID L. HAWKSWORTH MH Fig. 1 A, Paraphysoids (trabeculate pseudoparaphyses). B, Pseudoparaphyses (cellular pseudoparaph- yses). The ontogeny of the ascoma and structure of the ascus are now generally accepted as of principal importance in the classification of the Ascomycotina (Hawksworth, 19855). In the case of the main genera accepted in this revision, the ascus structure is typically bitunicate and discharge almost certainly fissitunicate. However, there is considerable variation in the structure of ascomata produced, and it is this character which proves to be of major importance for the recognition of genera here. The nature of the interascal tissue, or rather ‘hamathecium’ (Eriksson, 1981), present in the ascomata is related to its ontogeny. In one group of species this was found to consist of paraphysoids (trabeculate pseudoparaphyses) attached at the top and base of the ascomatal cavity; these are produced by a stretching of the former interascal cells as the ascomata mature and finally appear only 1-1-5 um thick, very rarely septate, and delicately branched and anastomosed (Figs la, 4c). The species with this type of hamathecium had carbonaceous ascomatal walls, often almost hyaline, and in several species multiseptate ascospores; all occurred on palms, bamboos or other stout grasses in the tropics and are non-lichenized. This assemblage of species is referred to Astrosphaeriella Sydow & H. Sydow (Hawksworth, 1981a; Figs 2a, 4). Astrosphaeriella belongs to the family Melanommataceae Winter, placed in the Melanommatales by Barr (1979a) and the Dothideales by Eriksson (1982). The ontogeny of the main saxicolous group of species referred to Microthelia was found to be most distinctive by Henssen & Jahns (1973: 113 fig. 4.18 k—-m) in that the ascomata are disc-like and superficial from a very early stage (Fig. 6B) with the asci finally arranged almost palisade- like in a hymenium separated by paraphyses-like branching filaments firmly attached at the base (Fig. 2D). In the other main groups, discussed below, the asci remain enclosed in a perithecium- like ascoma. The two species recognized are also of interest in that they have + smooth-walled ascospores with gelatinous sheaths swelling markedly in potassium hydroxide and spores which may become 3-septate; one has asci in which the outer wall turns blue in iodine (Fig. 6C), and they are biologically similar in forming a black superficial crustose ‘thallus’ but may be more appropriately regarded as algal parasites than as truly lichenized. The new generic name Lichenothelia D. Hawksw. was introduced for these species (Hawksworth, 1981b), the family placement of which within the Dothideales remains uncertain. Mycomicrothelia is distinguished from the other two genera recognized with cellular pseudo- paraphyses (Fig. 1B) and ascomata opening by a small circular ostiole by the development of the generative tissue beneath a clypeate involucrellum; this covering layer is made up of cells of the outer peridermal or epidermal layers intermixed and bound together by brown hyphae which form a strong covering tissue (Fig. 3C). In some cases the hyphae extend to form a conspicuous fringe extending beyond the involucrellum (e.g. M. melanospora, M. wallrothii; Fig. 2F), in which case the generative locule is often surrounded by a clearly differentiated thin exciple, while in others the hyphae are restricted to the area immediately surrounding the generative locule with no clearly differentiated exciple (e.g. M. confusa, M. thelena; Fig. 2G). For some SPECIES REFERRED TO MICROTHELIA 49 cs rf, F Fig. 2 Ascoma types of the principal genera included in the concept of Microthelia by previous authors (diagrammatic). A, Astrosphaeriella. B, Jahnula. C, Kirschsteiniothelia. D, Lichenothelia. E, Peridio- thelia. F, Mycomicrothelia (M. macularis type). G, Mycomicrothelia (M. thelena type). Not to scale. Substratum and(or) host tissue indicated by horizontal parallel rules. time I considered separating these two groups at the generic level but some species (e.g. M. socialis) are rather intermediate in structure and a broader generic concept is consequently more appropriate. The species grouped in this way are probably mainly non-lichenized and restricted to particular hosts, but some do seem to have trentepohlioid algae associated with them when fresh. This situation is strongly reminiscent of Arthopyrenia s.str. (Harris, 1975) and I endorse the view of Barr (1979a) that Mycomicrothelia should be referred to the family Arthopyre- niaceae W. Watson. Mycomicrothelia is very close to Arthopyrenia s.str. and is distinguished from it only in that the ascospores become brown while still within the asci and have a verruculose ornamentation. This affinity is further supported by the dimidiate form of the ascomata in almost all species and also by the presence of pycnidia with bacillariform conidia in 50 DAVID L. HAWKSWORTH Fig. 3 Tissue types in ascoma walls. A, Hyphal (textura intricata). B, Cellular (pseudoparenchymatous; textura globulosa or textura angularis). C, Clypeate (host cells, dotted, bound together and interspersed by fungal hyphae). several Mycomicrothelia species. Further, old ascospores within the ascomata of Arthopyrenia species may become brownish as they degenerate; it is not surprising therefore that some names included within Microthelia must be referred to Arthopyrenia (see ‘Excluded species’). Kirschsteiniothelia (Figs 2C, 6A) and Peridiothelia (Fig. 2E) both differ from Mycomicrothelia in that the ascomata are erumpent and finally + completely superficial. The ascomatal walls consist entirely of fungal tissue and do not have host cells as an integral part of their composition. In Kirschsteiniothelia the ascomatal wall is clearly pseudoparenchymatous with rather angular cells (Fig. 3B; textura angularis to textura prismatica) and a palisade of cells is often visible in the angles or at the base of the ascomata; in addition the ascomata become flattened at a late stage, the centrum is almost invariably unchanged with iodine, and the ascospores are often attenuated apically, with a thin gelatinous sheath. The six species accepted in this genus are essentially lignicolous saprophytes and where an anamorph is known this is a hyphomycete (Hawksworth, 1985a). In contrast, the ascomata of Peridiothelia have a wall consisting of small irregularly thickened rather rounded pseudoparenchymatous cells (textura globulosa), for the most part only becoming angular towards the base and especially in the angles of the ascomata (textura angularis); in addition the centrum tissues consistently turn blue in iodine and the ascospores have rounded apices and a conspicuous gelatinous sheath swelling in potassium hydroxide. Anamorphs are unknown in all the Peridiothelia species, which occur on the bark of living trees. Kirschsteiniothelia and Peridiothelia species do not appear to be lichen-forming. In the scheme of Barr (1979a), Kirschsteiniothelia would be referred to the family Pleosporaceae Nitschke as the ascomatal walls are + equal in thickness or thickened at the sides but further on the basis of the hyphomycetous anamorph, and Peridiothelia to the Phaeosphaeriaceae Barr as the ascomat- al walls are + equal in thickness or thickened above and no hyphomycetous anamorph is known. The main distinguishing features between the accepted genera are summarized in Table 1, and the differences in ascomatal structure are illustrated diagrammatically in Fig. 2. Key to the genera The following key includes the genera accepted in this revision (Table 1), together with other genera with ‘bitunicate’ asci and brown 1-septate ascospores which have been confused with these in the past. It is important to stress that this key does not purport to be a complete survey of genera with these characters. Many more taxa need to be studied before such a key can be produced. Further notes on the genera keyed out here are provided in the following section (pp. 52-62). “SUIPO!s [Oon'T - ‘aprxoipAy unissejod % OT , € 97 Z 9 I Or sajseds jo Joquiny] (sooxq (sdo1] snonploep (sooquieq snonplep yieq uo yoo! uo pure sioj1u0o pue sujed uo dso) y1eq onAydoides poziusyoly dso) poom (onenbe) poom ‘dso) o1ytsered uo onAydoides IO poziusyory] IO snojooIs;, Vy uo on Aydoides uo oy Aydoides Jo onAydoides ASoqorg (stsdoryddupuaq) umouUy o10yM UMOUY 919M umouyuy) snoj}a0AU0[907 uMouyuy.) snojooAmoydAF{ uMOoUuyUy-) umouyuy) ydioweuy snonoidsuos podojoaop-[[om pue podoyaaop Aj100g 0} AJayeIOpopy podo]aacp-]19M podoyaaop Aj100g podoyjaaap Aj100g podojaaap Aj100g yyeous (UMOIQ-pol (snosseal[o (rep (UMOIQ-pol (UMO1q-p2.) UMOIQ-pol 0} Ayyensn) umoig Ajyensn) umoig AI9A) UMOIG Ayyensn) umoig uMOIg Me Ijs ‘oulTeAyqns moog [enbo + IO Jo];euIs jenby + Ud}JO []20 IOMO'T jenby + jenby + jenbq jenbq sII2OD I (€-)I et T T oI eidag salodsoosy JoJOIA AJoIeI =—_ (anjq YJeoYs snose onyjq Ajore1 onig Jo posueyoug, Jo) posueyoup, Jo posueyouy) posueyouy posueyouy (-]) wn.qUaD sosAyderedopnesd soskyderedopnesg = saskyderedopnesg oyl-stshydeseg soshydeiedopnesg sploskydese gy sploshydeieg 9 wmpoyjemeEy posueyouy snosoeAl|O posueyoun posueyoug posueyouy) posueyouy, § (, 9) INojoD (e}eOLUI (eoneusiid (sluejnsue B1N}X9}) (eoneusid Jo suepnsue Jo eso[nqgoys o]dioxe yeydAy = Jo siiejn3ue (suepnsue B1N}X9}) BIN}X9} ‘]]eUIS ‘WIN][OION[OAUI B1N}X9} ‘WINIpoul B1IN}X9} SOBIE] Ie[NyI[I9 S][99) Ie[NIJID jeyddy oje0dATD jeydAyy S][99) Ie[N]TED S[[99) Tena: puw snosdeUOQIeD eM snonuljuoo o1nUue F yng Ajyensn aseq 9) RIPIuUIp pous}jey uso o1nue o1RIPUIp + nq oepruip F ‘WuoyMmweWw o1nuS eseq ‘wo0jiAd pue popunol Jo pous}jey F ‘WIopTMUePy Ajores ‘oyeuelddy oseq ‘O¥T]-OsIq 0} WIOFIWUWe Py oseq ‘WIOjLIAg aseq ‘[eoIUod odeys oseq juodunio agent ory £5) Sei Lees iag \ = ts ; Oo? ni epee a i ‘ = a ye, mt ’. “3 > Gains 7s 7 , ¢ y she! rr | 4, aes Ih Wee Gl Pes roe) Dh ye hae ~ 5, sae hoe git . if s dd aE , ' te 4 i Se et f am a f a, b* L * * Lf * =e rd British Museum (Natural History) Ferns of Jamaica A guide to the Pteridophytes G. R. Proctor This flora records and describes the 579 species and 30 varieties of ferns occurring in Jamaica. The succinct species descriptions include relevant synonymy and incorpo- rate distributional data both within and outside Jamaica. Special emphasis is given to the subtle distinctions between closely related species and all genera are illustrated. Keys to the genera and species facilitate a wider use of the flora in the West Indies and northern South America. The author, one time Senior Botanist in charge of the Herbarium of the Science Museum, Kingston, Jamaica, is an outstanding field botanist and his expertise is reflected in the practicality of the flora and especially in the habitat and ecological information. This volume represents an important addition to our knowledge of the flora of the West Indies. 1985, 631pp, 135 line illustrations, 22 maps. Hardback. 0 565 00895 1 £50.00 Titles to be published in Volume 14 Cytological observations on Indian subcontinent and Chinese Dryopteris and Polystichum (Pteridophyta: Dryopteridaceae) By Mary Gibby A redisposition of the species referred to the ascomycete genus Microthelia By David L. Hawksworth A classification of the genus Dryopteris (Pteridophyta: Dryopteridaceae) By Christopher R. Fraser-Jenkins Evolutionary cladistics of marattialean ferns By Christopher R. Hill & Josephine M. Camus Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk Printed in Great Britain by Henry Ling Ltd, Dorchester SSM 1Ol® Bulletin of the British Museum (Natural History) A classification of the genus Dryopteris (Pteridophyta: Dryopteridaceae) Christopher R. Fraser-Jenkins Botany series Vol 14 No 3 30 January 1986 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the Museum, both by the scientific staff of the Museum and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself, available separately, and individually priced. Volumes contain about 300 pages and several volumes may appear within a calendar year. Subscriptions may be placed for one or more of the series on either an Annual or Per Volume basis. Prices vary according to the contents of the individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History), Cromwell Road, London SW7 5BD, England. World List abbreviation: Bull. Br. Mus. nat. Hist. (Bot.) © Trustees of the British Museum (Natural History), 1986 The Botany series is edited in the Museum’s Department of Botany Keeper of Botany: Mr J. F. M. Cannon Editor of Bulletin: Mr J. R. Laundon Assistant Editor: Dr A. J. Harrington Editor’s Assistant: Miss M. J. Short ISBN 0 565 08009 1 ISSN 0068-2292 Botany series Vol 14 No 3 pp 183-218 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 30 January 1986 A classification of the genus Dryopteris (Pteridophyta: Dryopteridaceae) Christopher R. Fraser- Jenkins c/o Department of Botany, British Museum (Natural History), Cromwell Road, Stun Fk SW7 5BD Contents } GENERAL RT AR SC Tene aah chs vous band ides soveldd snsksvonadocncidusebesdacodaveaveasens 183 Higher classification and relationships of Dryopteris ...............cccccceceeceececeeceeeeeeenes 184 EM TAA AVIREEIR SIF VODICTES 5 600605 ccs lee ive ccdiscccctccvoeccvsceesecesdscssessevssoudeecscscenas 186 MIND, 00s cx WAN ek: da Ge Sedgs cab oas4K eae 06cadedee oss cveseradeccnvacssensscdddacsecdveed tens 187 MMMM ah oR ge ih a3 cvsag vag ip oad Uae cy'eg ja's bos b Dp nnd pbds onsababs eovsuseisasusas b4eb0 Fess 187 epmeruel. J ¥CHnOnlerts (P. MOOTE) CHING .c...02.scccceccsssccvscscvsesaevcceseoessccseaes 189 INE A ERNIE Ss a SN a sane ou snes vennvhsataeare asa ndiaveb bbs cayessdameareausesoes 189 RESET, Des REMPCMES E TASET “I CTKING 5.5.5 svcesescesatsesbapsesveseretedesdestesosieecutes 190 gee DO Sg 190 ETE FF AE T TANT SOPRIOS .. .saiscie sine deed cusnesdnsiac¥enededvisctaasoseenossesss 191 UNE IE UNION EE CAINS Lind ceSe\ yh y ds tse hde i dansasyadss veces steele tied iwecvereds vieed 191 OCHO (Ds -Bemmotde PrASCT-JENKINS «0... cis cece ceceseenecscssccdsovcsdecseascssacseess 192 A OF OMe FCOGES- SOURIS, 655i. cess cucetsieeoscetacenvsdacccstesscostsseoveces 192 OCtiothy 7: SUPNGENIES PIASET- JENKINS «000.666. .cisceceesescscsccsecesosceseacenscscces 193 Section 8, ‘Comamomeae Fraser-Jenkims. .............0.000scsescesescscessecscssesssess 193 pection- 9: Margingtae Fraser-Jenkins ........:..0..sscscesssscsiscssscscsesesesescssness 194 eRe CO, S RCIINEEE FTRDETSOREMIS «556505 scan scrccenisss sixenmaracssececsodedescscesauvs 194 Section 11. Lophodium (Newman) C. Chr. ex H. It6............... cece ee cec eee eee es 195 Subgenus 3. Erythrovariae (H. It6) Fraser-Jenkins .................0.ceececceeceeeneee ees 195 EY VTE OUT ENE liga isaol ees nko d tases nek bia TAs os ok ROUEN ORL 196 ERNE ee ORS FPOGET- FOTURING 50 oc ces cccesnetsessdcetvcuscereséveVendecvasceverteeetes 196 MEE oe CIPEE WTAGCTHIONMING £05 055s siccessses sccinaVesoutwedpeedsd cwacdeuysodevawscswes 196 Sabgenus 4. Nephrocystis (H. It6) Fraser-Jenkins ..................ccccccsssscscsecscoees 197 pecoon 1. Purpurascentes Fraser-Jenkins ................0000sccsscsseccscscnceescoscness 197 NI PEERY EMILE sg ch veep shots 8 enya dav elds oNia too ence PUT na tare eTes +a downs TORRES 198 Pee OMBOPND TAN TIRE WIASSINICATION -. «05.5000 ; Seq ty : pets ¢ pristopher R. fraser Jew pea Fig. 3 Dryopteris tingiensis Ching & S. K. Wu ex Fraser-Jenkins. Holotype (Chang Chung team 9525) (PE). 204 C. R. FRASER-JENKINS Tibet, from the single, immature specimen from Ting Tze which has immature spores. Although they have accepted other such specimens for publication as new species, often erroneously, they did not publish D. tingiensis along with their other Tibetan species. I agreed with their original recognition of it on seeing the specimen, and became reinforced in my view that it is a new species on studying the specimen that I have designated the type, which was unidentified in a new collection Professor Ching passed to me to examine while I was visiting him in Peking. It is named after Ting Tze county, Tibet, and is so far only known from SE. Tibet. 4. Dryopteris ruwenzoriensis C. Chr., sp. nov. Fig. 4 ‘Eudryopteris rhizomate virisimiliter obliquo; stipite valido, 60 cm vel ultra longo, brunneo-stramineo, paleis paucis (plerisque abrasis) luteo-brunneis, lanceolatis, integris, ad 2 cm longis, aliis multo minoribus filiformibus mixtis vestito, superne sulcato. Lamina ovata, magna (metrali?), 60 cm vel ultra lata, papyraceo-herbacea, laete viridi, tripinnata. Pinnis alternis, ad 10 cm inter se remotis, basalibus maximis, ad 35 cm longis, breviter petiolatis, 15 cm latis, latere basiscopico producto inaequilateralibus, acuminatis, sequentibus subaequilateralibus, subdeltoideo-oblongis, subsessilibus, costis sursum alatis; pinnulis II. ord. basiscopicis pinnarum basalium 11-12 cm longis 3 cm latis, pinnis primariis supremis similibus, acroscopicis iis pinnarum ceterum similibus, 4-5 cm longis 1-5-2 cm latis, sessilibus, e basi paulo aequali truncata latiore subgradatim in apicem acuminatum attenuatis omnibus alternis, 3-2 cm inter se remotis, ad costulam plus minusve late alatam profunde pinnatifidis vel majoribus versus basin pinnatis segmentis (vel pinnulis III. ord.) parum obliquis, lateribus parallelis apice truncato rectangularibus, 1-1-5 cm longis 0-5-0-7 mm latis ubique dentatis (dentibus brevibus, deltoideis, non aristatis), majoribus plus minusve profunde serrulato-laciniatis, laciniis subquadratis apice dentatis. Rachi, costis costulisque paleis tenuissi- mis parvis, lanceolatis, pallide luteo-brunneis sat sparse vestito. Venis in lobis pinnatis, plerisque furcatis, adscendentibus. Soris costularibus, sat magnis, globosis, indusiis non repertis.’ ‘This new species differs from the many African relatives by its large size, the broad, rectangular ultimate segments with many small teeth and in being (apparently) exindusiate. Small, early falling indusia may possibly be found in quite young sori, but in every case very different from the large indusia of D. Schimperiana (Hochst.) C. Chr., which comes nearest to our species, especially in scale-characters.’ Holotype: Herbarium Filicum Carl Christensen 5112, 22 Juli 1930, Mission H. Humbert-Plantes de l’Afrique Equatoriale — 1929, no. 8825 ter., ‘Dryopteris ruwenzoriensis nsp.’ [Christensen scripsit]. Massif du Ruwenzori, versant Ouest (Congo Belge, Ituri), Altitude vers 2000 m, Date de la récolte: Juillet (BM!). Paratypes: ditto nos. 8849 (P! BM!); 8900 bis (BM!); Mission Ch. Alluaud Afrique Orientale 1908-1909, no. 303, Ruwenzori (Est), hte. Vall. du Mobuku, montée du Col de Buamba, 3300-3400 m, Janv. 1909 (BM!). The above diagnosis and comments are typed on a sheet in a capsule mounted with the type specimen at the BM. From the lamina morphology alone it seemed to me that this was a good species, and this was confirmed on examining the spores, which are up to c. 42 wm long. These have a remarkable surface of somewhat dense blunt spines, unlike any other Dryopteris examined, and do not have clear wings of perispore. In Dryopteris schimperiana, which is close to D. ruwenzoriensis, the spores are smaller, and have short, more or less clear wings of perispore. The other major difference is the more prominent teeth at the apices of the segments in D. ruwenzoriensis, and the often somewhat reddish-brown stipe-base scales; the pinnule- apices are also slightly more obtuse than in many specimens of D. schimperiana, and the lamina is laxer. The indusium can be small in D. schimperiana on occasions, but is always either small or absent in D. ruwenzoriensis. The stipe contains a circle of about six or seven separate vascular strands. So far D. ruwenzoriensis is known only from the Ruwenzori (Mountains of the Moon, Uganda ' and Zaire), but it may well occur elsewhere in Africa, and a search in other herbaria may reveal it, identified most probably as D. schimperiana. Other specimens examined: Uganda, Western Province, Toro District, Ruwenzori, Namwamba valley, 3200 m, in heath forest, 6 Jan. 1935, G. Taylor 2954 (BM!), on rocks in river, 3170 m, 7 Jan. 1935, G. Taylor 2990 (BM!), under Podocarpus trees, 2900 m, 11 Jan. 1935, G. Taylor 3079a (BM!); Uganda, Western Province, Bigo, River Bujuku, riverside, 3450 m, 23 July 1952, H. A. Osmaston 1687 (BM!). CLASSIFICATION OF DRYOPTERIS 205 vPE SPECIMEN Digeptirvc_remengerivanin ©. Che HERR, CARL CHRISTENSEN. Purchased ¢ a4 SS eatin — Mission H. HUMBERT - Piantes de | AFRIQUE EQUATORIALE - 1929 Pea. peers tt ta nant Qret (CONGO BELGE, Tun 4 Woe HERBARIUM MUSE? BRITANNICI Fig.4 Dryopteris ruwenzoriensis C. Chr. Holotype (Humbert 8825) (BM). 206 C. R. FRASER-JENKINS Hybrids A list of 89 naturally occurring Dryopteris hybrids (excluding hybrid-derived apomictic or polypoid species), as known to the author, is given below in alphabetical order by regions. However, it should be noted that apart from North America, Europe (in the wide sense) and Japan, search for hybrids has been non-existent or only preliminary; see Fraser-Jenkins (1984: 39-40) for information about the identification of hybrids from spores, etc. The Japanese hybrids are taken from Nakaike (1970, 1975) and Hirabayashi (1974). A useful list of hybrids, though also containing polyploid species, is that of Knobloch (1976), and North American hybrids are taken from this source, and also from Wagner (1970) and Widén, Britton, Wagner & Wagner (1975). Knobloch’s list was intended to bring together all the available literature records and did not set out to decide the validity of the hybridity or nomenclature. It has been revised recently by Knobloch, Gibby & Fraser-Jenkins (1984). European hybrids are those verified by the present author after detailed long-term study; thus there are some modifications from published lists and the list here is believed to be definitive for the present. Where the hybrids have not yet been given binomials they are listed after the first parental species, the normal convention of placing the parents in alphabetical order being followed. Europe (including western Asia, Macaronesia, and north-west Africa) (23): D. X ambroseae Fraser-Jenkins & Jermy (= D. dilatata x D. expansa); D. X asturiensis ined. (= D. affinis subsp. affinis x D. corleyi); D. X brathaica Fraser-Jenkins & Reichst. (= D. carthusiana x D. filix-mas); D. X cedroensis Gibby & Widén (= D. guanchica x D. oligodonta); D. X complexa Fraser-Jenkins in press, and nothosubspecies (= D. affinis subspp. x D. filix-mas); D. x deweveri (J. Jansen) J. Jansen & Wachter (= D. carthusiana X D. dilatata); D. expansa x D. filix-mas; D. X euxinensis Fraser-Jenkins & Corley (= D. caucasica x D. filix-mas); D. x fraser-jenkinsii Gibby & Widén (= D. affinis subsp. affinis X D. guanchica); D. X gomerica Gibby & Widén (= D. aemula x D. guanchica); D. X graeca Fraser-Jenkins & Gibby (= D. pallida subsp. pallida x D. submontana); D. X initialis Fraser-Jenkins & Corley (= D. caucasica X D. oreades); D. X madalenae Fraser-Jenkins (= D. azorica x D. crispifolia); D. X mantoniae Fraser-Jenkins & Corley (= D. filix-mas x D. oreades); D. X martinsiae Fraser-Jenkins (= D. aemula x D. crispifolia); D. X picoensis Fraser-Jenkins (= D. affinis subsp. affinis x D. azorica); D. X pseudoabbre- viata Jermy (= D. aemula x D. oreades); D. X sardoa Fraser-Jenkins & Reichst. (= D. oreades x D. tyrrhena); D. X sarvelae Fraser-Jenkins & Jermy (= D. carthusiana x D. expansa); D. X sjoegrenii Fraser-Jenkins (= D. azorica x D. dilatata); D. X telesii Fraser-Jenkins (= D. crispifolia x D. dilatata); D. xX uliginosa (A. Braun ex Doell) Kuntze ex Druce (= D. carthusiana x D. cristata); D. x vidae Fraser-Jenkins & Gibby (= D. pallida subsp. pallida x D. villarii). North America (31): D. X algonquinensis D. Britton (= D. fragrans x D. marginalis); D. X atropalustris Small (= D. celsa x D. cristata); D. X australis (Wherry) Small (= D. celsa x D. ludoviciana); D. x benedictii Wherry (= D. carthusiana X D. clintoniana); D. x boottii (Tuckerman) L. Underw. (= D. cristata x D. intermedia); D. x burgessii B. Boivin (= D. clintoniana x D. marginalis); D. campyloptera x D. expansa; D. campylop- tera X D. intermedia; D. campyloptera x D. marginalis;? D. carthusiana X D. celsa; D. carthusiana x D. goldiana; D. celsa x D. clintoniana; D. celsa x D. goldiana; D. clintoniana x D. cristata; D. clintoniana X D. goldiana; ? D. cristata x D. goldiana; D. X dowellii (Farw.) Wherry (= D. clintoniana x D. intermedia); D. expansa X D. intermedia; D. expansa X D. marginalis; D. filix-mas X D. goldiana; D. filix-mas X D. marginalis; ? D. fragrans X D. intermedia; D. goldiana x D. intermedia; D. intermedia x D. marginalis; D. X leedsii Wherry (= D. celsa x D. marginalis); D. X neo-wherryi W. Wagner (= D. goldiana X D. marginalis); D. X pittsfordensis Slosson (= D. carthusiana x D. marginalis); D. Xx separabilis (W. Palmer) Small (= D. celsa x D. intermedia); D. X slossoniae Wherry ex Lellinger (= D. cristata X D. marginalis); D. X triploidea Wherry (= D. carthusiana x D. intermedia); D. x uliginosa (A. Braun ex Doell) Kuntze ex Druce (= D. carthusiana x D. cristata). Mainland China and Korea (4): D. X daliensis Z. R. Wang in press (= D. woodsiisora X D. zinongii); unnamed hybrid of D. fangii;? D. x saxifragi-varia Nakai (? = D. bissetiana x D. varia); D. X shuichangensis Chiu & Yao ex Ching (pro sp.) (= D. crassirhizoma subsp. whangshangensis X D. dickinsii). CLASSIFICATION OF DRYOPTERIS 207 Japan (23): Many of the Japanese hybrids have not been reported clearly enough or with sufficient detail of spores, etc., to ascertain that they are hybrids in the sense of this account; this list could, perhaps, contain some which are apomictic species. D. X fujipedis Kurata (= D. crassirhizoma subsp. crassirhizoma x D. lacera); D. X gotenbaensis Nakaike (= D. hondoensis x D. uniformis); D. X haganecola Kurata (= D. cycadina X D. uniformis); D. x hakonecola Kurata (= D. dickinsii x D. uniformis); D. X hisatsuana Kurata (= D. handeliana x D. lacera); D. X kidoana Kurata nom. nud. (= ? D. commixta x D. sp.); D. X kominatoensis Tag. (= D. monticola x D. tokyoensis); D. X mayebarae Tag. (= D. commixta x D. uniformis); D. x mituii Seriz. (= D. lacera X D. uniformis); D. X otomasui Kurata (= D. commixta X ? D. polylepis or ? D. uniformis); D. X pseudo-commixta Kurata (= D. commixta x D. dickinsii); D. X rarissima Kurata (= D. commixta X D. sparsa);? D. x shibisanensis Kurata (= D. commixta x D. cycadina); D. X sugino-takaoi Kurata (= D. lacera x D. polylepis); D. < tetsu-yamanakae Kurata (= D. commixta x D. sieboldii); D. x tokudai Sugim. (= D. crassirhizoma subsp. crassirhizoma X D. polylepis); D. X toyamae Tag. (= D. sieboldii x? D. lacera); D. X watanabei Kurata (= D. crassirhizoma subsp. crassirhizoma X D. uniformis); D. X yamashitae Kurata (= D. sparsa x D. subexaltata). According to Hirabayashi (pers. comm. 1982) a few other hybrids recently found in Japan are soon to be described, and Nakaike (1982) lists D. x kousaii Akas., D. X tonensis Kurata, D. X wakui Kurata, and D. X yasuhikoana Kurata. Indian subcontinent (9): D. x flemingii Fraser-Jenkins in press (= D. chrysocoma x D. juxtaposita); D. X ghatakii Fraser-Jenkins in press (= D. austro-indica x D. cochleata); D. x liddarensis Fraser-Jenkins in press (= D. barbigera subsp. barbigera x D. serrato-dentata); D. x loyalii Fraser-Jenkins in press (= D. caroli-hopei x D. marginata); D. X macdonellii Fraser-Jenkins in press (= D. filix-mas x D. ramosa); D. sparsa (2X) x D. sparsa (4x); D. x vidyae Fraser-Jenkins in press (= D. sparsa x D. splendens); D. wechteriana Fraser-Jenkins in press (= D. chrysocoma xX D. nigropaleacea); D. x zygo-parentalis Fraser-Jenkins in press (= D. darjeelingensis x D. scottii). List of synonyms The following is a list of synonyms within the genus Dryopteris. The species to which the synonyms are referred may be found listed after each section or in the index. In Christensen’s Index filicum and supplements (1905-1934), Pichi Sermolli (1965), and Jarrett et al. (1985) many more species are placed in Christensen’s subgenus Dryopteris than in the modern genus Dryopteris as he included Ctenitis and other closely related genera such as Nothoperanema, as well as many thelypteroid ferns, and many that are synonyms of Dryopteris species. It is hoped that the present list is nearly complete and will complement the Index filicum as far as Dryopteris (in the modern sense) is concerned. All the names in Index filicum and supplements have been checked and any not appearing in the present list have been excluded from Dryopteris. abbreviata (DC.) Newman ex Manton, non (Schrader) Kuntze = affinis subsp. affinis abbreviata auct., non (DC.) Newman ex Manton, nec (Schrader) Kuntze = oreades acutidens Ching in Ching & Wang, non C. Chr. = purpurella adenorachis C. Chr. = fructuosa adiantoides T. Suzuki = hasseltii affinis (Fischer & C. Meyer) Newman ex C. Chr., comb. inval. (in syn.) = caucasica alexeenkoana Fomin = dilatata alpicola Ching & Z. R. Wang, in press = chrysocoma ambigens (Franchet & P. A. L. Savat.) Koidz. = uniformis ambigens ‘(Nakai) Koidz.’ = uniformis ambigua Druce, nom. nud. = X deweveri ambigua Sledge = deparioides subsp. ambigua (Sledge) Fraser-Jenkins, in press americana (Fischer ex Kunze) Clute, nom. provis. (inval.) = campyloptera amoena Ching = kwanzanensis amurensis (Milde) Takeda, non Christ = amurensis aneitensis (Hook.) C. Chr. = ? hasseltii angustipinna Ching & S. K. Wu ex Fraser-Jenkins, nom. nud. = pulcherrima aperta (Fée) C. Chr. = patula apicifixa Ching, Boufford & Shing = fructuosa apicisora Ching & Y. T. Hsieh = peninsulae 208 C. R. FRASER-JENKINS aquilonaris Maxon = fragrans aristata (Villars) Druce, non (G. Forster) Kuntze = expansa assimilis S. Walker = expansa athyrioides (M. Martens & Galeotti) Kuntze = cinnamomea atrata (Kunze) Ching = hirtipes subsp. atrata (Kunze) Fraser-Jenkins, in press australis (Ten.) Guadagno = pallida austriaca auct., non (Jacq.) Woynar ex Schinz & Thell. = dilatata balearica (Litard.) Nardi = pallida subsp. balearica (Litard.) Fraser-Jenkins barbellata Fomin = sichotensis basiaurita Ching = dickinsii bernieri Tard. = bojeri bipinnata C. Chr. in A. Léveillé, non Copel. = fuscipes blepharolepis C. Chr. = sublacera blinii A. Léveillé = marginata Xx bohemica Domin, nom. inval. = filix-mas x borbasii Litard. = remota borreri (Newman) Newman ex Tavel = affinis subsp. borreri (Newman) Fraser-Jenkins x boydii (Stansf.) C. Morton = remota bullatipaleacea Ching = championii bulligera Ching = erythrosora x burnatii Christ & Wilczek = villarii buschiana Fomin = crassirhizoma subsp. crassirhizoma callolepis C. Chr. = antarctica canaliculata Ching = pulcherrima canaliculatis Ching ex Fraser-Jenkins, nom. nud. = pulcherrima caudifrons Ching in Ching & Wang = integriloba caudipinna Nakai = ? erythrosora cavalerii A. Léveillé, non (Christ) C. Chr. = fructuosa x cebennae Fraser-Jenkins = tyrrhena centro-chinensis Ching = rosthornii changii Ching, non zhangii Ching = championii changtouensis Ching ex Fraser-Jenkins, nom. nud. = pulcherrima chapaensis C. Chr. & Ching ex Ching = polita chichisimensis Nakai ex H. Ité = insularis chingii Nair = pulcherrima cochin-chinensis Ching, nom. nud. = integriloba confertipinna Ching & Shing = fuscipes constantissima Hayata = formosana cordipinnula C. Chr. = remotipinnulata coreano-montana Nakai = sichotensis cyrtolepis Hayata = wallichiana decurrentiloba Ching & C. F. Zhang, in press = uniformis dehuaensis Ching & Shing = bissetiana dentipalea Nakai = uniformis discreta Ching & S. K. Wu = pulcherrima dispar Ching & C. F. Zhang, in press = fuscipes x doeppii Rothm. = remota doiana Tag. = wallichiana x doluchanovii A. Askerov = remota doniana (Spreng.) Ching = wallichiana doshunglaensis Ching & S. K. Wu, nom. nud. = alpestris elongata (Aiton) T. Sim, non (J. Smith) Kuntze = aitoniana elongata ‘(Sw.) Chev.’ = aitoniana emigrans Copel. = deparioides subsp. concinna C. Chr. enneaphylla (Baker) C. Chr. = sieboldii espinosai Hicken = karwinskyana euspinulosa (Asch.) Fomin, nom. inval. = carthusiana extremiorientalis V. Vassiljev, nom. nud. = expansa CLASSIFICATION OF DRYOPTERIS 209 falconeri (Hook.) Kuntze = barbigera subsp. barbigera fatuhivensis E. Brown = ? hirtipes subsp. hirtipes fenyangshanensis Ching & C. F. Zhang, in press = cycadina fibrillosa (C. B. Clarke) Hand.-Mazz., non (Baker) C. Chr. = pulcherrima fibrillosissima Ching = pulcherrima floridana (Hook.) Kuntze = ludoviciana fournieri (Baker) C. Chr. = mexicana fuliginosa C. Chr. = cognata gamblei (C. Hope) C. Chr. = stenolepis gillespiei Copel. = aneitensis giraldii (Christ) C. Chr. [lectotype: ‘Nephrod. non elongatum, sed var. giraldiin. var. Christ, Alla meta del monte Si-Ku-Tzui-San, J. Giraldi, [18] Juglio 1894’ (P!)] = sericea gongboensis Ching = blanfordii subsp. nigrosquamosa (Ching) Fraser-Jenkins, in press gracillima Ching = deparioides subsp. gracillima (Ching) Fraser-Jenkins, in press grandissima Tag. = marginata grossa (Christ) C. Chr. = scottii gushanica Ching & Shing = bissetiana gushiangensis [‘gushaingensis’] Ching = blanfordii subsp. nigrosquamosa (Ching) Fraser-Jenkins, in press gutishanensis Ching & C. F. Zhang, in press = championii harae H. It6 = pulvinulifera hatusimae H. It6 = hasseltii hayatae Tag. = subexaltata heleopteroides Christ = cochleata x hercynica Rothm., nom. nud. = carthusiana heterolepia Alderw. = chaerophyllifolia heteroneura (Tag.) Ching = pseudosieboldii hirtosparsa Y. C. Wu, non Christ = integriloba hololepis (Hayata) Tag. = varia hopei Fraser-Jenkins ex Rush, nom. nud. = caroli-hopei huangangshanensis Ching = rosthornii hunanica Ching = kinkiensis [juvenile] hupehensis Ching = rosthornii hwangii Ching = integriloba hypophlebia Hayata = fructuosa immixta Ching = bissetiana indecora (Liebm.) C. Chr. = cinnamomea indusiata (Makino) Makino & Yamamoto ex Yamamoto = tenuicula infrahirtella Ching & Z. Y. Liu, in press = championii infrapuberula Ching, Boufford & Shing = namegatae integripinnula Ching = cordipinna inuyamensis H. It6 = fuscipes jiulungshanensis Chiu & Yao ex Ching = purpurella julaodongensis Ching & Z. R. Wang in Z. R. Wang & Z. X. Zhang, nom. nud. = rosthornii junlianensis Kung = lepidopoda kaihuaensis Ching & C. F. Zhang, in press = fuscipes kemulariae Mikheladze = remota kobayashii Kitagawa = sacrosancta kodamae Hayata = formosana komarovii Kossinsky = barbigera subsp. komarovii (Kossinsky) Fraser-Jenkins, in press kongboensis Ching ex Fraser-Jenkins, nom. nud. = blanfordii subsp. nigrosquamosa (Ching) Fraser- Jenkins, in press kooshiangensis Ching ex Fraser-Jenkins, nom. nud. = blanfordii subsp. nigrosquamosa (Ching) Fraser- Jenkins, in press koraiensis Tag. = gymnophylla kuratae Nakaike, nom. nud. = ? hangchowensis labordei (Christ) C. Chr. = gymnosora laeta (V. Komarov) C. Chr., non (Sw.) C. Chr. = goeringiana 210 C. R. FRASER-JENKINS lanceolato-cristata (Hoffm.) Alston = carthusiana lancifrons Hayata ex Fraser-Jenkins, nom. nud. = fructuosa lancipinnula Ching, nom. nud. = subimpressa x laschii E. Walter = x uliginosa laserpitiifolia (Scortech. ex Beddome) C. Chr., non (Mett.) Kuntze = hasseltii lastii (Baker) C. Chr. = kilemensis latibasis Ching = lepidopoda laurisilvicola T. Suzuki = polita x lawalreei Janchen = remota layardii (Baker) C. Chr. = aneitensis lepidorachis C. Chr. = championii lepinei (Kuhn) Kuntze = dicksonioides leveillei Nakai, non Christ = marginata libanotica (Rosenstock) C. Chr. = pallida subsp. libanotica (Rosenstock) Nardi likiangensis Ching, non lichiangensis (C. H. Wright) C. Chr. = montigena liliana Golicin [Golitsin] = aemula linganensis Ching & C. F. Zhang, in press = championii lingtzeensis Ching ex Fraser-Jenkins, nom. nud. = sublacera linyingensis Ching & C. F. Zhang, in press = erythrosora x litardierei Rothm. = oreades lividis Ching & C. H. Wang = polita liyangensis Ching & Lan = ? erythrosora longistipes Ching = lepidopoda ludens (Baker) C. Chr. = arborescens macaronesica Romariz = aitoniana macrocarpa R. Stewart = chrysocoma xX madeniana Kunkel, nom. nud. = oligodonta maderensis Milde ex Alston = intermedia subsp. maderensis (Milde ex Alston) Fraser-Jenkins makinoi Koidz. = fuscipes manshurica Ching ex Wang et al. = expansa matsuzoana Koidz. = varia maxima (Baker) C. Chr. = arborescens medioxima Koidz. = fuscipes mediterranea Fomin = wallichiana mehrae Khullar, nom. nud. = blanfordii subsp. blanfordii melanocarpa Hayata = platypus melanolepis (Alderw.) Alderw., comb. inval. (in syn.) = scottii metafuscipes Ching & C. F. Zhang, in press = erythrosora metcalfii Ching = marginata mimetica Ching & C. F. Zhang, in press = cordipinna mingetsensis Hayata ex Fraser-Jenkins, nom. nud., non mingetsuensis Hayata = fructuosa mingetsuensis Hayata = reflexosquamata minimisora Nakai = expansa minjiangensis Kung = sublacera xX mixta Rothm., nom. nud., non Rosenstock = X mantoniae monticola (Makino) C. Chr. = goldiana subsp. monticola (Makino) Fraser-Jenkins morrisonensis (Hayata) Hayata = expansa multijugata Ching & Shing ex Ching & C. F. Zhang, in press = fuscipes nakanensis Ching = microlepis nanchuanensis Ching & Liu, in press = tenuicula X neglecta Domin, non Brade & Rosenstock = ? X deweveri neoassamensis Ching = assamensis neochrysocoma Ching = woodsiisora neolacera Ching = peninsulae neolepidopoda Ching & S. K. Wu = lepidopoda neopodophylla Ching = podophylla nigra Ching = lepidopoda nigrisquama Hayata = scottii CLASSIFICATION OF DRYOPTERIS 211 nigrosquamosa Ching = blanfordii subsp. nigrosquamosa (Ching) Fraser-Jenkins, in press nilamoensis Ching & S. K. Wu ex Fraser-Jenkins, nom. nud. = pulcherrima nipponensis Koidz. = cystolepidota nyalamensis [‘nyalamense’| Ching & S. K. Wu = pulcherrima nyingchiensis Ching = sublacera obovata (Baker) C. Chr. = hasseltii obtusissima (Makino) Makino, non (Mett. ex Kuhn) Christ = fuscipes obtusissima (Mett. ex Kuhn) Christ, non (Makino) Makino = macrochlamys odontophora Copel. = aneitensis ogawae [‘ogawai’] H. It6 = varia okushirensis Miyabe & Kudé6 = dickinsii omeicola Ching = rosthornii oxyodon (Franchet) Kitagawa, non oxyodus (Baker) C. Chr. = goeringiana pachyphylla Hayata = wallichiana paleacea Hand.-Mazz., non (Lagasca ex Sw.) C. Chr., nec (T. Moore) Fomin = wallichiana paleacea (Lagasca ex Sw.) C. Chr., non Hand.-Mazz., nec (T. Moore) Fomin = wallichiana paleacea (T. Moore) Fomin, non (Lagasca ex Sw.) C. Chr., nec Hand.-Mazz. = wallichiana paleacea ‘(Don) Druce’ = wallichiana paleacea ‘(Don) Hand.-Mazz.’ = wallichiana paleacea ‘(Sw.) Hand.-Mazz.’, comb. inval. = wallichiana pallida ‘(Bory) Fomin’ = pallida pandurata Ching ex Fraser-Jenkins, nom. nud. = basisora para-chrysocoma Ching & Z. R. Wang = ? chrysocoma parallelogramma (Kunze) Alston = wallichiana parasparsa Ching & S. K. Wu = sparsa patagonica Diem = filix-mas patentissima (Wallich ex Kunze) Nair = wallichiana patentissima ‘(Franch.) Nair’ = wallichiana persimilis Ching & C. F. Zhang, in press = fuscipes petelotii C. Chr. = polita phaeocoma Ching & S. K. Wu ex Fraser-Jenkins, nom. nud. = lepidopoda phaeolepis Hayata = formosana platylepis Rosenstock = kilemensis poilanei Tard. = bodinieri x poyseri Wherry = clintoniana pseudatrata Ching = cycadina pseudoerythrosora Ching & C. F. Zhang, in press, non Kodama = erythrosora pseudo-erythrosora Kodama, non Ching & C. F. Zhang = championii pseudodontoloma Ching = lachoongensis pseudofibrillosa Ching = redactopinnata pseudo-filix-mas Nakai & Kodama, nom. nud., non (Fée) Rothm. = crassirhizoma subsp. crassirhizoma pseudo-lunanensis Tag. = dickinsii pseudomarginata Ching, nom. nud. = caroli-hopei pseudomas (Wollaston) Holub & Pouzar = affinis subsp. affinis pseudo-sabae [‘pseudo-sabaei’| Hayata = fructuosa pseudo-sikkimensis Ching & S. K. Wu = sikkimensis pseudosparsa Ching = purpurella pseudouniformis Ching = uniformis pseudovaria (Christ) C. Chr. = fructuosa psilosora Tag. = hasseltii pudouensis Ching, in press = pacifica qamdoensis Ching = pulcherrima quatanensis Ching = wallichiana raddeana (Fomin) Fomin = pallida subsp. raddeana (Fomin) Nardi raynalii Tard. = manniana reflexipinna Hayata = diffracta reholttumii M. Price, nom. nud. = pulvinulifera 242 C. R. FRASER-JENKINS remota ‘(A. Br.) Hayek’ = remota (A. Braun ex Doell) Druce, non Hayata remotipinnata Ching & C. F. Zhang, in press = fuscipes remotissima (Christ ex Matsum.) Koidz. = sabae resendeana Rezende-Pinto = affinis subsp. affinis retroso-paleacea Ching & C. F. Zhang, in press = cordipinna rhomboideo-ovata H. It6é = hondoensis rigida (Sw.) A. Gray = villarii rigida ‘(Hoffm. ex Sw.) Gray’ = villarii rigida ‘(Sw.) Underw.’ = villarii rossii (C. Chr.) C. Chr. = ? glandulifera rubristipes Ching & Z. Y. Liu, in press = purpurella saamingensis Ching = tokyoensis sakuraii (Rosenstock) Tag. = gymnophylla X satsumana Kurata = X pseudo-commixta schneideriana Hand.-Mazz. = sublacera x schorapanensis A. Askerov = affinis subsp. persica Fraser-Jenkins semipinnata Ching = lunanensis setosa (Christ) Kud6, non (Thunb.) Akas., nec (Blume) Kuntze, nec (C. Presl) C. Chr. = sichotensis setosa (Thunb.) Akas., non (Blume) Kuntze, nec (C. Presl) C. Chr., nec (Christ) Kud6é = bissetiana shensicola Ching & Y. T. Hsieh = peninsulae silaensis Ching = acuto-dentata simulans Ching, non (Baker) Kuntze = sri-lankensis sinoerythrosora Ching & Shing = erythrosora sinofibrillosa Ching = pulcherrima sinosparsa Ching & Shing = viridescens siranensis Nakai = expansa spinulosa (Roth) Kuntze = carthusiana spinulosa ‘(Miill. ex Roth) O. Ktze.’ = carthusiana squamifera [‘squamigera’| Ching & S. K. Wu = pulcherrima x subalpina (Borbas) Domin, non Alderw. = remota subassamensis Ching = subtriangularis subatrata Tag. = cycadina x subaustriaca Rothm. = remota subbarbigera Ching = barbigera subsp. komarovii (Kossinsky) Fraser-Jenkins, in press subdecipiens Hayata = scottii subintegriloba Seriz. = integriloba sublaeta Ching & Y. P. Hsu = goeringiana submarginalis Ching, Boufford & Shing = crassirhizoma subsp. whangshangensis (Ching) Fraser-Jenkins submarginata Loyal, non Rosenstock = subimpressa submarginata Rosenstock, non Loyal = tenuicula submonticola Nakai = goldiana subsp. monticola (Makino) Fraser-Jenkins subodontoloma Ching ex Loyal, nom. nud. = subimpressa subopposita Kodama ex Nakai, non Kuntze ex T. Mori = expansa subramosa Christ = goeringiana sunnii Ching = pycnopteroides supraimpressa Ching, Boufford & Shing = porosa tabacicoma Alderw. = subarborea tahmingensis Ching = integriloba taitoensis Tag., non taitunensis Koidz. = polita taiwanicola Tag. = lepidopoda takesimensis Kodama ex Nakai = erythrosora takeuchiana Koidz. = formosana taquetii Christ = erythrosora tarningensis Ching = fuscipes tasiroi Tag. = handeliana x tauschii (Celak.) Domin = x uliginosa x tavelii Rothm. = affinis subsp. borreri tenuissima Tag. = woodsiisora CLASSIFICATION OF DRYOPTERIS 213 thibetica (Franchet) C. Chr. = dickinsii thwaitesii (Baker) Kuntze, non (Hook.) C. Chr. = deparioides subsp. concinna C. Chr. tieluensis Ching & Y. P. Hsu = basisora tienmohshangensis Ching = uniformis triangularifrons Ching = gymnosora tsangpoensis Ching = redactopinnata tsaytiensis Ching & S. K. Wu ex Fraser-Jenkins, nom. nud. = pulcherrima tsutsuiana Kurata = namegatae undulata (Beddome) Kuntze = macrochlamys uropinna M. Price = subtriangularis ursipes Hayata = wallichiana venosa Ching & S. K. Wu = lachoongensis vescoi (Drake) C. Chr. = dicksonioides villarsii auct. = villarii viridis Ching = polita wenchuanensis Kung = ? habaensis whangshangensis Ching = crassirhizoma subsp. whangshangensis (Ching) Fraser-Jenkins wherryi Crane = celsa wladiwostokensis B. Fedtsch. = goeringiana X woynarii Rothm. = remota wuyishanensis Ching = kinkiensis xanthomelas (Christ) C. Chr. = rosthornii yabei Hayata = varia yakusilvicola Kurata = cacaiana yandongensis Ching & C. F. Zhang, in press = kinkiensis yaoi Ching = tenuicula yikungensis Ching ex Fraser-Jenkins, nom. nud. = yigongensis yui Ching = bonatiana yungtzeensis Ching = dickinsii Xx yuyamae Kurata = X hakonecola zayliensis Ching & S. K. Wu = pulcherrima zhangii Ching, in press, non changii Ching = tenuicula Acknowledgements I would like to thank the following for help in various ways: A. O. Chater, M. Gibby, A. J. Harrington, and J. R. Laundon (London), R. C. 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Taxonomic index Accepted names are given in roman and synonyms in italic; new names and principal page references are in bold. An asterisk (*) denotes a figure. See pp 207-213 for a list of synonyms. Acrorumohra 186, 187, 188, 197, 198 Arachniodes 185, 186, 188, 195, 197, 198 Arthrobotrys 187 macrocarpa 187 Aspidiaceae 184, 185 Aspidiineae 184 Aspidium 184, 188 Dichasium, subgen. 190 Diclisodon, group 197 Filix-mas, group 191 Fragrantia, group 191 patentissimum 188 Sparsa, group 197 Spinulosa, group 195 Aspleniaceae 184 Athyriaceae 184 Byrsopteris 188 Ctenitis 185, 186, 195, 198, 207 apiciflora 185 Dennstaedtiaceae subfam. Dryopteridoideae 184 Dichasium 188, 190 Diclisodon 188 deparioides 188 Dryopsis 185 Dryopteridaceae 184, 185 Dryopteris abbreviata 192 acutidens 198 acuto-dentata 191, 202, 212 aemula 194, 195, 210 Aemulae, sect. 183, 187, 194, 195 affinis 191 affinis subsp. affinis 207, 211, 212 affinis subsp. borreri 208, 212 affinis subsp. persica 212 aitoniana 193, 208, 210 x algonquinensis 206 alpestris 192, 208 X ambroseae 206 amurensis 195, 207 aneitensis 198, 209, 210, 211 angustifrons 194 antarctica 195, 208 approximata 194 aquilinoides 194 arborescens 198, 210 ardechensis 192 arguta 193 ascensionis 198 assamensis 195, 196, 210 x asturiensis 206 athamantica 194 X atropalustris 206 X australis 206 austro-indica 191 azorica 195 bamleriana 193 barbigera 192 barbigera subsp. barbigera 209 barbigera subsp. komarovii 202, 209, 212 basisora 193, 211, 213 X benedictii 206 bissetiana 197, 208, 209, 212 blanfordii 192 blanfordii subsp. blanfordii 210 blanfordii subsp. nigrosquamosa 209, 211 bodinieri 189, 211 bojeri 194, 208 bonatiana 191, 213 X boottii 206 x brathaica 206 Bulligerae, sect. 187, 196 burgessii 206 cacaiana 198, 213 campyloptera 195, 207 campyloptera X expansa 206 campyloptera X intermedia 206 campyloptera X marginalis 206 caroli-hopei 194, 209, 211 carthusiana 195, 208, 209, 210, 212 carthusiana X celsa 206 carthusiana X goldiana 206 caucasica 192, 207 caudipinna 196 X cedroensis 206 celsa 191, 213 celsa X clintoniana 206 celsa X goldiana 206 chaerophyllifolia 194, 209 championii 196, 208, 209, 210, 211 chinensis 195 chrysocarpa 191 chrysocoma 191, 207, 210, 211 cinnamomea 193, 194, 208, 209 Cinnamomeae, sect. 183, 187, 188, 190, 193 clintoniana 191, 211 clintoniana X cristata 206 clintoniana X goldiana 206 cochleata 187, 194, 209 cognata 198, 209 commixta 190 Xx complexa 206 conjugata 190, 199 cordipinna 196, 209, 210, 212 coreano-montana 192 corleyi 192 costalisora 191 crassirhizoma 191 crassirhizoma subsp. crassirhizoma 208, 211 crassirhizoma subsp. whangshangensis 183, 191, 212, 213 crinalis 198 crispifolia 195 cristata 191 cristata X goldiana 206 cycadina 190, 209, 211, 212 Cycadinae, subsect. 190 cyclopeltidiformis 195, 196 cystolepidota 196, 211 x daliensis 206 darjeelingensis 190 decipiens 196 deparioides 188, 198 Dryopteris contd. deparioides subsp. ambigua 207 deparioides subsp. concinna 208, 213 deparioides subsp. gracillima 209 x deweveri 206, 207, 210 Dichasium, subsect. 190 dickinsii 190, 199, 208, 211, 213 dicksonioides 198, 210, 213 Diclisodon, group 197 diffracta 197, 198, 211 dilatata 187, 195, 207, 208 x dowellii 206 dracomontana 194 Dryopteris, subgen. 186, 187, 189, 193, 195, 197, 198, 207 Dryopteris, sect. 187, 190, 191, 192, 193, 194 erythrosora 195, 196, 208, 210, 211, 212 Erythrovariae, sect. 186, 187, 196 Erythrovariae, subgen. 183, 186, 187, 188, 190, 195, 196, 197, 198 Erythro-variae, subsect. 196 esterhuyseniae 194 Eudryopteris, sect. 189, 190, 192 Eudryopteris, subgen. 189 X euxinensis 206 expansa 195, 208, 210, 212 expansa filix-mas 206 expansa X intermedia 206 expansa X marginalis 206 fadenii 194 fangii 191 fatuhivensis 190 fibrillosa 190 Fibrillosae, sect. 186, 187, 188, 190, 191, 192 filix-mas 188, 189, 190, 191, 192, 208, 211 filix-mas X goldiana 206 Filix mas, group 192 filix-mas X marginalis 206 X flemingii 207 formosana 197, 208, 209, 211, 212 Formosanae, subsect. 196 fragrans 192, 208 fragrans X intermedia 206 Fragrantes, subsect. 192 x fraser-jenkinsii 206 fructuosa, 193, 207, 208, 209, 210, 211 x fujipedis 207 fuscipes 196, 208, 209, 210, 211, 212 fusco-atra 191 futura 198 x ghatakii 207 glabra 198 glandulifera 194, 212 goeringiana 194, 209, 211, 212, 213 goldiana 192 goldiana subsp. monticola 184, 192, 210, 212 goldiana X intermedia 206 CLASSIFICATION OF DRYOPTERIS X gomerica 206 xX gotenbaensis 207 x graeca 206 guanchica 195 gymnophylla 195, 209, 212 gymnosora 196, 209, 213 Gymnosorae, subsect. 196 habaensis 191, 213 hadanoi 197 x haganecola 207 x hakonecola 207, 213 handeliana 190, 212 hangchowensis 190, 209 hasseltii 198, 207, 209, 210, 211 hawaiiensis 191 hayatae 197, 198 himachalensis 191 Hirtipedes, sect. 183, 187, 188, 190, 191 hirtipes 190 hirtipes subsp. atrata 208 hirtipes subsp. hirtipes 209 X hisatsuana 207 hondoensis 196, 212 huberi 194 inaequalis 188, 194 incisolobata 191 X initialis 206 insularis 197, 208 integriloba 195, 196, 208, 209, 212 intermedia 195 intermedia subsp. maderensis 210 intermedia X marginalis 206 juxtaposita 193 karwinskyana 194, 208 kenzoi 196 khullarii 191 X kidoana 207 kilemensis 198, 210, 211 kinkiensis 196, 209, 213 kinokuniensis 196 koidzumiana 196 X kominatoensis 207 X kousaii 207 kwanzanensis 193, 207 lacera 193 lachoongensis 193, 211, 213 laurisilvicola 196 X leedsii 206 lepidopoda 191, 209, 210, 211, 212 lewalleana 194 liankwangensis 190 X liddarensis 207 liyangensis 196 Lophodium, group 195 Lophodium, sect. 187, 192, 194, 195 X loyalii 207 ludoviciana 191, 209 lunanensis 190, 212 x macdonnellii 207 macrochlamys 197, 198, 211, 213 x madalenae 206 madrasensis 191 mangindranensis 194 manniana 194, 211 217 X mantoniae 206, 210 marginalis 187, 193 marginata 194, 208, 209, 210 Marginatae, sect. 183, 187, 190, 192, 194, 195, 198 x martinsiae 206 maxonii 194 xX mayebarae 207 meeteetseana 184 mexicana 194, 209 microlepis 190, 210 mindshelkensis 192 X mituii 207 Monticolarum, subsect. 192 montigena 192, 210 munchii 194 namegatae 190, 209, 213 napoleonis 198 neorosthornii 191 X neo-wherryi 206 Nephrocystis, sect. 186, 187, 197, 198 Nephrocystis, subgen. 183, 185, 186, 187, 193, 196, 197, 198 nigropaleacea 193 nobilis 191 nubigena 198 nuda 198 obtusissima 197 odontoloma 193 oligodonta 194, 210 oreades 192, 207, 210 Xx otomasui 207 pacifica 197, 211 paleacea var. madagascariensis 184, 191 pallida 192, 193, 208, 211 pallida subsp. balearica 208 pallida subsp. libanotica 210 pallida subsp. raddeana 211 Pallidae, sect. 183, 187, 190, 192, 193, 194 panda 191 Pandae, sect. 183, 187, 191, 193 papuana 198 parachrysocoma 191 parrisiae 184, 191, 199, 201* parvula 198 patula 194, 207 peninsulae 193, 207, 210, 212 pentheri 194 permagna 198 perrieriana 198 X picoensis 206 X pittsfordensis 206 platypus 198, 210 podophylla 189, 210 polita 195, 196, 208, 210, 211, 212, 213 Politae, sect. 183, 187, 196 polylepis 191 Polysticho-drys, sect. 186, 195, 196 porosa 194, 212 powellii 198 x pseudoabbreviata 206 218 Dryopteris contd. xX pseudo-commixta 207, 212 pseudo-filix-mas 191 pseudoparasitica 198 pseudosieboldii 189, 209 pteridiiformis 194 pulcherrima 190, 191, 207, 208, 209, 211, 212, 213 pulvinulifera 198, 209, 211 purpurascens 197, 198 Purpurascentes, sect. 183, 187, 197, 198 purpurella 196, 207, 209, 211, 212 Pycnopteris, group 189 Pycnopteris, subgen. 186, 187, 189, 190 Pycnopteris, subsect. 189 pycnopteroides 190, 199, 212 ramosa 194 X rarissima 207 redactopinnata 191, 211, 213 reflexosquamata 193, 210 reichsteinii 184, 191 remota 192, 208, 209, 210, 212, 213 Remotae, sect. 183, 186, 187, 191, 192 remotipinnulata 198, 208 rosthornii 191, 208, 209, 211, 213 rubripes 193 ruwenzoriensis 184, 194, 204, 205* ryo-itoana 196 sabae 198, 212 sacrosancta 197, 209 saffordii 194 sandwiciensis 198 X sardoa 206 X sarvelae 206 saxifraga 197 X saxifragi-varia 206 schimperiana 194, 204 schnellii 194 scottii 190, 209, 210, 212 x separabilis 206 sericea 193, 209 serrato-dentata 192 shibipedis 197 X shibisanensis 207 shiroumensis 194 x shuichangensis 206 sichotensis 192, 208, 212 sieboldii 188, 189, 208 sikkimensis 193, 211 simasakii 196 C. R. FRASER-JENKINS X sjoegrenii 206 sledgei 191 X slossoniae 206 sordidipes 197 sparsa 188, 197, 198, 211 sparsa X sparsa 207 spinulosa 195, 212 splendens 193 Splendentes, sect. 183, 187, 190, 193 sri-lankensis 198, 212 stenolepis 190, 209 stewartii 193 subarborea 194, 212 subcrenulata 198 subexaltata 197, 198, 209 subimpressa 194, 210, 212 sublacera 193, 208, 210, 211, 212 submontana 193 subpycnopteroides 184, 190, 199, 200* subreflexipinna 198 subtriangularis 196, 212, 213 X sugino-takaoi 207 X telesii 206 tenuicula 196, 209, 210, 212, 213 tenuipes 196 X tetsu-yamanakae 207 tingiensis 184, 192, 202, 203* xX tokudai 207 tokyoensis 191, 212 X toyamae 207 X tonensis 207 X triploidea 206 tsugiwoi 196 tyrrhena 192, 208 X uliginosa 206, 210, 212 unidentata 198 uniformis 193, 207, 208, 211, 213 varia 196, 197, 209, 210, 211, 213 Variae, sect. 183, 186, 187, 196 xX vidae 206 X vidyae 207 villarii 192, 208, 212, 213 viridescens 198, 212 xX wakui 207 wallichiana 188, 191, 199, 202, 208, 210, 211, 213 wardii 198 Xx watanabei 207 Xx wechteriana 207 whangshangensis 191 woodsiisora 191, 210, 212 x yamashitae 207 x yasuhikoana 207 yenpingensis 195, 196 yigongensis 191, 213 yoroii 198 zinongii 191 X zygo-parentalis 207 Elaphoglossaceae 184 Hypodematiaceae 184 Lastrea 184, 188 Arthrobotrys, subgen. 189, 191 Lastreopsis 185 Leptorumohra 185, 186, 197, 198 Lithostegia 185 Lomariopsidaceae 184 Lophodium 187 foenisecii 195 multiflorum 187, 195 Nephrodium 184, 187 marginale 187 monticola 192 Nothoperanema 185, 186, 195, 207 Onocleaceae 184 Peranemaceae 184 Peranemataceae 184, 185 Phanerophlebiopsis 186 Phegopteris 188 Polypodiaceae 184 Polypodium 188 filix-mas 188, 189 Polystichopsis 188 Polystichum 185, 188, 193, 195 Psidopodium 187 Pycnopteris 188, 189 sieboldii 188, 189 Rumohra 185, 186, 188 Acrorumohra, sect. 186, 197, 198 adiantiformis 185 aspidioides 185 Stenolepia 185 Stigmatopteris 185 Tectaria 184 Thelypteridaceae 184 Woodsiaceae 184 British Museum (Natural History) Ferns of Jamaica A guide to the Pteridophytes G.R. Proctor This flora records and describes the 579 species and 30 varieties of ferns occurring in Jamaica. The succinct species descriptions include relevant synonymy and incorpo- rate distributional data both within and outside Jamaica. Special emphasis is given to the subtle distinctions between closely related species and all genera are illustrated. Keys to the genera and species facilitate a wider use of the flora in the West Indies and northern South America. The author, one time Senior Botanist in charge of the Herbarium of the Science Museum, Kingston, Jamaica, is an outstanding field botanist and his expertise is reflected in the practicality of the flora and especially in the habitat and ecological information. This volume represents an important addition to our knowledge of the flora of the West Indies. 1985, 631pp, 135 line illustrations, 22 maps. Hardback. 0 565 00895 1 £50.00 Titles to be published in Volume 14 Cytological observations on Indian subcontinent and Chinese Dryopteris and Polystichum (Pteridophyta: Dryopteridaceae) By Mary Gibby A redisposition of the species referred to the ascomycete genus Microthelia By David L. Hawksworth A classification of the genus Dryopteris (Pteridophyta: Dryopteridaceae) By Christopher R. Fraser-Jenkins Evolutionary cladistics of marattialean ferns By Christopher R. Hill & Josephine M. Camus Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk Printed in Great Britain by Henry Ling Ltd, Dorchester , » 2) Bulletin of the British Museum (Natural History) Evolutionary cladistics of marattialean ferns Christopher R. Hill & _ Josephine M. Camus Botany series Vol 14 No 4 27 February 1986 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the Museum, both by the scientific staff of the Museum and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself, available separately, and individually priced. Volumes contain about 300 pages and several volumes may appear within a calendar year. Subscriptions may be placed for one or more of the series on either an Annual or Per Volume basis. Prices vary according to the contents of the individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History), Cromwell Road, London SW7 5BD, England. World List abbreviation: Bull. Br. Mus. nat. Hist. (Bot.) © Trustees of the British Museum (Natural History), 1986 The Botany series is edited in the Museum’s Department of Botany Keeper of Botany: Mr J. F. M. Cannon Editor of Bulletin: Mr J. R. Laundon Assistant Editor: Dr A. J. Harrington Editor’s Assistant: Miss M. J. Short ISBN 0 565 08010 5 ISSN 0068-2292 Botany series Vol 14 No 4 pp 219-300 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 27 February 1986 Evolutionary cladistics of marattialean ferns Christopher R. Hill ‘\ Department of Palaeontology, British Museum (Natural History), Cra DW i Roac C1 ondon- SW7 5BD Ne 6 ry ES OF eet Josephine M. Camus Department of Botany, British Museum (Natural History), Cromwell Road, London SW7 5BD Contents SIERRAS SUR SR RD OAR er ANT EE Pee UT n ey einen eT NEE a REA Ooi BES OT ie Rene i oe 219 NESE co on rene acetate edad sades wes cate oh daca suey UNals eoraacae Tie ei ade Casa e wees 221 ape INATIRNG CO PACOMSIIIG 55 oo oa.cs cicits cca con sno oad ad Chu ack bon ua INK 48. A Gana alee ens 222 PR 8 Se 19 Fase cco eo nna n a8 FUE ERs pecs tae ou tee ena ue OLED eMedia TEs bate eee tes ee Method of cladogram construction and introduction to terminology ............... 222 Rane Ce Ce IT LBRO 10) fy go cnn Sanctstcetins) ehaehentuniec dee Pa nese nentee eens 226 RM CNR ROIS 3 2.5. cored Gosctn 74 poSeC DE Tula ee vlavobs helen ty dec aabieaeehwales Cecuueveyes Z2r PCr e ME OE CR ATACICES ecco icisuniares oa eae aesten ie ened dad ae 2s Soak cheese aaa 228 BREE MABE LOE 5 foi6 08 555.54 oh oceans erases coc dase erepAds ees Ge asa ee een ARON AAG 228 MRIs COAT ACCES Ses cos gt oo ala oar ae Pasa oan eee eCOpRA OSSUR SIG Ee RAED 231 ERO MU CH CRAP OC LOEB ord s sete ccs wes ieidss sarees wa con ey nese aeeneee aeons 255 MitieaM? WRUUMEIE MII CUSCUSEIONY 55s oe fc acts dd iid Non aveu dh tae edocs buvunutn Ciancubehenssanieuatpeds 266 : Comparison of the cladogram with previous classifications ...................0.000008 266 Definition and relationships of the internested taxa ............ 0... .cceceeeeeeeeee eee es 22 SEIETINIEY OF FSAPOMIUS -FSCOLECODIETIS © 5 occ ccc ovo eoe renee ceeuetca gat oaiwees an eR 276 Definition of genera of extant Marattiales (Marattiaceae) ...................cceeeeeee 276 Provisional written classification of Marattiales .................cccececeeneeeeeeee seen 278 II. Phylogenetic relationships (evolutionary cladistics) .................ccececceceeceeeeeeeeeees 278 BRP oct cai y 225s one Crieverte vacunenacaca, Keck as pate es weer aaeeatecwe et At acanenneecnamnes 278 Principles of evolutionary cladistics (sequence analysis).............:s.0eceeeeeeeee ees 279 PreuMentation for invoOKing a tM AXIS'.>: 924525555 ojaser as accssnseeaarenires andes anes 281 PUPRNRTORUY Of TElOVANELOSSUS:. f500 Sr oscteauss Ge codeesna ee cioeohe cece a Seen eee 282 POET NY rk SF ccs cate sow se ws atid Sie ics Una Sale gua Malas ck serra 285 ig TAT Cos Date OTE OEE guERAY CRO) GENT RAR Stet oo SE Rey Cr plore RN AND Mere Ey gan 288 Peoey atid COMPparative ONYSIOIORY 6: .5 <2. 600. i ceseeinnsaissdadecesaSsn.uaeouteebeeed 290 Co Ca SAN BE Se RED CON leet oe SES PIT MAE SACRE Fe ree Men Aree PT eR ae 201 PNR ose oon nos see ice a a Or ee DATE Se AOR vw walt Se aulhiak See wdeas Wen cer meuoe eeeeeeNtees 291 MAAN MEDC SEUSCIITIO TTS SUUIIONN oo. Fn can ys oxo vcaavree ne dx cie ira pt eu diane ens tia Zaaeeee nee ATAN Ss HPaREAS 292 ae MMAR ECEIIOTIOS 055 ouice 2 ici Hs csoeedseavinniwbesdederg haps Rusa ai dedigeeoessinyetpuarnane tasers 294 SEN MNNTE nn aes ps.sn rien eL trode ta ees cara ihiaeoe Cee eabuae tera ahle Conde secareaye cleanest teantoes 294 Synopsis A new classification of the marattialean ferns is proposed, based on an overview of the characters of extant species compared with Ophioglossales and the fossil Psaroniaceae. The new classification shows consider- ably greater resolution of taxonomic relationships than previously proposed classifications, of which the scheme most widely adopted had no resolution of relationships and was therefore totally uninformative. The new scheme is not only relatively informative taxonomically, but is also consistent with independent evidence from ontogeny, biogeography, and stratigraphy of fossils. This was either not the case or was at best scarcely so with the previous classifications. The general principles for testing phylogenetically interpreted cladograms utilising independent evidence (‘evolutionary cladistics’ or ‘sequence analysis’) are discussed. Bull. Br. Mus. nat. Hist. (Bot.) 14 (4): 219-300 Issued 27 February 1986 C. R. HILL & J. M. CAMUS CLADISTICS OF MARATTIALES Zo Introduction In his incisive survey of ferns, Edwin Bingham Copeland (1947) drew attention to their unsatisfactory classification at ranks higher than the genus, particularly at family rank. Marat- tiales is no exception. At least nine family classifications have been proposed for the order in the past, of which the one embraced by most authors (Fig. 25A) is totally uninformative. It merely groups the seven extant genera into the order as a whole, with no further subdivision. Such a classification is uninformative because it fails to indicate relationships. This paper provides a general overview of extant Marattiales prior to more detailed taxonomic revision and is mainly about relationships. It concerns them in two senses, firstly (in part I of this paper) in terms of classification (taxonomic relationships), secondly (in part IT) ina wider, explanatory context, in terms of evolutionary history (phylogenetic relationships). To evaluate taxonomic relationships we use the now well-established method of biological classifica- tion termed cladistics. Cladistics is preferred to other methods of classification, such as phenetics, because for certain purposes it is superior to them: it has a greater potential for unambiguous definition of higher taxa and it has in particular a greater potential for detailed resolution of their relationships. Thus we provide a classification of Marattiales that enhances understanding of generic and higher taxonomic limits for this group and that has a demonstrably greater resolution of ranks than any previous classification. One of many aspects of these ferns requiring further study is the need for a more appropriate circumscription of the genera. This is briefly discussed and attention is drawn to the complex relation between methods of classification and pragmatic considerations of taxonomic judge- ment: cladistics in our view has considerable value in this regard but should not be misunder- stood. It has limits and provides a no more universally practical system for delimiting genera and species than any other method of classification. The most important taxonomic use of cladistics is for defining taxa at ranks higher than the genus and for assessing their relationships. In terms of phylogenetic relationships the Marattiales are of special interest. Not only are they the most primitive filicopsids surviving today, but also they are well represented by fossils from Palaeozoic times onwards. This makes them an ideal subject of study from the explanatory viewpoint of evolutionary history and relationship. The great resolution of taxonomic (particu- larly internested) relationships provided by cladistics is of particular relevance in this regard. In the context of general evolutionary theory such relationships may be interpreted as predicting relative ages of origin of the taxa. We show that these predictions for Marattiales, based in part on studies of the characters of living genera, are in fact corroborated by the relevant fossil (stratigraphic) evidence. Such evaluation of independent fossil evidence could not even be contemplated from the uninformative classification formerly so widely adopted, which, inter- preted in these terms, implied that all extant species of Marattiales arose at the same time. Indeed it is precisely such a consideration, set in the broader context, that makes cladistics of much greater empirical value than merely an alternative method of classification. Whilst Marattiales therefore provides an example for promulgating the theory and practice of evolutionary cladistics, these ferns are also inherently challenging. Although widely discussed at textbook level (Bierhorst, 1971; Eames, 1936; Copeland, 1947; Sporne, 1975) they are poorly known at the research level. As plants of tropical and warm temperate environments and therefore under some threat of extinction, this is particularly unfortunate. Thus, although we aim here mainly to provide an overview of the group, a number of new observations have been made. These range from characterisation of such features as the intercellular rods that occur on the walls of the spongy mesophyll cells to preliminary consideration of a possibly new species of Danaea, which is remarkably similar in habit to the Carboniferous fossil Psaronius. In addition some persistent errors are corrected, for instance the assertion that the walls of marattialean sporangia have stomata (Sporne, 1975). Definition of extant Marattiales has hitherto been rather vague, but their often huge size (with Frontispiece Crozier of Angiopteris evecta (G. Forster) Hoffm. C. D. Sayers 4, New Guinea (K. hort. 723-63.72306). Xc.0-7. 222 C. R. HILL & J. M. CAMUS the fronds of some species up to 7 m long being the largest of any fern), pulvini of the fronds and peculiar fleshy stipules at the base of the stipe, are well known (e.g. Copeland, 1947, Sporne, 1975), as in particular are the synangia of the genera Christensenia Maxon, Danaea Smith, and Marattia Swartz. Christensenia is limited in its geographic distribution to tropical forest clearings in south-east Asia and, with its ‘palmate’, reticulate-veined fronds, rounded synangia, and huge stomata (the largest of any vascular plant), it differs markedly from all other genera. The fronds of the other genera are pinnate with open venation and have elongate synangia or sori. Danaea, known only from the western hemisphere, has synangia deeply sunken in the lamina tissue, whereas the only pan-tropical genus, Marattia, has the synangia borne on the surface of the lamina, as is characteristic also of all the other genera (sometimes borne ona pedicellate placenta in this genus). The remaining genera, Macroglossum Copeland, Angiopteris Hoffmann, nom. cons., Archangiopteris Christ & Giesenhagen, and Protomarattia Hayata are limited in their distribution, like Christensenia, to the eastern hemisphere. All are normally free-sporangiate (i.e. objectively with true sori when mature), though those of Protomarattia show fusion of the dorsal walls of the sporangia so as to be partly synangiate. In summary, the aims of this study are 1. To assess the taxonomic relationships of extant Marattiales using cladistic method. 2. To compare this assessment with previous classifications, especially regarding previous family classifications and also with preliminary reference to generic circumscription. 3. To assess taxonomic relationships in general evolutionary (i.e. phylogenetic) terms of descent with modification, with reference to stratigraphic, ontogenetic, and biogeographic evidence provided both by the extant and the fossil members of the order. Brief reference is also made to the importance of ecological evidence in an evolutionary context. I. Taxonomic relationships Methods Method of cladogram construction and introduction to terminology This section is intended to introduce some basic concepts and terms for readers unfamiliar with cladistics. It is not intended to be a comprehensive summary, and fern workers wishing to learn more about the method should study Wiley (1981), Hill & Crane (1982), and Hennipman & Roos (1982, 1983). Readers already familiar with cladistics should start at p. 226. The method of cladistics' in the strict sense used here and also by most other workers today is entirely comparative: it is based solely on taxonomic characters and excludes reference to biogeography, ontogeny, or stratigraphy (see Hennig, 1966), though many authors still make some reference to ontogeny (e.g. Wiley, 1981; Hill & Crane, 1982) and to some extent this is unavoidable. The main object of the method is to produce a hierarchical classification in which taxa are defined by characters shared uniquely by their members and so far as possible not shared with any other taxon. In logical terms (Hill, 1981) the subclasses in the classification hierarchy are defined both necessarily and sufficiently by their characters, so that these ' Cladistics may be defined as a method of hierarchical biological classification in which groups are defined so far as possible only by reference to characters that distinguish a group from all others and that are shared by every member of the same group. For instance, the group ‘Seed plants’ encompasses all plants that share the character of the life-cycle ‘reproducing actually or potentially by means of seeds’. Fig. 1 A, Cladogram showing the relationships and defining characters (synapomorphies) of the cladistically natural taxa Spermatophyta and Angiospermae. Synapomorphies are shown as black bars. Synapomorphies abbreviated from Hill & Crane (1982), Hesse (1980, 1984) and Kubitski & Gottlieb (1984). The taxon ‘gymnospermae’ is not cladistically valid. It is a paraphyletic (grade) group. B, Diagram in three dimensions to show the same relations in terms of class inclusion. The subclass Angiospermae is internested within the more inclusive class Spermatophyta. Diagram A may be thought of as a vertical section of diagram B. C, Diagram showing relations of internesting of taxa. The relations of taxa 1, 2 and 3 are of class inclusion, i.e. of internesting, as is that of 4 to 1. The relation of taxon 4 to 2 or 3 is not an internested relation and cannot be compared in such terms. A-F are the individual members of taxon 1 and its various included sub-groups 2-4. Segment X is a paraphyletic taxon. Taxa 14 are Cladistically natural. CLADISTICS OF MARATTIALES 223 SPERMATOPHYTA "GYMNOSPERMAE' ANGIOSPERMAE lI | high reduction-oxidation potential pollenkitt pollen lacking prothallials pollen-receptive stigmatic surface pollen endexine not laminate 4-16 nuclei in megaprothallus embryo sac wall lacking sporopollenin double fertilisation companion cells in phloem sieve tubes reproducing by means of seeds (character A) PERMATOPHYTA 224 C. R. HILL & J. M. CAMUS characters in principle are individually sufficient for group definition*. Such characters are commonly termed synapomorphies (Hennig, 1965, 1966). Examples of cladistically valid, i.e. cladistically natural, taxa (Fig. 1A) are the taxon Seed Plants (Spermatophyta), which is defined uniquely as a taxon by the capacity of all its members to reproduce by means of seeds; and the Flowering Plants (Angiospermae), defined by at least ten uniquely shared characters, for instance double fertilisation and the occurrence of axially aligned companion cells in the phloem, derived during development from the same mother cells as the sieve elements (Hill & Crane, 1982, give eight of these ten Angiosperm synapomorphies, for the other two cited here see Hesse (1980, 1984) and Kubitzki & Gottlieb (1984)). Within the seed plants, the gymnosperms (‘Gymnospermae’) provide an example of a cladistically unnatural taxon. Gymnospermae is considered unnatural because these plants possess no uniquely shared characters (Fig. 1A). The group is only definable on a phenetic basis, by a combination of characters of two types: a) a character shared with other taxa, in this example with Angiosperms, i.e. character A, the capacity to reproduce by means of seeds; and b) characters that are lacking compared to such other taxa, i.e. the lack of all ten of the synapomorphies of Angiosperms. In other words, gymnosperms are plants that reproduce by means of seeds but do not have double fertilisation or companion cells in the phloem, and so on. Or still more plainly, such groups can be defined as groups only by a combination of the characters they have and those they do not have, and even the characters they have are not unique to them, such that the characters can only collectively be sufficient for group definition. Other botanical examples of such non-cladistic (‘grade’ or paraphyletic) taxa often referred to in current literature are cyanobacteria, psilophytes, rhyniophytes, trimerophytes, pteridophytes, progymnosperms, pteridosperms, proangiosperms and dicotyledonous Angiosperms (Bremer & Wanntorp, 1978; Hill, 1981a; Hill & Crane, 1982; Crane & Hill, in press; Hill, in preparation; Crane, in preparation). Characters such as character A of the gymnosperms, referred to above and in Fig. 1A, are too generalised in their distribution to function as synapomorphies at the rank in question (i.e. of gymnosperms) and are termed plesiomorphous or plesiomorphies (Hennig, 1965, 1966). Plesiomorphies are of no value in defining cladistically natural groups because they are characters applied at an inappropriately low rank, i.e. at too low a rank in the classification hierarchy to uniquely define a group. Whilst character A functions as a clear synapomorphy at the more inclusive rank of Spermatophyta (Fig. 1A), it is too generalised in its distribution to be cladistically valid for defining any particular sub-group such as gymnosperms or Angiosperms: it simply becomes redundant at these ranks for cladistically valid definition of those taxa and, at these ranks, is therefore a plesiomorphy. A cladistic classification is logically best expressed in a diagram that is three-dimensional (Fig. 1B) but for practical purposes is generally expressed diagrammatically in two dimensions as a dichotomously forking diagram termed a cladogram (Figs 1A, 23). Cladograms sensu stricto explicitly show a) the synapomorphies used to define the cladistic taxa and b) the taxonomic relations of the taxa, i.e. the pattern of and extent to which they form internested sequences (in Figures 1A and B the taxon Angiospermae is internested within the more inclusive taxon Spermatophyta). Such diagrams may also be summarised in written form (Eldredge & Cracraft, 1980; Wiley, 1979, 1981), in which case the synapomorphies themselves are in general no longer shown, though the internested relations of the taxa are carefully preserved by means of certain conventions (see Wiley, 1981 for a comprehensive account). It follows from the definition of cladistics given above that production of a cladogram involves a) observation of characters and b) an assessment of their relevance as synapomorphies. (This means that what is initially ‘observed’ to be a character may, after study, in some cases later cease to be recognised as a character, or at any rate as one of cladistic relevance. Platnick (1979: 542-543) comments appropriately that cladistic characters (synapomorphies) as such are not * In practice cladistics is not quite so simple, owing to the widespread occurrence of homoplasies and also to interpretative difficulties that can occur in recognising which characters are cladistic homologies (synapomorphies). The sentence in the text assumes, therefore, that characters have been diagnosed correctly as synapomorphies. For pertinent discussion of the use of characters in cladistics the reader is referred to Platnick (1979: 542-543). CLADISTICS OF MARATTIALES 225 observable as ‘given’ but are empirical, i.e. theory-laden, properties.) An important additional requirement is appropriate selection of the taxonomic units to be compared. These may range from subspecies to higher taxa, depending on the problem in hand, yet they are in each case (regardless of their rank) the lowest rank to be analysed. They are therefore generally termed terminal taxa, since they occupy the ends of the ‘branches’ of the cladogram and are of variable rank. They are equivalent in function to the operational taxonomic units of some authors (e.g. Sneath & Sokal, 1973) and are not necessarily themselves definable by having unique (autapo- morphous) characters, though many cladists insist on this requirement (see Hennig, 1966 vs Wiley, 1981; Hill & Crane, 1982; 308-310 and this paper p. 277 for a range of views). The choice of terminal taxa for Marattiales is discussed below. Assessment of the relevance of characters as synapomorphies requires study of the distribu- tion of characters amongst the terminal taxa selected. If two or more terminal taxa uniquely share the same character, that character is potentially a synapomorphy. As will be evident from Fig. 1 and the foregoing discussion of it, synapomorphy and plesiomorphy are relative, one to the other, depending on the particular rank in the hierarchy. Equally, vice versa, a particular synapomorphy determines a particular rank (Hill & Crane, 1982). The main initial requirement in practice is therefore to determine the relative generality of the distribution of characters amongst the terminal taxa. For reasons given by, for example, Hill & Crane (1982: 275-280) and others, this is usually achieved by reference to a group having relatively more generalised (i.e. plesiomorphous) characters in comparison to the group being analysed. Such a group is termed an outgroup (Stevens, 1980; Wiley, 1981; Watrous & Wheeler, 1981). In the present study, the order Ophioglossales provides an outgroup for Marattiales as a whole, and the family Psar- oniaceae (Psaronius + Scolecopteris) an outgroup for the extant Marattiales (as discussed below). The class inclusion relation provides the logic behind hierarchical classification. It involves (by definition) the idea of transformation®, from more general to less general characters in terms of their distributions. These characters respectively define more inclusive to less inclusive taxa, at respectively higher to lower ranks (Platnick, 1979). The inclusion relation necessarily involves hypotheses of polarity of characters into two or more states, from character states that are more generalised (—) in their distribution (relatively plesiomorphous character states) to those that are less generalised (+) in their distribution (relatively apbomorphous character states). Deter- mination of + vs — character states is based on the knowledge or assumption that most characters of the outgroup or outgroups relevant to establishing relationships are relatively generalised compared to those of the taxa being analysed in detail, in this case the extant Marattiales. Thus the spores of the outgroup Ophioglossales are typically round and trilete, the assumed — character state, whereas in some Marattiales they may be bean-shaped and monolete (regarded as the + character state). In drawing up a cladogram the individual characters nearly always prove to conflict to some extent with others as potential synapomorphies. For instance character 66 (Table 2, Fig. 23) is uniquely shared by (and thus is potentially a synapomorphy of) a group composed of Christen- senia + Danaea + Marattia, whilst characters 40 and 48 are (disregarding Psaronius + Scolecopteris) uniquely shared by, and therefore also potentially define, a group of Marattia + Macroglossum + Angiopteris + Archangiopteris + Protomarattia. These groupings are in conflict with one another over the appropriate higher taxon for Marattia. It could fit in either. Because of such conflicts it is heuristically appropriate to select the cladogram that most economically (parsimoniously) represents synapomorphies. This is done with reference to the characters considered as a whole (Wiley, 1981; Hill & Crane, 1982: 281-2). Essentially, a taxon defined by three characters as synapomorphies would be preferred to one in conflict with it supported by only two potential synapomorphies. Such a choice may be made in practice (as in > The term transformation here is used solely in a technical sense, such as that of mathematical transformations (in which, for instance, ellipses can be transformed into circles by decreasing the distance between the two foci). This meaning does not imply temporal transformation (see p. 281 for that different usage). Nor is the term used here in the same sense as Platnick (1979) in relation to schools of cladistic thought, though our views coincide with his (Platnick, 1979: 543). 226 C. R. HILL & J. M. CAMUS this study) by drafting up several alternative cladograms, totalling the number of conflicting characters in each case and finally selecting the diagram with the least number of conflicting characters: consistent, of course, with maximising taxa defined by synapomorphies. The residual conflicting characters, termed homoplasies, are indicated by asterisks in Fig. 23. Concentration on method of construction rather than on the aims of producing cladograms is not the main object of this paper (Feyerabend, 1978), and the foregoing is merely intended to briefly introduce terms and concepts for the general reader. However, it should be added that the method we have used (as with that of Hill & Crane, 1982) differs slightly from some others. Unlike Compatibility methods, for instance, (see e.g. Duncan, 1984) some of the data have not been initially discarded, though in the sense of minimising homoplasies (which represent potential synapomorphies) all methods eventually select from and to that extent exclude some of their data. Equally in the perceived interests of objectivity, our approach differs slightly from Parsimony method (e.g. Farris, Kluge & Eckhardt, 1970; Humphries, 1983; Churchill, Wiley & Hauser, 1984, 1985; Farris & Kluge, 1985). Unlike Parsimony method we have not treated characters which are near-synapomorphies, such as character 42, as if they were true synapo- morphies of more inclusive taxa (of Marattiales as a whole for character 42) by invoking reversals. For this reason the resulting cladogram is overall probably somewhat less parsimo- nious than might be obtained by such method. Further work is in progress on these important details of method. The main thrust of the present paper, however, is different, and claims that the efficacy of the various methods for producing cladograms can only be assessed meaningfully for the biologist in the broader context of evolutionary theory, i.e. in the light of empirical evidence independent of that used to produce a cladogram. Choice of terminal taxa Although more than 300 extant species of -Marattiales have been named, mostly in the genera Angiopteris and Marattia, a limited range of 23 species was selected to provide the terminal taxa for the cladogram. This selection was based on a background study of more than 150 named species and was designed to represent the main range of variation for each of the extant genera. For instance, although only five species of Marattia were selected (out of about 80 described species), these adequately represent the range of frond form and synangial structure for this genus (see Copeland, 1947, for relevant discussion). To cover the range of variation still more effectively, species at the boundaries of the genera as currently circumscribed were also selected besides those more characteristic of the genera; for example Angiopteris monstruosa Alderw. (intermediate between typical Angiopteris and Macroglossum) and A. annamensis C. Chr. & Tard. (intermediate between Angiopteris and Archangiopteris). Apart from the ‘filmy’ species of Danaea, D. trichomanoides, the other named species of Marattiales studied but not shown in the cladogram differ in no important way for the purpose in hand than those selected. Indeed, as Christensen (1905) and Holttum (1978) remark for Angiopteris, the 200 or more species recognised by some authors have been regarded by others as a single variable species: in reality there may perhaps be between 10 and 30 natural species of this genus. Species discrimination in Marattiales has in the past been excessively fine. Species have been separated on minute details that almost certainly in most cases represent merely intraspecific variation within a relatively few evolutionary species. At present this can only, however, remain a matter of opinion, since detailed field and laboratory study is an essential requirement for proper understanding of the range of variation of characters within the whole plants and within populations. What is absolutely certain is that the fragmentary herbarium material of many of the type specimens, and the sometimes inadequate descriptions based largely or entirely on such material, means that many described species are of questionable value. The numerous named species of De Vriese in De Vriese & Harting (1853) and Ching (1959), for example, are mostly concepts applied, like the name, to fragments of fronds on herbarium sheets rather than to whole plants or populations of whole plants. Unlike the exemplary work of Holttum (e.g. 1978) no useful account is made of variation in the field or of cultivation under uniform conditions. A major problem is the large size of many of these plants: herbarium sheets are too small to individually represent the range of variation of pinna or CLADISTICS OF MARATTIALES Dit pinnule form even within a single frond (often up to 7 m long!). Only a few collectors, notably A. H. G. Alston and C. J. Brooks, have tackled this problem realistically by collecting whole fronds and preserving them on an annotated series of sheets. As regards variation in the field and in populations raised in cultivation from spores, still less is known. Some of our terminal taxa are probably unsatisfactory species, though we have tried as far as possible to use ones that are well circumscribed. All the extant terminal taxa of Marattiales selected for the cladogram are species. However, the two outgroups Psaroniaceae and Ophioglossales are higher taxa. Information for the fossil Psaroniaceae (Stewart, 1983) was taken almost entirely from the literature. Several genera of fertile fronds and axes are now included in this family but we have limited our choice to the best known genera, Psaronius Cotta and Scolecopteris Zenker emend. Millay (1979). Psaronius is a genus of stem material attributed to the same whole plant as Scolecopteris (which is a genus of fertile frond material). We refer informally to the reconstructed fossil plant, which was a tree fern in habit, as “Psaronius + Scolecopteris’. Since there is considerable variation amongst the species of these genera there are several —/+ ([d4) entries for them in the data matrix (Table 2) and cladogram (Fig. 23). Choice of outgroups Amongst extant plants the nearest outgroup to Marattiales is probably Ophioglossales (Bierhorst, 1971; Sporne, 1975). These plants are in some respects similar but in many are very different: for instance their ‘aerial complexes’ (Bierhorst, 1971) and exannulate sporangia, each sporangium having its own individual vascular supply, are more readily compared with ‘trimerophyte’ or ‘aneurophyte’ branching systems (Stewart, 1983) than with the laminate fronds and sori, composed of more or less annulate sporangia, possessed by most filicopsids. Nonetheless, they can be used as an outgroup for Marattiales as a whole, and so far as it is possible to assess homologies, we have referred to them as the main outgroup for the order. In relation to extant Marattiales however, Psaroniaceae is by far the most similar group, even though exclusively fossil. Although Psaroniaceae has a number of unique characters it has several generalised characters compared to extant Marattiales, yet is more closely related to them than to Ophioglossales by at least three characters of cladistic value. It may be regarded as a plesiomorphous sister group of Marattiales and thus as an outgroup (Stevens, 1980; Watrous & Wheeler, 1981). We have found Psaroniaceae of considerably more use than Ophioglossales in evaluating relationships amongst extant Marattiales. We emphasise that Psaroniaceae is used as an outgroup purely in terms of character comparison: its Upper Carboniferous to Permian age is of no importance in assessing taxonomic relationships. Also, the range of characters used for the living genera has not in any significant way been narrowed to facilitate comparison with these fossils (cf. Patterson, 1981a). An estimate of relative completeness of knowledge of characters of the fossil vs the living genera (information gap index) was calculated as follows. Using Table 2, the number of characters for Psaronius + Scolecopteris excluding those not applicable was totalled (69). This was divided into the number of characters for which information may potentially become known but which are at present unknown because of imperfect preservation (e.g. of root hairs) or incomplete study (e.g. the surface cell layer of the sporangium wall). Such characters for Psaronius + Scolecopteris in Table 2 are represented by blanks and total 9. Multiplication x 100 gives a percentage figure, the information gap for the fossil compared to its extant relatives: 9/69 x 100 = 13% for Psaronius + Scolecopteris, a remarkably small gap. The position is considerably less favourable for other fossils discussed below, and for which therefore no formal inclusion in the cladogram has been attempted. This index may equally be applied to lesser known extant species in comparison with better known ones. Thus for Danaea sp. A the index is currently 28/71 x 100 = 39%, for Archangiop- teris itoi Shieh 16/69 x 100 = 23% and for Angiopteris elliptica Alderw. 9/68 X 100 = 13%. These figures indicate that gaps in knowledge for some of the extant species may equal or even considerably exceed that for the fossil outgroup. 228 C. R. HILL & J. M. CAMUS Observation of characters Herbarium specimens have provided the main material and for many species were the only source of material for direct study currently available to us. As far as possible the type specimens were examined as well as other material of a species. The specimens studied for the cladogram are listed in the section “Taxa and specimens studied’, p. 292, others are indicated in the text. Observations on the dried material were supplemented both by spirit-preserved material at the British Museum (Natural History) (BM) and Kew (K), and with reference to live plants mainly at K, including Macroglossum which has recently been brought into greenhouse cultivation there. JMC was able to examine Christensenia, Angiopteris and Marattia in the field during a visit to Borneo in 1980. Several methods were used to observe the anatomy of the herbarium specimens. The scales, frond hairs, and root hairs were removed and mounted in glycerin or gum chloral. Epidermal characters were examined by clearing pinnae or pinnules in a 10% aqueous solution of commercial bleach (‘Domestos’) for 6 to 24 hours, the exact time depending on ambient temperature as well as the resistance of the lamina to bleaching and the freshness of the bleach. After rinsing well in distilled water the preparations were stained with either gentian violet in ethanol or an aqueous solution of methylene blue. Surface details of synangia and sporangia were studied similarly, using 10% Domestos to bleach and restore them to the shape shown in life. (That the restored shape matches closely that in life was checked for a living plant of Angiopteris at K). After thorough rinsing the bleached material was stained with 1% safranin in ethanol and examined by optical microscopy. Sporangia and synangia were also examined in the air-dry and critical-point dried states using scanning electron microscopy. SEM was used to examine the projections on the walls of the mesophyll cells, and the spores (which were examined without acetolysis). Sections of rhizome from the herbarium specimens were mostly made by hand-razor (in one instance requiring a hack-saw), and were stained with phloroglucinol + concentrated hydro- chloric acid. Serial sections of fertile pinnules were made by microtome sectioning of herbarium material rehydrated in approximately 1% aqueous tri-sodium orthophosphate. Standard de- hydration and infiltration procedures were used to embed the material in wax, staining with safranin and fast green or alcian blue. Sections were mounted in euparal. Permanent slide preparations and the SEM stubs are in the Department of Botany, BM. List of characters This section lists the characters used to construct the cladogram of Marattiales shown in Fig. 23, using the method outlined above. For each character the character state considered to be relatively plesiomorphous (generalised) is given first, connected by an arrow (—) to the relatively apomorphous (less generalised) character state. This information applied to the terminal taxa selected for the cladogram is summarised in Table 2, which records the characters as — for the generalised state and + for the less generalised (derived) one. Where an entry fora character is not applicable this is indicated by /. For the reconstructed fossil plant Psaronius + Scolecopteris blank entries in Table 2 indicate characters for which information is as yet unknown owing to incomplete preservation or study. For the extant taxa, characters of this status are represented by a predicted state, which is indicated by enclosure within parentheses. Several of the characters listed require explanation and discussion. This is dealt with on p. 233-266, preceded by a discussion of certain characters that were not used for the cladogram on p. 231-233. 1. Stelar anatomy of adult stem: solenostele — dictyostele. (Kuhn, 1889; Farmer & Hill, 1902; West, 1917b; Schoute, 1926; Chang, 1975; DiMichele & Phillips, 1977; Millay, 1979.) 2. Number of cycles in dictyostele as seen in T.S. of adult stem: one (monocyclic) — three (tricyclic) > more (polycyclic). (Kuhn, 1889; Shove, 1900; Farmer & Hill, 1902; Gwynne-Vaughan, 1905; Campbell, 1911; West, 1917b; Schoute, 1926; Stidd & Phillips, 1968; Chang, 1975; DiMichele & Phillips, 1977; Rothwell & Blickle, 1982; Mickle, 1984.) No 1s. 16. CLADISTICS OF MARATTIALES 229 Meristeles in T.S. of adult stem: in clearly visible cycles, distinct from a helix — cycles not clearly distinguishable from a helix. (Kuhn, 1889; Shove, 1900; Farmer & Hill, 1902; Gwynne-Vaughan, 1905; Campbell, 1911; West, 1917b; Schoute, 1926; Morgan, 1959; Stidd & Phillips, 1968; Chang, 1975; DiMichele & Phillips, 1977; Rothwell & Blickle, 1982; Mickle, 1984.) Protophloem of stem meristeles: external to the metaphloem (i.e. exarch) — internal to metaphloem (endarch). (Shove, 1900; Brebner, 1902; Gwynne-Vaughan, 1905; Campbell, 1911; West, 1917b; Esau, 1969; Chang, 1975; Smoot & Taylor, 1981; Smoot, 1984.) Stipules borne in pairs at base of rachis: absent — present. (Christ & Giesenhagen, 1899; Hayata, 1919; Ching, 1958a; Morgan, 1959; Bierhorst, 1971; DiMichele & Phillips, 1977; Johns, 1981; Smoot & Taylor, 1981.) Pulvini in frond: absent — present. Mucilage canals in stem: absent —> present. (Kuhn, 1889; Gwynne-Vaughan, 1905; Campbell, 1911, 1914b; West, 1915; Stidd, 1971; Chang, 1975; DiMichele & Phillips, 1977; Rothwell & Blickle, 1982.) Mucilage canals in root: absent > present. (Kuhn, 1889; Gwynne-Vaughan, 1905; Campbell, 1911; Morgan, 1959; Chang, 1975; Ehret & Phillips, 1979.) Mucilage canals in axial components of frond: absent —> present. (References as for character 8; also Brebner, 1902; Stidd, 1971.) Mucilage canals following course of lateral vein in lamina beneath sorus: absent —> present. (Campbell, 1914a, b.) Sclerenchyma in stem cortex: absent — present. (References as for character 2, excluding Schoute, 1926, and including Goeppert, 1864-5; Kuhn, 1890; Brebner, 1902; Stidd, 1971; Smoot & Taylor, 1981.) Sclerified pith in stele of root: absent — present. (West, 1917b; Ehret & Phillips, 1977.) Continuous band of sclerenchymatous fibres in cortex of aerial or subterranean root: absent — present. (West, 1917b; Morgan, 1959; DiMichele & Phillips, 1977; Ehret & Phillips, 1977; Rothwell & Blickle, 1982.) Apparently sclerenchymatous cells occurring externally to cylindrical traces in stem or rachis: absent > present. (Shove, 1900; Brebner, 1902; Gwynne-Vaughan, 1905; Campbell, 1911; Morgan & Delevoryas, 1952a, b; Morgan, 1959; Stidd, 1971; DiMichele & Phillips, 1977.) Multicellular root hairs: absent — present. (Brebner, 1896; Campbell, 1911; West, 1917a.) Roots numerous and supported by parenchymatous tissue surrounding the mature stem at least towards its basal region: absent —> present. (Morgan, 1959; Stidd & Phillips, 1968; Ehret & Phillips, 1977; Smoot & Taylor, 1981; Rothwell & Blickle, 1982.) Habit of adult stem: erect — creeping. (Mettenius, 1856; Campbell, 1911; West, 1917b; Morgan, 1959; Shieh, 1970; DiMichele & Phillips, 1977; Rothwell & Blickle, 1982.) Length of erect adult stem: squat — elongate. (References as for character 17.) Symmetry of adult stem: radial or scarcely dorsiventral — dorsiventral. (Shove, 1900; Campbell, 1911; West, 19176; Morgan, 1959; DiMichele & Phillips, 1977; Rothwell & Blickle, 1982.) Extent to which mature fronds are pinnately divided: 3 or more-pinnate — 2-pinnate. (Stidd, 1971; Croft, pers. comm. 1983.) Extent to which mature fronds are pinnately divided: 2-pinnate — 1-pinnate. Extent to which mature fronds are pinnately divided: 1-pinnate — simple. Extent to which mature fronds are pinnately divided: pinnate — palmate. More or less spiny transverse ridges on stipe of mature fronds: absent — present. Soft spines on veins of pinnules: absent — present. (Stidd, 1971; Stidd & Phillips, 1980.) Stipe of juvenile frond: without a pulvinus > with one pulvinus a short distance proximally to laminate region of frond. (Christensen & Tardieu, 1935.) 230 C. R. HILL & J. M. CAMUS Stipe of adult frond: without a pulvinus — with one pulvinus a short distance proximally to laminate portion of frond. (Christ & Giesenhagen, 1899; Hayata, 1919, 1928; Ching, 1958b; Shieh, 1970.) Number of pulvini per stipe in adult frond: never more than 1 — some fronds with 2 or more. (References as for character 27.) Microscopic projections from exterior of walls of mesophyll cells: absent — present. (De Vriese & Harting, 1853; Luerssen, 1873a, b, 1875b; Schenk, 1886; Kuhn, 1889.) Stomata of lamina: guard cells functional — permanently open. (Campbell, 1911; Scott & Holden, 1933.) Stomata of lamina: anomocytic or weakly cyclocytic > more or less strongly cyclocytic. (Scott & Holden, 1933; Van Cotthem, 1970.) Stomata of lamina: subsidiary cells typically dicyclic — typically dicyclic but with lateral subsidiary cells in a paracytic position. (Van Cotthem, 1970.) Stomata of lamina: subsidiary cells typically dicyclic — typically tri- or tetracyclic. (De Vriese & Harting, 1853; Luerssen, 1873a; Van Cotthem, 1970.) Diameter of stomatal apparatus (excluding subsidiary cells): in general < 80 wm — in general > 80 pm. (Scott & Holden, 1933; Van Cotthem, 1970.) Anticlinal walls of ordinary epidermal cells (adaxial epidermis): more or less straight, curved or at most very coarsely sinuous — uniformly and markedly sinuous. (References as for character 32, including Pant & Khare, 1969.) Idioblasts in abaxial epidermis: absent > present. (Christ & Giesenhagen, 1899; Van Cotthem, 1970; Ogura, 1972.) Lamina margin of pinnae or pinnules, excluding apex: distinctly toothed or lobed — scarcely and inconspicuously toothed or lobed. (Morgan & Delevoryas, 1952a; Millay, 1979, 1982a, b.) Fertile pinnae or pinnules: barely recurved — recurved fully to protect sori. (Morgan & Delevoryas, 1952a; Millay, 1979; 1982a, b; Stubblefield, 1984.) ‘Fibre’ bundles alternating with and parallel to veins of lamina: absent — present. (Campbell, 1914a; Millay, 1979, 1982a, b.) Scales composed of: more or less isodiametric cells — elongate cells. (Scott & Holden, 1933; Stidd & Phillips, 1968; Stidd, 1971.) Depth of anticlinal walls of cells of scales, in scales with anticlinal walls more or less isodiametric as seen in surface view of the scale: deep — shallow. Margin of scales: more or less entire — lobed and/or fimbriate. (Scott & Holden, 1933; Stidd, 1971.) Division of scale margin: lobed —> fimbriate. Uniseriate multicellular hairs on frond: absent > present. (Scott & Holden, 1933; Millay, 1979, 1982a, b.) Veins of lamina: not normally reticulate — reticulate. (Millay, 1979, 1982a, b.) Veins of lamina: free to margin > anastomosing near margin to form a submarginal bundle. Sterile and fertile fronds: monomorphic — distinctly dimorphic. Uniseriate multicellular branching hairs attached to placenta: absent — present. (Millay, 1979, 1982a, b.) Scales or hairs borne on synangial or sporangial wall: absent — present. (Millay, 1979, 1982a, b.) Tissue of lamina between sori: scarcely expanded — expanded ™%4 of the way up the height of the synangia Or sporangia. (Morgan & Delevoryas, 1952a; Millay, 1979, 1982a, b.) Placental tissue: not incorporating tracheids — incorporating them. (Luerssen, 1871-4, 1875a; Strasburger, 1874; Bierhorst, 1971; Millay, 1979, 1982a, b; Stubblefield, 1984.) Placenta: without > with apparent epidermis. (References as for character 51.) Placenta: relatively weakly developed — relatively well-developed. (Millay 1982a; Stubblefield, 1984.) Sporangia: individually vascularised by one trace — sorus of several sporangia (or synangium) collectively vascularised by one individual trace. CLADISTICS OF MARATTIALES Zou 55. Arrangement of sori in ultimate divisions of frond: two rows parallel to the midrib > approximately in two rows but rows parallel to the main lateral veins. (Morgan & Delevoryas, 1952a; Millay, 1979, 1982a, b; Stubblefield, 1984.) 56. Arrangement of sori in ultimate divisions of frond: in two rows — relatively scattered. 57. Position of sori relative to margin: near to margin > midway between midrib and margin. (Morgan & Delevoryas, 1952a; Millay, 1979, 1982a.) 58. Symmetry of sorus in plan view: in general radial — in general elongate. (References as for 57, including Millay, 1982b; Stubblefield, 1984.) 59. Spore-bearing loculi of synangia (or sporangia of sori) packed together: loosely — closely. (Millay, 1979.) 60. Composition of surface cell layer of walls of sporangium or synangium: all walls composed of more or less isodiametric cells — lateral (and ventral) walls of sporangium, or synangial septa between locules, composed of elongate cells. (Millay, 1979, 1982a, b.) 61. Alignment of cells of ventral wall of sporangium or synangium with dehiscence slit: parallel — oblique. (Millay, 1979, 1982a, b.) 62. Group of more or less isodiametric, thick-walled cells at apex of sporangium or synangial locule: absent — present. (Millay, 1979, 1982a, b.) 63. Number of sporangia or locules per sorus typically: > 10 > < 10. (References as character 57, including Millay, 1982b; Stubblefield, 1984.) 64. Ventral walls of opposing rows of sporangia or locules in the sorus: free — adnate. (Morgan & Delevoryas, 1952a; Millay, 1979, 1982a, b.) 65. Ventral walls of sporangia or of locules of synangia: laterally partially adnate — laterally free. (References as for character 64.) 66. Ventral walls of sporangia or of locules of synangia: laterally partially adnate > completely adnate. (References as for character 64.) 67. Dorsal walls of sporangia or of locules of synangia: laterally partially adnate — laterally free. (References as for character 64.) 68. Dorsal walls of sporangia or of locules of synangia: laterally partially adnate — laterally completely adnate. (References as for character 64.) 69. Dehiscence of sporangia or synangial locules: by longitudinal slits > pores. (References as for character 64.) 70. Shape and tetrad mark of spores (where visible): typically round and trilete — typically elongate and monolete. (References as for character 64, including M. D. Crane, 1973; Millay, 1976; Stubblefield, 1984.) 71. Exine ornamentation: fundamentally granular or warty — with cones or spines. (References as for character 70.) 72. Exine ornamentation elements: more or less free — commonly fused laterally, e.g. warts forming rugulae. (References as for character 70.) 73. Exine spines: in general simple — bifurcating or further branched. (References as for character 70.) Excluded characters Although we have attempted to study as many characters as possible, several have been excluded from the data matrix and cladogram, some because they proved too generalised (plesiomorphous) to be of use, such as the mesarch to endarch differentiation of protoxylem in the meristeles of the stem. This character state is shared with Ophioglossales and differs from the exarch maturation of most ‘pteridophytes’ (Ogura, 1972). In relation to ‘trimerophytes’ (which are centrarch) the condition in Ophioglossales and Marattiales is relatively derived but appears to be a grade character rather than one of genuine cladistic value, i.e. it is comparable to the gymnospermous xylem characters of ‘progymnosperms’, an unnatural group. This argument for exclusion applies in particular to the several generalised characters of the gametophyte, some of which are shared with bryophytes. A few characters were considered but ultimately rejected because no sense could be made of them. One such character is the extent of elongation of the meristeles as seen in T.S. of the adult 232 C. R. HILL & J. M. CAMUS stem. Characteristically these form tangentially elongate bands in Psaronius, giving relatively few gaps per cycle whilst in extant Marattiales the meristeles are mostly strands, short in T.S. and generally with more gaps per cycle. However, the work of Morgan (1959) and others shows that although some species of Psaronius (e.g. P. melanedrus Morgan) have all the meristeles tangentially elongated, in some (e.g. P. brasiliensis Brongn. and P. magnificus (Herzer) Rothwell & Blickle) those of the inner cycles are of varied shapes and can be short (G. W. Rothwell, pers. comm.). It may also be noted that Mettenius (1864) records an interesting individual of Angiopteris that died after enduring unfavourable conditions of cultivation. The meristeles were almost continuous laterally in the upper part of its stem, as in Psaronius melanedrus. A few characters were considered initially but later rejected through lack of uniformity of data. This applies particularly to biochemical information, mainly on flavenoids but also on a few other chemicals (e.g. Wallace, Story, Besson & Chopin, 1979; Wallace, Yopp, Story, Morris, Gomez & Stone, 1979; Wallace, Yopp, Besson & Choppin, 1981; Wallace, Markham, Giannasi, Mickel, Yopp, Gomez, Pittillo & Soeder, 1982; Richardson, 1984). It appears that Marattia and Angiopteris have been examined frequently but with only patchy reference to a few of the other genera, thus rendering this aspect at present of little use for the present study. Since herbarium material is in general adequate for such work, sampling could readily be made more representative in the future and we have sent samples to Dr M. Richardson with this in mind. The position is different for certain other kinds of information such as karyotype analysis, certain aspects of ontogeny, comparative ecology and physiology, and interbreeding behaviour. Knowledge of these for Marattiales is at present scanty, partly because live plants are an essential requirement for their study. Although chromosome numbers are known for all genera except Archangiopteris and Protomarattia, thorough karyological study has only recently been initiated. Interesting aspects of ontogeny remain unexplored, such as whether the sori are already developed in the crozier stage of frond development, whether the stipules can be understood as reduced pinnae, and the development of pulvini in the stipes of juvenile fronds of sporelings. Comparative physiological work is, so far as we know, at a primitive stage of progress on Marattiales, as also is experimental breeding and knowledge of their ecology. With such longer-term possibilities in mind we are currently cultivating gametophytes of Marattia, Macroglossum and Angiopteris. Considering this against a background of continuing indigenous habitat destruction we have also attempted, with the help of colleagues, to bring representative material into greenhouse cultivation. Marattia and Angiopteris have been cultivated widely for a century or more, but other genera much less so. Danaea and Christen- senia have been grown at K for some years, though optimum conditions for growth have yet to be achieved in this country and the plants remain at present entirely vegetative (Danaea has been grown successfully in N. America (J. M. Sharpe and J. E. Skog, pers comm.)). Recently, however, Macroglossum (collected by T. G. Walker and A. C. Jermy in 1978) has been very successfully cultivated at K, and from there is being actively distributed internationally. At the time of going to press, live plants of Archangiopteris henryi var. somai had just been received at BM from Taiwan, through the kindness of Tsai Linn-lai, and they appear to be growing satisfactorily at K. Protomarattia, which came from northern Vietnam and has not, to our knowledge, been collected for some 50 years, may well be extinct or almost so; if it survives, plants in cultivation would be scientifically invaluable. We should add that at any rate for species of large size, the small plants produced adventitiously on the stipules are usually sufficient for successful propagation of these ferns. The character entries in parentheses in Table 2 indicate those for which the character state was merely predicted. Such instances, together with scant knowledge of their ecology, reveal a need for more thorough fieldwork on these plants than has formerly been the case (with the notable exceptions of collecting by A. H. G. Alston, C. J. Brooks, and R. E. Holttum). Sufficient information about the larger sized species has rarely been recorded (or material collected), and most existing herbarium specimens are mere fragments of the frond alone. These frequently give no information on even such basic characters as the outline of the lamina or degree of pinnation, nor of the stem, stipule and root form and structure. The internal tissue of the base of the stipe, and of the stipules and stem, is sometimes coloured a striking deep purple, a feature rarely noted CLADISTICS OF MARATTIALES 233 by collectors. Thus its true extent cannot be judged exactly. This applies also to the extent of variation of characters from plant to plant within a stand, and from stand to stand. Only such work, combined with cultivation under uniform conditions and study of variation in sporelings, is likely to reveal the true extent of intraspecific variation and thus to assist in verisimilitudinous discrimination of species-level differences. Evolutionary study (as opposed to speculation) in a Darwinian (adaptationalist) context is precluded without a sophisticated knowledge of physiology and ecology. Discussion of characters This section refers to characters requiring explanation and discussion for several reasons: (a) of observations that are new; (b) where accounts in the literature are scattered and in some cases confused, and thus in need of synthesis or correction; (c) where our choice of polarities of the character-states requires clarification, particularly in comparison with the outgroups Psaronius + Scolecopteris and Ophioglossales; (d) to set the 23 terminal taxa chosen for the cladogram into the context of the broader sample of species studied as background material. The object of the discussion is to attempt to clarify exactly what we have done and thus to facilitate criticism and improvement by others in the future (Stevens, 1980). 1. Stelar anatomy of adult stem. Despite the work of Beck, Schmid & Rothwell (1983) and Schmid (1983) in the United States and of Sporne (1975) in England there is as yet no single clear system for classification of all steles of vascular plants. Sporne’s system is in general simpler, it is more explanatory in biological terms and to that extent is preferable, but the American system is easier to apply as a classification in certain instances. The work of Kuhn (1889), Farmer & Hill (1902) and West (19175) suggests that the stele of Christensenia may best be described as a perforated solenostele sensu Sporne (Ogura, 1972, refers to it as a dictyostele), though further work is needed. Steles of adult stems of Ophioglossales are variable and mostly defy straightforward classifica- tion but may be considered to range from medullated protosteles through a solenostele to rudimentary perforated dictyosteles sensu Sporne (Bierhorst, 1971; Ogura, 1972: 285-7; Sporne, 1975). Millay (1979: 55-56) discusses Psaronius which also has a wide range of stelar anatomy. The range of adult stelar anatomy of the stem in Ophioglossales, Psaronius and extant Marattiales is similar in each taxon, though the occurrence of simple medullated protosteles in Botrychium (which we consider the most plesiomorphous genus of Ophioglos- sales) suggests that the simpler steles should be considered plesiomorphous in these taxa. This polarity is consistent with ontogenetic evidence. The alternative, however, that a dictyostele is the generalised condition for Marattiales (with consequent reversal to a solenostele in Christen- senia) cannot be entirely excluded as a possibility. This polarity might well be selected by Parsimony method. 2. Number of cycles in the dictyostele as seen in T.S. of adult stem. Information on this and other characters of the stem is meagre because stems of the larger sized plants, the size of barrels, are rarely collected. Living material in cultivation is so scarce that few plants have been sacrificed for anatomical study. The few anatomical observations that have been made, published some time ago, are of limited use because the ontogenetic stages of the individuals studied were varied and the exact stage was rarely specified. This provides a poor basis for comparison of equivalent developmental stages, termed semaphoronts (Hennig, 1966) and which is a basic requirement of cladistics. In some of the early studies only young stages were examined, for example of Macroglossum (Campbell, 19146) and Marattia alata Sw. (Charles, 1911). Gwynne-Vaughan (1905) and Chang (1975) examined stems of Archangiopteris henryi Christ & Giesenh., and Hayata (1928) illustrated the vascular system of A. henryi var. somai (Hayata) Tagawa (under the name of Protangiopteris somai Hayata). Each study shows a monocyclic dictyostele but with an ‘internal strand’ (Gwynne-Vaughan, 1905) or ‘connecting vascular bundle’ (Chang, 1975). These steles are treated here as monocyclic (i.e. dictyosteles sensu stricto of Schmid (1983)), although clearly incipiently dicyclic. 234 C. R. HILL & J. M. CAMUS Sectioning of the apparently new species of Danaea referred to in this paper as Danaea sp. A (Figs 2, 3), revealed a dicyclic stele (Fig. 4A). The steles of adult stems of Ophioglossales are never more than monocyclic and thus they provide an unambiguous basis for the polarity adopted here. Their occasional internal strands of xylem tissue are considered to be of no biological significance (Ogura, 1972: 287). Fig. 2. Danaea sp. A., P. J. Edwards & J. R. Woodhams KER 1026, Guyana (Kew hort. ref. 174-78.01484): showing erect growth habit, pulvini (p, character 6), and mid-stipe pulvini (msp, character 28). x 0-5. Fig.3 Danaea sp. A., P. J. Edwards & J. R. Woodhams, KER 1026, Guyana (BM pteridophyte spirit collection B.72, Kew hort. ref. 174-78.01484): showing aerial production of roots. The horizontal part of the stem (arrowed) is the original portion collected in Guyana where it grew erect. It was pinned horizontally on the soil surface by the gardener at Kew. The new growth resumed the erect habit. Scale bar represents 5 cm. 235 CLADISTICS OF MARATTIALES 236 C. R. HILL & J. M. CAMUS 3. Meristeles in T.S. of adult stem. The monocyclic steles of Ophioglossales could hardly be otherwise than cyclic and thus are not considered applicable for determining polarity of this character. Psaronius + Scolecopteris is therefore used as the outgroup. 4. Phloem maturation of stem meristeles. In Ophioglossales, protophloem cannot usually be distinguished from metaphloem but where evident it is exarch as in most other ‘pteridophytes’ (Esau, 1969; Ogura, 1972). The endarch phloem maturation of Marattiales is therefore considered apomorphous compared to the outgroup. ae 3 i D ‘ 7 : : Fig. 4 A, Transverse section of stem of Danaea sp. A., P. J. Edwards & J. R. Woodhams KER 1026, Guyana (BM pteridophyte spirit collection B.72, Kew hort. ref. 174-78.01484): dicyclic dictyostele (character 2) in centre, with meristeles of outer cycle in short arcs, mucilage canals (m, character 7), sclerenchyma in hypodermal layer of stem (s, character 11), in root pith (s, character 12) and in root cortex (s, character 13). Traces to a frond are visible at right. x 12-5. B, Detail of A to show sclerenchyma in hypodermal layer of stem (s, character 11). C, D, Danaea alata Smith, A. C. Jermy 10936, Trinidad (BM): multicellular root hairs (character 15). D shows the nuclei clearly. C x 150, D x 120. Fig.5 Stipules of extant Marattiales (character 5). A, B, Angiopteris sp., A. Paterson 362, Sumatra (K. hort. 118-74.01395): showing shape of the stipules, their fimbriate-lacerate margin and lenticels. Note also the elongated lenticels on the frond stipes. A also illustrates the broad base of the stipules, and B the pair of stipules clasping the swollen basal region of the rachis. A x c.0-70; B x c.0-75. C, Angiopteris hypoleuca Vriese, origin unknown (K. hort. 000—73.13071): stipules intimately shielding the unfurled laminate region of young fronds. In this species, at the stage of frond development when photographed, the abaxial and lateral regions of the rachis were left exposed. Note the differently divided margin and surface texture compared to the stipules in A and B. X c.0-85. n ca) J ne) > < < = a oe) Nn oO — ee 2 Q < a S) 238 C. R. HILL & J. M. CAMUS 5. Stipules (Fig. 5). These are expanded flap-like organs attached in a pair to the stem at the extreme base of each rachis. On anatomical grounds they are considered to be part of the frond (Bierhorst, 1971). Their shape is orbicular to oval, attached by a broad base (Fig. 5A). Numerous vascular traces enter the base and, where clearly seen (e.g. in Danaea) they run parallel (with slight divergence from one another) to the margin. Such venation may be described as approximately cyclopteroid but details are poorly known. The free margin of the stipules ranges from more or less entire to uniformly fimbriate (Fig. SC); sometimes it is more unevenly divided (Fig. 5A, B), and may be divided into broad lobes or even become broadly crenulate (Frontispiece). The divisions are never very deeply incised. Further work on the range of stipule form and structure of extant Marattiales is in progress. In the young crozier the stipules intimately clasp the unfurled laminate region of the frond, sometimes leaving the abaxial and lateral region of the rachis exposed (Fig. 5C). They thus function to shield the laminate region, which at this stage is very delicate and presumably is appetizing to predators. At an earlier stage of stem development the stipules also function to protect the next frond developed (Bierhorst, 1971) whilst later,.after unfurling of the crozier, they more or less tightly clasp the swollen basal region of the rachis (Fig. 5B; Shove, 1900; Brebner, 1902; Ogura, 1972). They persist permanently on the stem after detachment of the fronds. We have found that the cells of the stipules contain abundant starch grains. Stipules are therefore recognised here as a form of trophopod, sensu Wagner & Johnson (1983). It is widely accepted that all extant Marattiales possess stipules, though for many described species they have not been recorded by the authors and have rarely been collected. Since their form is constant for a plant but varies from plant to plant, particularly in the detailed characters of the margin, they may be useful for discriminating species (Holttum, 1978). The term ‘stipule’ has also been applied to the sheathing structure formed from the expanded basal region of the ‘aerial complex’ (sensu Bierhorst, 1971) in Ophioglossales. Differences from Marattiales are as follows: (a) In Ophioglossales the so-called ‘stipules’ are thin, non- vascularised and scarious, whilst in Marattiales they are vascularised and relatively massive. (b) In Ophioglossales they do not store starch (determined from thin sections stained with iodine in potassium iodide solution). (c) The ‘stipules’ of Ophioglossales, as we understand them, are at least initially in their development a single sheath. This completely encloses the next aerial complex developed, to form a ‘bud’ (Bierhorst, 1971). Subsequent growth of the complex ruptures the sheath and this often gives it the appearance of two stipules (Bierhorst, 1971; cf. Sporne, 1975). Further work on live plants is needed, however, to clarify these observations for each of the three genera. (d) Adventitious buds and lenticels often occur on the stipules of Marattiales but never on the sheaths of Ophioglossales. Of these comparisons, (c) in particular suggests that the sheath of Ophioglossales is not homologous with the stipules of Marattiales, except in the functional sense of an organ providing protection during early frond development. Apomorphous features (not shown in Fig. 23) of extant marattialean stipules compared to the sheaths of Ophioglossales are (a) their clear occurrence in pairs; (b) their function (besides protection) for photosynthesis and for storage of starch; (c) their relatively massive substance; (d) possession (in some species) of lenticels; (e) their ability to produce adventitious buds; and (f) their vascularisation. Although radically different in form from ordinary pinnae, it is generally held that the stipules of extant Marattiales represent reduced pinnae. Their association with the basal pulvinus of the rachis, and the association generally of pinnae and pinnules with pulvini (characters 6, 26-28), provides some indirect evidence for this belief. There is also some indirect support from certain fossils referred to the form genus Aphlebia Presl (Scott, 1920; Kidston, 1924; Crookall, 1976). These range in form from some species rather similar to the stipules of extant Marattiales to others that are pinnate in construction and therefore more clearly represent reduced pinnae, such as A. spinosa Lesquereux, A. acanthoides Zeiller and A. crispa (Gutbier) Presl (see e.g. Crookall, 1976). Some species of Aphlebia are known to occur in field association with Scolecopteris, and also with Pecopteris Brongniart and Asterotheca Presl (Asterotheca and some species of Pecopteris are attributed to Psaroniaceae). However, the only forms known in actual attachment to CLADISTICS OF MARATTIALES 239 pecopterid fronds are securely attributed to other families, such as the remarkable aphlebiae which occur at the bases of the pinnae in the fronds of Senftenbergia plumosa (Artis) Stur, figured by Kidston, 1924, Pl. XCIII, figs 2-4. These (for S. plumosa) resemble Aphlebia spinosa and therefore this very deeply dissected form of aphlebia may be dismissed in connection with Marattiales. Aphlebia acanthoides Zeiller is considered by Crookall (1976) on evidence of field association to have been ‘almost certainly borne’ by fronds of Asterotheca daubreei Zeiller. It is essentially bipinnate and to that extent pinna-like, but the divisions are merely cut into lobes. Such a form may be considered intermediate between a pecopterid pinna and the form of stipule seen in extant Marattiales. Aphlebia crispa (Gutbier) Pres] (Gutbier, 1835: 13, pl. 1, fig. 11 and ? pl. 6, fig. 18; see also Crookall, 1976), although two to three times pinnate, has a broader axis and has lobes that are rather more flabelliform and decurrent than in A. acanthoides, the lobes having lacerate margins. Each primary lobe approaches (more closely than in A. acanthoides) the most deeply divided form of stipule found in extant Marattiales, though considerably more deeply incised in the fossils than in extant material. Aphlebia membranacea Lesquereux (see Crookall, 1976) has a broad main axis, is only once-pinnate and the divisions are distinctly flabelliform lobes, irregularly divided and with a fimbriate margin. Therefore in many respects it resembles a stipule of living Marattiales. Aphlebia flabellata Sternberg is still more flabelliform, without a clear ‘midrib’ (Hirmer, 1927) and also has a partly fimbriate margin. To this extent we consider A. flabellata, together with some other species such as A. linearis (Gutbier) Presl, may show the nearest approach to stipules of extant Marattiales. An Aphlebia species (cf. A. crispa) with lobes somewhat similar to those of some extant marattialean stipules is illustrated for comparison with them in Fig. 6. Although a sequence of forms of Aphlebia could be proposed by which pecopterid pinnae may have become progressively reduced, and although there is some evidence from field association that such forms were borne by psaroniacean fronds, there is no direct evidence for either. In contrast there is direct evidence that several different groups of ferns and fern-allies bore aphlebiae of varied form. Therefore any proposed evolutionary sequence based on detached forms alone must be viewed with extreme caution. Only two generalisations are at all clear from the fossil evidence: firstly, that aphlebiae generally functioned in the same way as the stipules of extant Marattiales to protect fronds and/or individual pinnae at the crozier stage and during unfurling of the frond. They were located at the sites where pulvini generally occur today in marattialean fronds (excluding mid-stipe pulvini). Secondly, it seems that possession of stipules and aphlebiae is a plesiomorphous feature of ferns and their allies. We speculate that they served to protect megaphyllous fronds at their most tender (and palatable) stage of ontogenetic development, at a stage in their evolutionary development before the plants had evolved the battery of chemical defences that today deters many animals from eating their young fronds. Such a supposition is testable to an extent from the comparative biochemistry of extant taxa. Marattiales and Ophioglossales at crozier stage would be expected to have fewer such chemical deterrents, or at any rate less sophisticated ones, than Filicales. More particularly we cannot conclude that Psaroniaceae definitely had stipules. The evidence that they had aphlebiae is merely based on general field association and is at best of structures that are only approximately stipular in form. Stipules (or aphlebiae) were lacking from the young Psaronius sporophytes described by Stidd & Phillips (1968), one of which had five leaf traces and some attached petiole bases. (In extant Marattiales, stipules do not occur on the first frond of the young sporophyte but are soon developed on later ones, the second in Christen- senia, the third in Marattia and Angiopteris, though sometimes not until the fourth in Danaea (Farmer, 1892; Campbell, 1908, 1911)). Older stems of Psaronius are difficult to evaluate, since only the apical region of the stem surface is at all directly comparable with that of extant Marattiales. So far as we are aware, these apical regions of Psaronius lack persistent stipules. There is equally no evidence of stipules in the more proximal regions, which is to be expected in view of their dense development of external root mantle. In conclusion, the overall evidence as to whether Psaroniaceae had stipules (or aphlebiae) is not clear. Until such organs may be found in attachment or very convincingly associated, we suggest Psaronius lacked them. Even if it had them they were clearly not persistent. 240 C. R. HILL & J. M. CAMUS 6. Pulvini (Fig. 2). These are regions of the frond axis that in live plants are more or less pale in colour and swollen but in herbarium specimens are usually dark and shrunken. They occur above the stipules at the base of the stipe, sometimes elsewhere on the stipe but at some distance proximal to the lamina and at the junctions of pinnae or pinnules with the frond axes. Unlike other parts of the stipe and frond axes they have a hypodermal layer of collenchyma rather than sclerenchyma, as illustrated by Chang (1975), and a ring of mucilage canals may occur between the band of collenchyma and the epidermis (Sachs, 1882; Gwynne-Vaughan, 1905; Brebner, 1902; Chang, 1975). Takhtadzhyan (1956) and Chang (1975) consider the pulvini function to orientate the frond lamina in relation to sunlight. According to Chang, Hayata’s (1928) discussion of Archangiop- teris ‘pointed out the fleshy nodose swellings on the middle of the stipes as resulting from ecological adaption, helping the plant to drop its leaves at this point under dry conditions’. Holttum (1978) comments for Angiopteris: ‘In dry periods of weather. . . the loss of water from cells in the pulvini causes the whole frond to droop, and also each separate pinna’. The pulvini can also act as points of detachment of senescent pinnules as shown by Croft’s field observations (pers. comm. 1983) and our own observations of Marattiales grown at K. Holttum (1978) writes that the size of pulvini is a distinctive specific character in the living plant. 7-10. Mucilage canals (Fig. 4A). The lysigenous origin and development of these in extant Marattiales is documented by West (1915). Mucilage cells are known to occur in some other distantly related filicopsids (especially Cyatheaceae), but canals are rarely formed. Since they are absent from Ophioglossales the occurrence of mucilage canals of the kind seen in Marattiales is considered to be apomorphous for the order. Mucilage canals have been recorded from some Psaronius stems (Morgan, 1959; Stidd, 1971; DiMichele & Phillips, 1977) but not others (Rothwell & Blickle, 1982). They have not been reported from the roots of Psaronius. Millay (1979: 39, pl. 15, fig. 108) however, shows such canals for pinnules of Scolecopteris and Stidd’s (1971) ‘lacunae’ indicate their occurrence in the axial components of the Scolecopteris frond. In our root sections of some species of Archangiop- teris not included in the cladogram, mucilage canals were not visible: A. tamdaoensis (Hayata s.n., Vietnam, 4 August 1917 (TI)), A. ‘variabilis’ Ching (ms name) (Wu Ligong 4196, SE. Yunnan (PE)), A. henryi var. somai (collector not known, Taiwan (PE)), A. subintegra Hayata (collector not known, Tonkin, Vietnam, September 1932 (P)) and A. ‘brevipetiolata’ Ching (ms name) (Feng Guo-Mei 5078, SE. Yunnan (PE)). Campbell (19145) did not report mucilage canals in the roots of young Macroglossum plants, though we have seen them in the roots of a mature plant. Mucilage canals running parallel with and below the veins under the sori were considered by Campbell (1914a, b) to be unique to Macroglossum. However, we have found them also in our sections of five other of the terminal taxa shown in Fig. 23 and in Angiopteris polytheca C. Chr. & Tard. (Poilane 2370, Laos (BM)). 11. Sclerenchyma in the stem cortex. Holle (1876) reported a layer of sclerenchyma in the outer cortex of the stem of a specimen of Danaea trifoliata Reichb. Later authors (Kuhn, 1890; Brebner, 1902; Campbell, 1911) did not see this feature in their studies of Danaea, and since Holle had not mentioned stipules on his specimen, it was assumed that it was not a Danaea. However, this layer of sclerenchyma definitely does occur in Danaea. We have found it both in creeping and erect rhizomes of specimens of D. elliptica Smith (C. Whitefoord 1215, Belize (BM)), D. fendleri L. Underw. (N. Y. Sandwith 1786, Tobago (BM)), and D. moritziana C. Presl (A. H. G. Alston 8539, Colombia (BM)). In these the layer is one to three cells thick. In a specimen of arborescent habit this layer is well-developed and may be up to eight or nine cells thick. (Persaud 76, Guyana (K, NY, UG): D. sp. A of this paper, illustrated in Figs 2,3, and 4A, B). Sclerenchyma in the stem cortex has not, however, been seen in other extant Marattiales. In Fig.6 A fossil Aphlebia, cf. A. crispa (Gutbier) Presl. Upper Carboniferous, locality unknown, V.2865. A, Showing the broad midrib with several pinnae. x 1. B, Close-up of bottom pinna at left in A to show lacerate margin. X 2-5. Compare with the stipule margins in Fig. 5. CLADISTICS OF MARATTIALES 241 DAL C. R. HILL & J. M. CAMUS Psaronius, the peripheral sclerenchyma of the cortex may occur either as longitudinal bands, or, as in extant Danaea, as a cylinder, and is absent from some specimens. Sclerenchymatous tissue is absent from the stems of Ophioglossales but occurs in those of many other pteridophytes, e.g. Polypodiaceae s.s., Cyatheaceae, Hymenophyllaceae. We consider these to be homoplasious occurrences with Marattiales. Unlike the Polypodiaceae and Hymenophyllaceae the development of such tissue in Marattiales may be correlated with an arborescent habit, at any rate ancestrally. 12-13. Root sclerenchyma. We have examined the roots of five Angiopteris species, five of Marattia, three of Archangiopteris and Macroglossum smithii: Angiopteris canaliculata Holttum (R. E. Holttum s.n., New Guinea [BM + pteridophyte spirit collection W. 393; K hort. 578-63.57807]), A. elliptica, A. evecta, A. boivinii Hieron. (M. J. Whitehead 56, Seychelles [BM pteridophyte spirit collection $.280; K hort. 301—73.04156]), Marattia laevis, M. douglassii (C. Presl) Baker (A. C. Jermy 8345, Hawaiian Islands [BM pteridophyte spirit collection S. 276; K hort. 272-70.02662]), M. novoguineensis Rosenstock (Sands, Pattison & Wood SPWX 1429, Papua New Guinea [BM pteridophyte spirit collection $.282; K hort. 330-75.03192] and J. R. Woodhams s.n., New Guinea [BM pteridophyte spirit collection $.275; K hort. 003-74.00163]), M. laxa Kunze (R. I. Beyer RIB.108, Mexico [BM pteridophyte spirit collection W.394; K hort. 175—78.01730]), M. rolandi-principis, Archangiopteris caudata Ching, A. hokouensis Ching, and A. tamdaoensis (Hayata s.n., Tamdao, Vietnam, 4 August 1917 [TI]). Not one of these shows sclerenchymatous tissue in either pith or cortex of the root. Protomarattia tonkinensis Hayata has hypodermal collenchyma which contains some sclerified cells. In contrast, all the species of Danaea examined had sclerenchymatous tissue in the root pith and also as a hypodermal layer in the root cortex, as shown in Table 2. These two zones of sclerified tissue are illustrated in Fig. 4A, in a root passing vertically through the stem cortex prior to its emergence, as in the apex of Psaronius. Unlike the occurrence of such tissue in the stems, there is no obvious adaptive explanation for this occurrence of sclerenchyma in marattialean roots, excepting possibly that greater stress on roots (as prop-roots) of forms with arborescent stems, coupled with evolutionary conservatism, might be considered to indicate that Danaea was primitively arborescent. Root sclerenchyma has not been reported from Ophioglossales. By outgroup comparison we therefore consider it to be the derived character state. 15. Root hairs. Brebner (1896) reported multicellular root hairs in a young sporophyte of Danaea simplicifolia Rudge and comments that they had not been reported for Angiopteris or Marattia. Campbell (1911) interpreted them as septate hairs and recorded them for Danaea elliptica, D. jamaicensis L. Underw., and Christensenia. West (1917a) illustrates the same feature in a transverse section of the root of Danaea alata Smith. Since we have seen nuclei in some of these hairs (Fig. 4C, D) we accept Brebner’s interpretation of them as multicellular. They appear always to occur mixed with ordinary unicellular hairs and are easy to overlook. We have seen them in Angiopteris canaliculata (R. E. Holttum s.n., New Guinea [BM pteridophyte spirit collection W.393, K hort. 578-63.57807]), A. evecta, A. boivinii (M. J. Whitehead 56, Seychelles [BM pteridophyte spirit collection S. 280, K hort. 301-73.04156]), Macroglossum smithii, Marattia novoguineensis (Sands, Pattison & Wood SPWX 1429, Papua New Guinea [BM pteridophyte spirit collection S.282, K hort. 330-75.03192]) and J. R. Woodhams s.n., New Guinea [BM pteridophyte spirit collections S. 275, K hort. 003-74.00163]), M. laevis (A. C. Jermy 8345, Hawaiian Islands [BM pteridophyte spirit collection $.276, K hort. 272—70.02662]), M. laxa (R. I. Beyer RIB.108, Mexico [BM pteridophyte spirit collection W.394, K hort. 175-78.01730]), Christensenia aesculifolia, and Danaea sp. A. We therefore predict that they are a universal character of extant Marattiales. Since root hairs in pteridophytes are typically unicellular, this character is also considered apomorphous (Bary, 1884; Bierhorst, 1971; Ogura, 1972). Root hairs have not been recorded from Psaronius, nor from Ophioglossales, which have mycorrhizal roots (Bierhorst, 1971). We have therefore adopted the polarity absence = plesiomorphous and presence = apomorphous. CLADISTICS OF MARATTIALES 243 17-19. Habit and symmetry of adult stem. West (1917b) and Chang (1975) interpreted Danaea alata and Archangiopteris henryi as having dorsiventral rhizomes, but we consider only Christensenia to be truly dorsiventral. Danaea, Archangiopteris and Protomarattia all show fronds emerging both from the lateral and ventral surfaces of the rhizome as well as from the dorsal surface, as opposed to solely from the dorsal surface in Christensenia. Amongst Ophioglossales, Helminthostachys has a dorsiventral rhizome. Our choice of polarity for this character is therefore based on ‘pteridophytes’ as a whole, for which the dorsiventral state is generally considered apomorphous. Observation of populations in the field is needed to clarify the extent of the distinction drawn between erect rootstocks vs creeping rhizomes, and which we have had to base largely on limited collections of dried specimens. Figures 2 and 3 illustrate the elongate erect stem of Danaea sp. A, which in some respects shows a remarkable resemblance to the fossil stem Psaronius. 20-23. Pinnation of mature fronds. The ‘mature’ frond is considered here to be the stage of development that bears sporangia. This is usually also the most mature stage in terms of degree of pinnation. Rare instances are known, however, where earlier stages of pinnation may also bear sporangia, for example simple-leaved herbarium specimens (BM) of Christensenia and once-pinnate fronds of Angiopteris lygodifolia Rosenstock (Asama, 1960). This causes prob- lems in describing the degree of pinnation of the mature developmental stage of A. annamensis C. Chr. & Tard., because it is known from only a few specimens which show equally both once-pinnate and twice-pinnate mature fronds. The developmental sequence of frond forms in extant Marattiales is summarised in Fig. 7. Although a binary treatment is used for pinnate vs palmate frond development, the palmate frond of Christensenia is probably the result of evolutionary condensation of a complex pinnate frond form. There is, however, no positive evidence for this from the vasculature of mature fronds. There is a comparable range of ‘frond’ dissection within the aerial complexes of Ophioglossales. Botrychium is pinnatified to 3-pinnate, Helminthostachys approximately pal- mate, whilst Ophioglossum is simple or dichotomously forked. We consider (by reference to ‘trimerophytes’) that the highly dissected frond form of some species of Botrychium is the most generalised state in Ophioglossales, and therefore also in Marattiales. 24. Transverse ridges on stipe of mature fronds. An apparent homologue of this feature occurs in Marattia brooksii Copel. Neither it nor Angiopteris annamensis have been examined anatomi- cally and the ridges may merely represent some kind of pathological growth response. This character does not occur in Ophioglossales. By comparison within Marattiales its occurrence is considered to be the derived state and its absence as generalised. 25. Soft spines on the veins of the pinnules. These occur in Marattia laevis, mainly on the upper surface, but in some specimens also on the lower. Measurements of 35 spines from several herbarium specimens gave their length as (0:22—)0-56(-1-3) mm, and their width just above the decurrent base as (0-04—)0-12(-0:2) mm. No homologous feature has been reported from fossil foliage, but appendages of similar size and shape have been described from Psaronius (termed ‘scales’) by Stidd & Phillips (1968). This character does not occur in Ophioglossales. By comparison within Marattiales its occurrence is considered to be the derived state and its absence as generalised. 26-28. Pulvini on the frond stipe (Fig. 2). Hayata (1928) transferred Archangiopteris somai Hayata, A. subintegra and A. tamdaoensis to his now obsolete genus Protangiopteris partly because he considered a pulvinus was lacking from the mid-region of their stipes. However, he had earlier (1919) recorded a pulvinus in this position for the type specimen of A. tamdaoensis (TI). Archangiopteris henryi is figured and described by Christ & Giesenhagen (1899) as lacking this pulvinus in the mid-region of the stipe, but Hayata did not transfer this species to Protangiopteris. In fact all specimens of Archangiopteris and Protomarattia do show a pulvinus C. R. HILL & J. M. CAMUS 244 VILLVYUVAN a}euurd-g <———— "soyeI}] BIR] JULX9 JO SULIOJ pUOI Jo dduaNbes [ejusuIdojoaop Jo WeIseIG LZ ‘Bly “fd Sid¥31dOlIONVHOYV ‘fd vavnvad SIYALdOISNV VILLVYVAN f “td SIYSLdOIDNVHOYV SIHALdOIDNV ‘dd yavynva 4 “dd SIYSLdOIDNWHOUV WNSSOTDOYOWN VILLVYVA ‘dd vavNVad ae VINSSNALSIYHOD (VITOSIOITdWIS VAVNVQ }d90x9) ei9uab |e 4? eiaueb |e CLADISTICS OF MARATTIALES 245 in the mid-region of the stipe of fertile fronds. It is very clear in the type specimen of A. henryi (P), and Gwynne-Vaughan (1905) reported it for the material of A. henryi collected by Henry and preserved in spirit. (Gwynne-Vaughan’s preparations are in the Department of Botany, University of Glasgow.) Ching (19585) and Chang (1975) also consider a pulvinus in the mid-region of the stipe to occur in all individuals of Archangiopteris. Chang (1975) suggested that the occurrence of this character in all Archangiopteris fronds but only in the juvenile, once-pinnate fronds of Angiopteris (confirmed by Hayata’s and our own observations) indicates neotenous descent of Archangiopteris from Angiopteris. Angiopteris annamensis is the only species of Angiopteris normally having fertile fronds that may be either once or twice pinnate. It might provide such a link between these two genera. With so few species of Marattiales in cultivation there is little information available on this character for juvenile fronds. However, plants produced from adventitious buds on the stipules apparently pass through the same ontogenetic stages as plants raised from spores and therefore this evidence has been used provisionally instead. Of the live Marattiales observed in this way at K, none of the Marattia, Christensenia, or Macroglossum plants produced juvenile fronds with a pulvinus. However, all the Angiopteris juvenile fronds were found to have a pulvinus in the middle region of the stipe. Fig. 7 illustrates the developmental sequence of frond forms in extant Marattiales, with the positions of the pulvini indicated. Vareschi (1969) comments that the number of pulvini developed on the stipe is very variable in any one species of Danaea. However this does not seem to hold for all species of the genus. Vareschi possibly confused pulvini on the stipe with pulvini left on the rachis after detachment of pinnae and it is also possible that he treated several species as one. In this study the pulvinus on the stipe of D. simplicifolia is interpreted as the pulvinus basal to the lamina rather than the one in the middle region of the stipe. Pulvini do not occur in Ophioglossales and Psaroniaceae. By outgroup comparison their occurrence in extant Marattiales is therefore considered the derived character state. 29. Microscopic projections from exterior of walls of spongy mesophyll cells. These extraordin- ary intercellular rods illustrated in Figs 8 and 9 were first reported by De Vriese in De Vriese & Harting (1853) who saw them through the stomatal openings of Kaulfussia assamica De Vriese (now synonymised with Christensenia aesculifolia). Luerssen (18756) found them in the petioles, stems and roots of Angiopteris evecta and of Marattia cicutaefolia Kaulf. but was uncertain whether the structures seen in Danaea elliptica were truly such rods or fungal mycelia. He thought their composition was ‘cuticular cellulose’ and also reported their occurrence in members of Cyatheaceae, Polypodiaceae s.1., Osmundaceae and Ophioglossaceae. Bary (1884) considered them to be local thickenings of the cell wall membranes, whilst Gardiner (1885) thought that in ‘Aspidium filix-mas, Blechnum brasiliense and other ferns’ they were composed mainly of mucilage. Schenk (1886) concludes that they are neither pure cellulose nor wax and comments that their biological significance is completely enigmatic. Campbell (1911) claims they are ‘mainly composed of calcium pectinate’. Dr J. M. Pettitt has examined these structures cytochemically and with TEM. He found that alcian blue at pH 2:5, dialysed iron and the periodic acid — Schiff (PAS) method were all positive (the first two strongly so, PAS faintly positive), indicating the presence of polysaccharides. Staining with coomassie blue and oil red O were negative, suggesting that the projections do not contain protein or lipid. His electron micrograph (Fig. 9) shows that these structures are composed of microfibrils orientated the same way as those of the mesophyll cell wall, but that they seem to have been formed by successive layers of material laid down after the formation of the cell wall. We have also seen these projections in Helminthostachys (Ophioglossales) but not in other members of this group. We therefore regard their occurrence as a derived character state within both Ophioglossales and Marattiales. 30. Guard cells and lenticels. Campbell (1911) writes ‘The stomata of Kaulfussia (= Christen- senia) have long been known for their great size and the fact that they form permanently open C. R. HILL & J. M. CAMUS LY Fig. 8 Christensenia aesculifolia (Blume) Maxon, Griffith s.n., Assam, (BM): projections from exterior of walls of mesophyll cells (character 29). SEM photographs, Hitachi S 800, A x 1500, B x 4000, C x 10000, D, x 50000. CLADISTICS OF MARATTIALES 247 ao: call Fig. 9 Macroglossum smithii (Racib.) Campbell, A. C. Jermy 14223, Sarawak (Kew hort. ref. 279- 78.03064): projections from exterior of walls of mesophyll cells (character 29). TEM (Hitachi HS 9). x 40000. Photograph by J. M. Pettitt. pores, a peculiarity already developed in the cotyledon’. They are the largest of any vascular plant. Although De Vriese in De Vriese & Harting (1853) does not actually comment that Christensenia stomata are permanently open, his observations indicate that he was the first to observe this (see discussion of character 29). SEM photographs of herbarium material show open stomata in Christensenia and closed ones in other extant genera. Scott & Holden (1933) illustrate the stomata of Scolecopteris oliveri Scott with preserved guard cells. Holttum (1978) comments incisively on the different pattern of lenticel distribution in Angiopteris compared to members of the Filicales. In this genus, and also in large plants of Marattia and Danaea, l\enticels are found scattered all over the stipe and rachis (Fig. 5). In contrast in large plants of, for example, Cyatheaceae, Dicksoniaceae, Dennstaedtiaceae, and Thelypteridaceae, the aerating tissue occurs in two lateral bands on the frond axes. We have found neither stomata nor lenticels in the epidermis of the stipe of Christensenia. 31-33. Subsidiary cells of the stomata. (Fig. 10B, D, F). Although Van Cotthem (1970) describes the stomata of Ophioglossaceae and a few genera of Polypodiaceae as cyclocytic, his illustrations appear to us to show anomocytic to only weakly cyclocytic stomata. Strongly cycloctyic stomata appear to be unique to Marattiales amongst extant filicopsids. The ontogeny of the guard cells and subsidiary cells has been studied in only four species of 248 C. R. HILL & J. M. CAMUS = . 25 saa WO thong) ty he Fig. 10 Epidermal characters (characters 30-36). A, B, Christensenia aesculifolia (Blume) Maxon, Zollinger 1902, Java (BM). A, Upper epidermis of frond and B, lower epidermis. x 200. C, D, Danaea elliptica Smith, G. Aldridge 1834, Trinidad (BM). C, Upper epidermis of frond and D, lower epidermis. x 200. E, F, Marattia fraxinea Smith, H. Humbert 8417, Zaire (BM). E, Upper epidermis of frond and F, lower epidermis; idioblasts arrowed. x 200. C-F, Interference contrast, photographs by C. H. Shute. CLADISTICS OF MARATTIALES 249 Angiopteris (Kondo, 1962; Pant and Khare, 1969), with some apparent conflict in the develop- mental stages seen. Our preparations indicate that the difference in the arrangement of the mature subsidiary cells in Danaea compared to Marattia, Angiopteris, Macroglossum, Archan- giopteris, and Protomarattia is slight (character 32). 34. Stomatal dimensions (Table 1). Different cytological races are known to occur in Marat- tiales (Braithwaite, 1977; Walker, 1979), including Christensenia. Stomatal dimensions of two cytological voucher specimens (7. G. Walker T 13319, n = 40 [BM]; A. F. Braithwaite R.S.S. 4220, n = 80[K]) were compared with those of five other specimens of Christensenia. The results did not indicate that the level of ploidy is expressed in terms of stomatal dimensions. 35. Walls of epidermal cells. Van Cotthem (1970) surveyed the epidermal characters of Pteridium aquilinum (L.) Kuhn from Europe, the Americas, Africa, and Asia. His results showed that most characters of the epidermal cells vary greatly, for example cell shape and dimensions and sinuosity of their anticlinal walls. Such variation is also seen in Marattiales and Ophioglossales. However, the kind of sinuosity seen in the anticlinal walls of the ordinary epidermal cells of Christensenia (Fig. 10) is consistently different from that shown by all the other extant genera of these two groups and is therefore regarded as an apomorphous character state. Details of the walls of the epidermal cells of Scolecopteris oliveri are not obtainable from the slides of Scott & Holden (1933). 36. Idioblasts. These are approximately isodiametric cells with more or less granular contents. They frequently occur in pairs amongst the commoner, larger, and more oblong ordinary cells of the lower epidermis of most extant species of Marattiales (Fig. 10F) and also occur in the stem parenchyma. Christ & Giesenhagen (1899) termed them ‘kieselzelle’ and recorded their presence in Archangiopteris and Angiopteris but absence from Danaea. This term has been translated as ‘silica-cell’ and is so used by Pant & Khare (1969), Van Cotthem (1970), and Ogura (1972). There is, however, no evidence to suggest the cells contain silica. Their contents are dissolved in NaOCl solution as used in this study and by Van Cotthem (see his figs 13 & 18, where the characteristic cells are shown without contents) and silica is not soluble in aqueous NaOCl. Translation as ‘stone-cell’ is equally inadvisable as these cells bear no resemblance to sclereids. The contents of these cells have also been variously considered to consist of gypsum, calcium oxalate and ‘Kalilauge’ (potassium hydroxide) (Hansen, 1888; Rothert, 1887; Christ & Giesenhagen, 1899). Idioblasts are not known to occur in Ophioglossales. 37-39. Lamina margin and ‘fibre’ bundles. The Marattiales used for the cladogram all have toothed or lobed lamina margins. In some species, however, the division of the margin is barely Table 1 Summary of stomatal dimensions of extant genera of Marattiales. Angiopteris Macroglossum Archangiopteris Marattia Christensenia Danaea Length of stomata um (stoma + guard cells) Van Cotthem (1970: 97) 35-65 125-145 65-80 Pant & Khare (1969) (Angiopteris evecta) 47-73 this study 36-76 65-270 56-92 Breadth of stomata um (stoma + guard cells) Van Cotthem (1970: 97) 20-35 80-110 20-35 Pant & Khare (1969) (Angiopteris evecta) 13-17 this study 28-44 60-213 28-44 250 C. R. HILL & J. M. CAMUS noticeable below the apex of the pinna or pinnule and it is this to which the character states in Table 2 refer. The teeth or lobes each have a lateral vein running to the point, and (except in Christensenia) the sori occur over these veins in the fertile fronds. Millay (1976, 1979) reconstructs species of Scolecopteris a) with apparently fibrous lobes between or corresponding in position to the synangia (e.g. S. saharensis Millay, S. calicifolia Millay, S. latifolia Graham), and b) with more lobes than sori (e.g. S. mamayi Millay). In the fibrous forms, the lobes appear to be composed of ‘very elongate thick-walled cells of the pinnule hypodermis’. In these the lobing is not easy to compare with that of extant Marattiales but their fibrous construction may have some interest in relation to development of fibre bundles. A toothed lamina margin (character 37) is considered plesiomorphous because it is associated with the least reduced aerial complexes of Ophioglossales (though seen only in certain species of Botrychium). The fertile structures of Ophioglossales are such that character 38 (fertile pinnae or pinnules: barely recurved — recurved fully to protect sori) cannot be applied to them. The fully recurved state is considered derived based on comparison within Marattiales. Macroglossum, most species of Angiopteris and a few of Archangiopteris all have ‘fibre’ bundles (character 39). (The report of their occurrence in Marattia squamosa is erroneous, see Alderwerelt, 1911). These bundles, about five cells wide, are composed of elongate, sometimes thick-walled cells and run parallel to and alternate with the veins of the lamina. Mettenius (1856) accurately described their composition and criticised the still prevalent use of ‘nervi (or venuli) recurrentes’ (recurrent nerves) since they do not show any connection to the lateral veins. When their occurrence is not obvious as a dark line, visible from the lower surface of pinnules, a longitudinal cross-section of the pinnule lamina shows whether they are present or absent. Holttum (1955) writes that the structure of these ‘recurrent veins. . . is the same as that of the edge of the leaflet, and apparently they indicate that once Angiopteris had fronds more finely divided, but the finer divisions have joined together, the edge-character persisting at the junctions’. Holttum’s statement implies that margins of the pecopterid pinnules of Scolecopteris show an identical structure to fibre bundles, possibly by reference to the tissue of enlarged cells at the pinnule margin referred to as ‘hydathodes’ by Scott & Holden (1933). If so, there is in fact no close similarity to fibre bundles and Holttum’s view is difficult to understand. We have seen a few of these fibre bundles extending from sinuses of pinnule lobes in a partially tripinnate specimen of Marattia purpurascens De Vriese (Rawson 1998, Ascension Island [BM]]). The cells of these (as in other extant species) are not, however, identical with any of the cells of the pinnule margin. Hill, Wagner & El-Khayal (1985) have described Qasimia, a dimorphic Marattia-like fern with mainly taeniopteroid pinnules, from the Permian of Saudi Arabia. It may possibly explain the origin of fibre bundles more accurately. The vegetative fronds of Qasimia have more than twice the concentration of lateral veins seen in the fertile ones (the veins being correspondingly more densely forked in the vegetative pinnules). Interestingly, only the fertile pinnules appear to show fibre bundles between the veins. This suggests that the apparent fibre bundles of Qasimia represent merely vestigial ordinary vein branches that were surplus to the number needed to service the synangia in fertile pinnules. Thus they would not directly represent evidence of fusion from a more divided ancestral frond. In Mesozoic Marattia, which may be regarded as directly descended from Qasimia, fibre bundles occur both in the vegetative and fertile pinnules, whilst extant Marattia scarcely shows them at all. Fibre bundles do not occur in Ophioglossales and are therefore interpreted as apomorphous for Marattiales. 40-43. Scales (Fig. 11). In both Christensenia and Danaea the cells in surface view of the scale are more or less isodiametric, but those of Christensenia are considerably deeper than those of Fig. 11 Frond scales: illustrating cell shape and scale margin (characters 40-43). A, Archangiopteris henryi var. somai (Hayata) Tagawa, C. M. Kuo 6700, Taiwan (BM). x 50. B, D, Christensenia aesculifolia (Blume) Maxon, A. C. Jermy 8316, Philippines (BM). x 75. C, Danaea elliptica Smith, A. C. Jermy 10864, Trinidad (BM). E, Danaea alata Smith, A. C. Jermy 10936, Trinidad (BM). x 50. F, Marattia fraxinea Smith, Newman & Whitmore 475, Malawi (BM). x 50. CLADISTICS OF MARATTIALES 251 252 C. R. HILL & J. M. CAMUS Danaea, appearing box-like under the microscope (character 41). In Marattia, Macroglossum, Angiopteris, Archangiopteris, and Protomarattia the scale cells are elongate and shallow, the cells appearing very thin under the microscope. The lobed margin of Christensenia scales is markedly different from the characteristically fimbriate margin of scales of the other extant genera. Scales of Scolecopteris and Psaronius are very varied. They include multiseriate ‘hairs’ (Scott & Holden, 1933; Millay, 1979) which are here termed scales. Scott & Holden also described larger scales which are very thin and expanded, with a microscopically toothed margin much like that of scales of extant Marattiales (Fig. 11F). Stidd (1971) describes more substantial scales with dominantly multicellular transverse sections but which appear also to be expanded, at any rate towards their apices, and there become only one cell-layer thick. The massive, more or less conical appendages on a young sporophyte of Psaronius (Stidd & Phillips, 1968) are better referred to as spines. In extant Marattiales the larger scales often have a podium at the base and (particularly in Christensenia) are multicellular in transverse section for some way above this. Millay’s (1979: pl. 17, fig. 119) ‘scales’ seen on the pinna axis of Scolecopteris mamayi look remarkably like the podia of Christensenia. Scales are not known in Ophioglossales. Presumed transition from simple to complex has been used to determine the plesiomorphous vs apomorphous states of characters 40-43. 44. Uniseriate multicellular hairs. Scott & Holden (1933) figure these on pinnules of Scolecopter- is oliveri, and their occurrence in Scolecopteris is also noted by Millay (1979) and by Stubblefield (1984). Amongst extant Marattiales they occur on the abaxial surface of the lamina of Christensenia and Archangiopteris hokouensis. Hairs are characteristically lacking from the ‘aerial complexes’ of Ophioglossales. Their occurrence is taken to be the derived character state mainly on the basis of absence (plesiomorphous) — occurrence (apomorphous). 45,46. Veins of lamina. The usual venation type seen in the least reduced species of Ophioglos- sales is open dichotomous with reticulate venation in Ophioglossum which is considered reduced. Therefore reticulate and occasionally anastomosing veins are considered derived character states. 47. Sterile and fertile fronds. Bierhorst uses the non-commital term ‘aerial complex’ for the ‘fronds’ of Ophioglossales. If interpreted as true fronds (see Bierhorst, 1971, for discussion) their pinnae are strongly dimorphic, though the least so in species of Botrychium such as B. virginianum, considered by us the most plesiomorphous members of the order. Comparison is difficult with Marattiales even on this basis, since it is generally whole fronds of Marattiales rather than their pinnae that are dimorphic. We have chosen (for simplicity) to polarise the states of this character solely with reference to extant Marattiales, thus regarding dimorphism as relatively apomorphous for the order. Note on characters 48-68. For many of these characters Ophioglossales provide no useful guidance for determining the polarity of character states. They do not in general have sori (the nearest analogue to Marattiales being the ‘synangium’ of Ophioglossum) and in all cases each sporangium has its own vascular supply, even in Ophioglossum (Bierhorst, 1971; Sporne, 1975). As aresult, this group has nothing that could be homologised with the placenta of Marattiales or Filicales. The polarity of the states of characters 48, 50-56, and 62-67 has therefore been determined by comparison amongst Marattiales. Figure 12 illustrates the terms used for discussion of characters 48-52 and 57-66. 48-49. Placental hairs and synangial scales. Presl (1845) segregated genera within Marattia partly on the presence or absence of an ‘indusium’ — a term applied wrongly to Marattiales and CLADISTICS OF MARATTIALES 253 ie gE lamina uniseriate sm ss multicellular branching x Y hair placenta 4 pnt of midrib synangial dorsal wall S valve or ventral wall mS sporangium Fig.12 A, Plan of undersurface of an idealised fertile pinnule of Marattiales; sm, synangium; ss, sorus of separate sporangia; lv, lateral vein. The lines X—Y indicate the plane of the vertical section shown in B. B, Vertical section through a generalised marattialean sorus or synangium, illustrating terms used for discussion of characters 48-52 and 57-66, in particular ‘dorsal’ vs ‘ventral’ wall and ‘placenta’. referring merely to the hairs attached to the placenta. Mettenius (1856) was the first to criticise this misuse of the term. Ching (1958a, b) referred to the hairs as paraphyses or paraphysate hairs, as did De Vol (1973) and Holttum (1978). Mettenius (1856) pointed out that Presl’s concept of these hairs as always present or absent in a species was incorrect. More recently, Bienfait (1968) concluded that as well as their presence or absence depending on the age of the specimen examined, it also depended on the ecological or geographical ‘forms’ or ‘races’ studied. We have recorded great variability in development of placental hairs in successive fronds of a specimen of Angiopteris evecta growing at K (Birdsey B, Pahang, K hort. 371—77.02808). Millay (1976) comments on Scolecopteris ‘the quantity of hairs present on any given specimen varied widely, and is thought to reflect environmental influences’. Chang (1975) and Holttum (1978) consider the degree of branching of the placental hairs may be a useful character at the specific level. In Danaea, scales (rather than hairs) occur on the walls of the lamina troughs that envelop the synangia, but they are not found on the placenta. Uniquely amongst extant Marattiales, we have found that Christensenia has scales on the dorsal wall of the synangium (Fig. 13A, B). These synangial scales are easily shed, and commonly only the slightly protruding region of attachment is seen. Millay (1976, 1979) found uniseriate multicellular hairs on the sporangial wall of eight species of Scolecopteris and multiseriate as well as uniseriate hairs on those of S. saharensis and S. mamayi. His illustration of S. mamayi (1979: pl. 17, fig. 116) suggests that the ‘multiseriate hairs’ are referable to scales. Fig. 13 A,B, Christensenia aesculifolia (Blume) Maxon, A. C. Jermy & J. M. Rankin 14907, Sabah (BM): scales on dorsal wall of synangium (character 49). The synangia were rehydrated in 10% commercial bleach (‘Domestos’), then critical point dried. SEM photographs. A x 20. B, Detail of scale from another synangium of same specimen as in A. X 150. C, D, Vertical sections of synangia of Danaea showing lamina between synangia ‘expanded’ to varying degrees (character 50). C, Danaea alata Smith, C. Wright 1066, Cuba (BM). x 60. D, Danaea nodosa (L.) Smith, A. F. Skutch 4180, Costa Rica (BM). x 60. Fig.14 Tracheids in placental tissue of synangium (character 51). A, B, D, Marattia laevis Smith, J. Miers s.n., 1840, sine loc. (BM). A, Vertical section through a synangium. x 40. B, D, Detail of tracheids from a successive serial section to A. B x 150. D, Close up of B. x 400. C, E, Christensenia aesculifolia (Blume) Maxon, Griffith 1696, Assam (BM): vertical section through a synangium. X 20. F, Tracheids at base of synangium (arrowed in C). Interference contrast. x 600. 254 C. R. HILL & J. M. CAMUS CLADISTICS OF MARATTIALES z Ay 255 & ey ‘ he a ae, .. 256 C. R. HILL & J. M. CAMUS 50. Expanded lamina tissue between sori. This interesting feature of Danaea is illustrated in Fig. 12C, D. It also occurs, but to a lesser extent, in Scolecopteris mamayi (Millay, 1979) and in extant Macroglossum and Angiopteris elliptica. In the fossil Millaya Mapes & Schabilion (1979b) the expansion is as strongly marked as in Danaea. We do not consider the expanded lamina tissue of Marattiales to be homologous with the intersporangial tissues of Ophioglossum (Bierhorst, 1971). 51. Tracheids in placental tissue. Millay (1979, 1982a) has observed fibres and tracheid-like elements with scalariform thickening in the pedicellate placenta of some species of Scolecopter- is. (He (19826) appears to indicate that S. conicaulis Millay has fibres rather than tracheids.) Millay regards all such elements in the placenta — whether fibres or tracheids — as primitive, based on interpretation of the synangium as a fertile telome truss (Zimmermann, 1952). To this extent the argument for polarity, whilst reasonable, rests as much on palaeontological evidence and preconceptions as on character comparison, though to the extent that Ophioglossales have vascularised axes bearing the sporangia, Millay’s view may be.considered to receive general support from character comparison. We have adopted the reverse polarity, based solely on character comparison within Marattiales. Figure 14A, B, D, shows short tracheids in the pedicellate placenta of extant Marattia laevis, comparable to those of Scolecopteris and (so far as we know) reported for the first time in this paper. In Christensenia there is a peculiar disc of such cells, at the base of the central column only, with no clear connection to the underlying vein (Fig. 14C, E) (originally figured, but not commented on, by Luerssen, 1875a). 53. Development of placenta. The pedicellate placenta (receptacle) of some extant species (Figs 14C, D; 17A, C) shows a remarkably close resemblance to the ‘pedicel’ of Scolecopteris (Millay, 1979, 1982a; Stubblefield, 1984). 55-56. Arrangement of sori. Mettenius (1856) writes that Kaulfussia (Christensenia) bears synangia in an approximately biseriate manner (character 56, illustrated in Fig. 15B). The developmental sequence of frond form (Fig. 7) has been separated by some authors into two species of Christensenia, C. cummingiana Christ and C. aesculifolia s.s. The distribution of synangia in Christensenia may be linked with apparent ‘concertina-reduction’ of the frond from fundamentally pinnate to approximately palmate. In the other extant genera, the sori typically occur in two rows parallel to the midrib (Fig. 15A). 57. Position of sori relative to margin. A phyletic slide in the position of sori on the lamina was proposed by Bower (1926), from marginal to superficial, but Wagner (1973) does not include soral position in his table of ‘Directionality of character states’. Danaea and some species of Archangiopteris have long sori that cover almost the full width of the lamina. They seem to differ. In Danaea the inner ends of the sori form a uniform line near the midrib and the sori extend for varying lengths outward towards the pinna margin. In Archangiopteris the converse arrangement occurs: the outer ends of the sori form a uniform line near the margin and the sori extend for varying lengths inward, towards the midrib (Fig. 16). However, this is a tendency only, which seems slightly stronger in Archangiopteris. 58. Symmetry of sorus in plan view. Whilst typically round in Christensenia and elongate in the other extant genera, abnormal radial synangia or sori occur occasionally in Danaea, Marattia, and Angiopteris and somewhat elongate ones in Christensenia. They mutually counteract any taxonomic value that might be placed on them. 59. Packing together of spore-bearing loculi of synangia (or sporangia of sori). The close packing of sporangia in the sorus (or of the synangial loculi) is unique to Marattiales amongst the ferns (see p. 272 for discussion). CLADISTICS OF MARATTIALES 257 ra alli “« S 8 FY x “ a. Lrchremar Tobeinete? AmdAn Be Rife riven fom. Mav tems 15212 Devrwere tr FLORA OF THE PHILIPPINES Ab Henne wlitelte (My) bexen Prarie mani Mount Sarawag Samar 1a ge Shxdetensen sgt Ee Fig. 15 Arrangement of sori (character 54). A, Protomarattia tonkinensis Hayata, Hayata s.n., 30 July 1917, Vietnam (TI): sori in two rows which lie parallel to midrib and margin. B, Christensenia aesculifolia (Blume) Maxon, G. E. Edario 15112, Philippines (BM): sori mainly in two rows, the rows parallel to the main lateral veins rather than midrib and margin. 60-62. Cell differentiation in sporangial or synangial walls. Stomata are not seen in the sporangial or synangial walls of any extant Marattiales or Ophioglossales, contrary to the frequently quoted statement of Eames (1936) e.g. by Sporne (1975). Nor have they been seen with certainty in these structures in Scolecopteris (Scott, 1932; Millay, 1979, 1982a, b). Millay has studied the reproductive structures of Scolecopteris species in detail, based on peel sections of permineralised material. For some characters the range of variation he describes 258 C. R. HILL & J. M. CAMUS Fig. 16 Arrangement of sori. A, Danaea nodosa (L.) Smith, A. Hombersley 240, Trinidad (BM). x 1. B, Archangiopteris henryi Christ & Giesenh., Henry 11544A, Yunnan (BM). Xx 0-5. In Danaea the inner ends of the sori form a uniform line near the midrib and the sori extend for varying lengths outwards towards the pinna margin. In Archangiopteris the converse arrangement occurs: the outer ends of the sori form a uniform line near the margin and the sori extend for varying lengths inwards towards the midrib. occurs also to some extent in extant Marattiales. For example, sporangial/synangial apices may be long or short, blunt or acute, though the range is smaller in the extant species. The ratio of the height of the placenta to the height of the sporangium/synangium may be low or high, which is very similar to that of the extant taxa. Some species of Scolecopteris are similar to extant species of Marattiales in the number of layers of cells in the dorsal wall of the sporangium or synangium, but there is greater variation in the detailed structure of this dorsal wall than in extant Marattiales. For example, in S. parvifolia (Mamay) Millay and S. majopsis Millay there is a middle layer of cells with thickened walls. However, a layer of thickened cells below the outermost layer of cells may also occur in the dorsal wall of Marattia costulisora Alston, M. werneri and M. rolandi-principis, as indicated by sections made from rehydrated herbarium specimens (Fig. 17A, B), but the quality of the rehydration is far from perfect and this requires checking from fresh material. The extant species examined for this study have some characters not described by Millay (1979, 1982a, b) for all Scolecopteris. These may well prove to be very useful at the subgeneric level. For example, in Angiopteris the outermost cells of the dorsal wall may be elongate, similar to those of the ventral walls, or almost isodiametric (Fig. 18B, D). Where isodiametric, the outer periclinal walls may bulge out conspicuously as in other genera. In Marattia the shape of the surface cells of the inner-facing walls of the synangial valves can be strikingly different from one CLADISTICS OF MARATTIALES Bw > ee enna : *. Fo ee Phe pen Fig. 17 A, Marattia costulisora Alston, Clemens 5157, New Guinea (BM): vertical section through a synangium, obliquely so of right hand valve. x 40. B, Detail of A showing cells with apparently thickened walls in dorsal wall of synangium (tc). x 200. C, Archangiopteris henryi Christ & Giesenh., Henry 11544A, Yunnan (BM): vertical section through sorus. x 50. D, Protomarattia tonkinensis Hayata, Hayata s.n., 30 July 1917, Yunnan (BM): vertical section through sorus. x 50. species to another (Fig. 18A, C). Our sections of synangia of Marattia fraxinea Smith and M. werneri show scattered cells in the outermost layer of the dorsal wall, the contents of which resemble those of the idioblasts of the lower epidermis (character 36). In Archangiopteris and Angiopteris the arrangement of the apical thickened cells can differ markedly in different species. Our concept of the outermost cell layer in a generalised Angiopteris sporangium, based on comparison amongst extant Marattiales and Ophioglossales (see p. 285), is one with a rounded apex with thick-walled, more or less isodiametric cells; outermost cells of the upper half of the dorsal wall more or less isodiametric with thin periclinal walls which bulge outwards; cells of the 260 C. R. HILL & J. M. CAMUS Fig. 18 Variation in surface layer of cells of dorsal and ventral walls of synangia and sporangia. A, Marattia brassii Copel., L. J. Brass 12117, Irian Jaya, New Guinea (BM). x 35. B, Angiopteris laciniata Vriese, E. Smith 2208, Thailand (BM). x 100. C, Marattia alata Smith, D. Moore s.n., Jamaica, 1857 (BM). x 50. D, Angiopteris ankolana Vriese, Junghuhn 698, Sumatra (BM). < 100. SEM photographs. ventral wall elongate and parallel to the dehiscence slit. The characters of Scolecopteris described by Millay (1979, 1982a, b), other than to some extent for S. minor Hoskins emend. Millay, and some other species do not appear, however, to correspond to this. In Ophioglossales the surface cell layer of the sporangial wall is composed uniformly of more or less isodiametric cells. The cells of the ventral walls of the sporangia are aligned parallel with the dehiscence slit and there is no group of more or less isodiametric, thick-walled cells at the apices of the sporangia. The polarity of characters 60-62 was decided by comparison with this group, partly because Psaroniaceae shows an unhelpfully wide range of variation of these characters in this context. We note, however, that ‘trimerophytes’, in contrast to Ophioglos- sales, had elongate more or less pointed sporangia and the surface cells of the wall were elongate (Stewart, 1983). 63. Number of sporangia or locules per sorus or synangium. This is not a constant character, even allowing for those taxa in which the sori almost cover the lamina width. An extreme example is the individual of Angiopteris evecta mentioned under character 48 above, where an average of seven sporangia per sorus was found on one frond and sixteen on the successive frond. In Christensenia (and possibly other taxa) a bimodal score obtained for the number of locules in the synangium might be considered to reflect polyploidy, although we did not obtain such a score for the cytological voucher specimens referred to for character 34. Our observations show a marked difference in placenta height of different individuals assigned to Marattia fraxinea. Millay (1976) also points out a difference in height of synangia in different ‘species’ of Christensenia. CLADISTICS OF MARATTIALES 261 64-69. Partial or complete lateral fusion of the dorsal or ventral walls of the sporangia (and dehiscence). Fusion is interesting in relation to whether the free sporangiate or the synangiate state should be considered the generalised condition for Marattiales. Early work by various authors on the ontogenetic development of marattialean sporangia is summarised by Bower (1898), with more recent studies by Bienfait (1969) and Bierhorst (1971). Bienfait considered that Angiopteris shows a tendency towards being synangiate whilst Bierhorst concluded that the free sporangiate condition is the derived one. As both synangiate and free sporangiate forms are found in Scolecopteris and other relevant fossils of Carboniferous age, palaeontological evidence has not been used by the various authors to determine polarity. In this study we have initially chosen as a compromise to consider the partially fused state to be generalised by using outgroup comparison with Psaronius + Scolecopteris, but our overall results refute this choice for the extant genera (p. 289). Millay (1976) observed that sporangia of Angiopteris crassipes Wallich and A. faurieri Hieron. may be fused laterally at the base. This feature has also been seen by us in Angiopteris fokiensis Hieron., A wallichiana C. Presl (Fig. 19A, B), Archangiopteris cadierei (Cadiére C, Vietnam [P]), and A. caudata (specimens ex Herb. Kwangsi University, Southern Kwangsi [PE]). We consider these mostly to be abnormalities. Presumably because he did not see such fusion in Archangiopteris, Millay (1976) writes that its sporangia are truly free, comparable to those of Eoangiopteris Mamay (Mamay, 1950). He (1976, 1979, 1982a, b) interprets the sporangia of Scolecopteris as being fused laterally for part or all of their length prior to dehiscence, and considers that the sporangia would separate laterally before shedding their spores. If so, this is unlike the extant Marattiales, as is the oblique arrangement of cells associated with the dehiscence slit in at least some species of Scolecopteris. Figure 20 shows the cells of the outer layer of the dorsal and ventral walls of sporangia and synangia in some extant Marattiales, all characteristic species of each genus. The photographs are of herbarium material rehydrated with 10% ‘Domestos’ and stained with gentian violet or safranin differentiated by brief immersion in ethanol. The thick-walled cells (stained darker) are similar in Marattia, Angiopteris, Macroglossum, Archangiopteris, and Protomarattia. In Danaea the cells around the pore have only their tangential walls thickened in relation to the pore. No such differentiation of cells was seen in preparations of Christensenia. Another feature seen in sori of Macroglossum, Angiopteris, Archangiopteris, and Protomar- attia, apparently also not recorded before, is that the immature sporangia on opposite sides of the placenta are touching, with the ventral walls of the sporangia vertical. Thus they have the same attitude as the two valves of Marattia synangia. As they mature they spread apart, the dorsal sporangial walls becoming closer to the lamina before the sporangia finally open. Similarly, the synangia of Christensenia seem also to open out prior to dehiscence. It is possible that Scolecopteris showed this as well. 70-73. Spore characters. Spores of seventy species of extant Marattiales have been examined by Mettenius, 1856; Tchistiakoff, 1874; Jonkman, 1880; Selling, 1946; Harris, 1955; Erdtman, 1957; Ching, 1958a, b; Nayar, 1964; Nayar, Lata & Tiwara, 1964; Tardieu, 1966; Welman, 1970; Erdtman & Sorsa, 1971; Lugardon, 1971, 1972; Kremp & Kawasaki, 1972; Bobrov, 1973; M. D. Crane, 1973; Morucchio, 1973; Devi, 1977; Millay, 1976; Huang, 1981; Pant, Misra & Singh, 1982; Tryon & Tryon, 1982 and Pettitt pers. comm. Their surface morphology and structure are described in varying degrees of detail and with varying accuracy. The spores of some of the taxa studied here are illustrated in Figs 21 and 22. Mettenius (1856) was the first to comment on the occurrence both of long and spherical spores, with mono- and trilete marks respectively, within the synangia of any one plant of Marattia. This has been recorded by later authors for Christensenia, Angiopteris, and Archangi- opteris, as well as for Marattia. Millay (1979) who has noted this also in Scolecopteris , comments that it indicates genetic instability and ‘is by implication a primitive trait’. It has been noted also in Jurassic species by Van Konijnenburg-Van Cittert (1975a). The spores often, however, appear alete. M. D. Crane (1973) discusses the occurrence and possible formation of these. He suggests that they may not truly lack some sort of aperture. But very little is known at present of 262 C. R. HILL & J. M. CAMUS CLADISTICS OF MARATTIALES 263 ee. Fig. 19 Fusion of dorsal walls of sporangia (characters 66 and 68). A, B, Abnormal specimens of Angiopteris. A, Angiopteris fokiensis Hieron., J. Linsley Gressitt 589, Kwangtung (BM). x 90. B, Angiopteris wallichiana C. Presl, Herb. Lacaita 15866, Sikkim (BM). x 150. C, Protomarattia tonkinen- sis Hayata, Hayatas.n., 30 July 1917, Vietnam (TI). x 40. SEM photographs. Compare with the sections shown in Fig. 16. Fig. 20 Cell differentiation of surface layer of sporangial and synangial walls. A, Angiopteris evecta (G. Forster) Hoffm., Leeds University Expedition, s.n., Sri Lanka, (in cultivation at Leeds University Botanic Garden + BM). x 50. B, Marattia alata Smith, D. Moore s.n., Jamaica, 1857 (BM). x 40. C, Protomarattia tonkinensis Hayata, Hayata s.n., 30 July 1917, Vietnam (BM). x 40. D, Danaea nodosa (L.) Smith, A. Fendler 129, Trinidad (BM). Xx 50. E, Archangiopteris henryi var. somai (Hayata) Tagawa, Ogata 1, Taiwan (BM). x 50. i \ Zr ane 3 Se oh 2 d i ato dl i _ i — _ aad — rei — co . as) i H | i ll Ca CLADISTICS OF MARATTIALES 266 C. R. HILL & J. M. CAMUS Sorsa, 1971; Lugardon, 1971, 1972; Wagner, 1974; Verma & Khullar, 1979; A. F. Tryon, MS). Bierhorst (1971) adopts the opposite polarity. Millay (1979, 1982a, b) has shown that there are two major differences between the spores of Scolecopteris and those of extant Marattiales. One is the range of spore size (c. 13-88 um in Scolecopteris vs c. 20-40 wm in extant genera). The other is the layer of the spore wall that develops ornamentation. Millay (loc. cit.) and Millay & Taylor (1985) consider that in most species of Scolecopteris it is the perispore that is ornamented; in a few (S. majopsis, S. altissima (Mamay) Millay and S. alta (Watson) Millay) it is the exospore. In extant Marattiales it is the exospore that shows ornamentation, though the perispore in Marattia, Angiopteris, Macroglos- sum, Archangiopteris, and Protomarattia more or less moulds and therefore replicates the exospore. Such replication of the exospore ornamentation by the perispore is seen in some Scolecopteris species (Millay, 1979). In Danaea and Christensenia it seems that the perispore is only seen as a scant layer between the spines or ridges of the exospore (Tchistiakoff, 1874; Lugardon, 1971, 1972; M. D. Crane, 1973; Millay, 1976; Gomez & Walter, 1980; Pettitt pers. comm.). T.E.M. work on extant Marattiales and Ophioglossales (Lugardon, 1971, 1972) shows that their exospore is three-layered, with cavities in the middle layer. Preliminary study of ‘trimerophytes’ (Gensel, 1980) shows that the exospore is two-layered, the outer, sculptured layer separating readily from the inner one. The exospore ornamentation of ophioglossalean spores ranges from warty through spiny (as in Marattiales) to distinctly foveolate, a more complex condition than occurs in Marattiales though some species of Archangiopteris show an approach to this. Psaroniaceae shows a similar range of ornamentation to extant Marattiales. Our choice of outgroups for characters 71-73 therefore proved inadequate to achieve a hypothesis of polarity of the various character states of relevance. We justify our choice only weakly, on the basis of 1. intuitive hypothesis of a simple to complex transformation series (i.e. warts — short spines — long spines — branched spines > reticulate/foveolate) and 2. wider outgroup comparison with ‘trimerophytes’. However, the evidence from ‘trimerophytes’ (Gensel, 1980) suggests that short spines are plesiomorphous, in which case the warts of some marattialean spores would be better interpreted as apomorphous (character 71). Results and discussion Using the method outlined on p. 226, the information in Table 2 was used to construct the cladogram shown in Fig. 23, which is summarised in skeletal form in Figs 24 and 25J. The pages which follow discuss the cladogram within the constraints of taxonomic relationships and compare it with previous classifications. The cladogram proves to be more highly resolved than these previous classifications in terms of internested taxa and enables clear definitions to be formulated of Marattiales as a whole, of the Psaroniaceae and of extant Marattiales (Marat- tiaceae), each of which taxa is shown to be cladistically natural. Within the Marattiaceae, except for Christensenia and Danaea, the current generic circumscriptions are not cladistically natural: Marattia, Macroglossum, Angiopteris, and Archangiopteris are all ‘paraphyletic’. This reveals a need for further work on these genera. In the interim a provisional written classification is provided which incorporates some paraphyletic taxa. Comparison of the cladogram with previous classifications The scope of the term classification is generally limited to mean ‘systems of words’ (Wiley, 1981). Such classifications are here referred to as ‘written classifications’ or ‘classifications in writing’. In our opinion diagrams such as cladograms are also classifications (Eldredge & Cracraft, 1980; Hill & Crane, 1982: 277), and may be referred to as ‘diagrammatic classifica- tions’. An important property of diagrammatic classifications is the means they provide for compari- son which overcomes the limitations traditionally imposed on written classifications by the Linnaean hierarchy. To this extent, adherence to the Linnaean hierarchy can be viewed as a CLADISTICS OF MARATTIALES 267 weakness of classifications in writing (but see p. 278). Diagrammatic classifications also facilitate demonstration of the characters used to define taxa, which is not economically possible (nor desirable) with written classifications. The distinction between written and diagrammatic classfications is not, however, absolute. In this paper our results on Marattiales are presented in four versions of classification. There is the entirely diagrammatic cladogram in Figs 23 and 24. Figure 23 shows the synapomorphies and homoplasies (a cladogram sensu stricto), Fig. 24 is without them (a cladogram sensu lato). A classification entirely in writing is given on p. 278 and a classification intermediate between it and a cladogram (an annotated cladogram) is given in Fig. 25J. This figure shows the taxa (lettered U-Z) that most directly represent the sequence of internested taxa of the diagrammatic classification. In principle, these taxa could all be named (as for taxon U, Marattiales and V, Marattiaceae), and to the extent that they are lettered the classification may be regarded as being partly in written form. We do not, however, formally name the taxa (other than for taxa U, V, and X) since to do so would merely clutter the literature with excess words (Harris, 1964). The main importance of these taxa is their internested relations and not their names, though clearly some names and letters are needed for discussion. It is a matter of balance. Figure 25J may be compared with nine previous classifications of Marattiales in the literature, which are represented diagrammatically in Fig. 25A—I to facilitate comparison. (Taxon U of Fig. 25J is disregarded for this purpose since none of the previous classifications includes reference to Psaroniaceae). Comparison shows a) That the previous classifications are very varied, b) The classification favoured by more authors than any other, Fig. 25A, is the least resolved in terms of internested taxa. Thus, although the least controversial classification, it is also the least informative in these terms. It is in fact totally uninformative. c) The cladogram is considerably more resolved than any of the previous classifications. Excluding taxon U, Fig. 25J has five internested taxa (ranks), whilst the maximum in the other classifications is two ranks (Fig. 25B-I), with the minimum of zero internesting, i.e. one rank only, in Fig. 25A. d) The classifications nearest to the cladogram are those of M. D. Crane (Fig. 25H; 1973) and Devi (Fig. 251; 1981). The previous classifications vary mainly because they do not fully resolve taxon W of Fig. 25J separately from V, nor X from W. Marattia is the ‘joker’, resolved with difficulty amongst these schemes as well as in ours. It appears in more different positions relative to other taxa than any of the other genera. Grouping X remains essentially constant throughout except for Pichi Sermolli’s (1957, 1977) scheme (Fig. 25F), which reflects the erroneous view that Protomarattia resembles Marattia more closely than Archangiopteris. We do not wish to imply that when cladograms of Marattiales are produced by different authors they will prove to lack variety. Results will vary according to the scope and number both of terminal taxa and of characters utilized, and depending on the exact method and polarities chosen. Some of the taxa U—Z are based on few synapomorphies and may well prove sensitive to change. This is another reason for our reluctance to formally name some of these taxa. Nonetheless, we would be surprised if cladograms produced in the future (assuming they are based on a comparably wide range of characters) prove to vary as widely as the previous classifications in terms of the sequence of internesting of the taxa. Two criticisms should be levelled at our comparisons. Firstly, the authors of the previous classifications did not intend them to be presented diagrammatically in class inclusion terms as if they were cladograms. Secondly, and more importantly, since the previous classifications are tied to the Linnaean hierarchy at the ranks of order and family, it is unjustified to expect more than two internested taxa to have resulted from them. We would accept this criticism, and merely state that we know of no other useful grounds for comparison: to refrain from such comparison for these reasons would clearly lead nowhere, whereas to make the comparison leads somewhere. Indeed, regardless of these considerations there are two important conclu- sions. Firstly, the cladogram unequivocally achieves a considerably greater resolution of internested relations of the taxa than the previous, more traditional classifications. Secondly, this greater resolution is a result of freedom from the restricted internesting of ranks maintained by conventional usage of the Linnaean hierarchy. This greater resolution of class inclusion , relatively apomorphous aracter is not applicable. Table 2 Data matrix for the two-state characters of Marattiales listed in the text. Plesiomorphous state shown as — as +. Those marked —/+ show both plesiomorphous and apomorphous states in the same taxon. / indicates a ch Parentheses are used where data are more or less conjectural, blanks are left where prediction seemed ill-advisable; both symbols indicate where further observation is needed. Wd}WD ueA-sinquouluoy ue, myonj{q siuajdoisup jeAeyy-[q wz J9Use AA ‘][IH (QUSIOWII’]) avwus{{yos DIUIS”DO ojAR] NUOISp1Y ,P1YIOISPoY, SISUBUIYUO] DIJDADULOJ OL sisuanoyoy siaajdoisunyoy pvpnvd Sidajdo1suvyo4y pyouuidig siaajdoisunyoay 12491PV9 S14a]dO1SUDYIAY 10N1 Sldajdo1suvyoaudy 1d4uay S1dajdo1subyoy NYMNUS WNSSO]ZOLIDW vsonajsuou siajdoisupy voydyyja sidajdoisuy sisuawouud Ssiuajdoisupy pjdada Sldajdoisuy S1AaD] DIDDY DAOSYNISOD DINIDADIN 1dULIM DIYIDADIY sidiguiid-1puvjod D1IIDADIY DaUIxvAl DINIDADI “vy ‘ds vavung pIDIJD DaDUDGq pyofiojduas pavuvg psopou vavuvg voudija pavung pyofynasav piuasuajsi4yy $14ajd02aJ09§ + snluosvsSg sayessojso1ydg, Jojovieyg +? ) + + CC) ie) Ce). OG) GORGE) Gn) + + ( + aa i) CE ere Ci) oe) CP yets) OH) Ci) + + ( + es SCE)! AGE) OCP eCe), sth) o Gr) a UTS) + + Ch) eA) + + + + + Ca eG te Gor eo up ee Ga) ie a) en a CE GE Gry iC) + + + (+) CoCo CEC) + + + + + + + + + + + (+) + toa aa a oe + oa a oa = + + + + + + + + + + + + + + + + — — +) (+) + ie ITI —_, ay, Ny, ieee a — — _~ —_~— dora Dee Wii eee — — ON (mm BOOS a —~—~ — ec eae ta toe Vw — Tad ae a ee —_ a Pare iater a —_v ich Ane — Se i haa NA a oe arena a in Mee es ie eae aia fo je eS ae oe —_~ b++ 04+ ae a pe eee Ti++ i te ea eo + / CE ea CEy A) + / 17 18 YO ~O WY” Cry ey A) (/) +? / / / / / 20 =/+ —? / / =e + + (+) + + + - - = = + + + CEC) GE GEE Ca CE) CE) + ie eo (suk Say ema (\ ie CO Gl Ce Cay a) he) ob + — + ase) + + + + task: les SS LO ste lbeat —“"_ oo + + berrterti 1% b+) ltteey + b+ibi+t | Serb r eter ti b+ l++t+t+i¢i b+] i+tttt+i +i b+ i++tt+i4i P+1ir+ttit¢i b+) btttti + b+) btttti +i b+) i rrttti¢i b+ibtttti ti iti i itttit¢i b+ critter 4t b+ibtttti ti t++r~r ieee it b+ i++tti cil }+~~~+H+4+ 1/11 b+trttttill ber rreetrrt Cee b+iitttt+i~i b+ittttt+i~i+ b+) btttt+i~ b+) b+t+tt++1~ 1 ++tter errs Ter i+i+etit + eet Dh tacks 1+7400~n | + i tee +~ 1 ltt =) ie ee +~ litt Seed +~1 1+ +~ll+++t +~llit~ +~Il1l+~ +~1l+4i +~11lt~ +~I11l+~ b+i++ i ee b+ 1+4+4+1 bt+i++t+1 b+)1+1+~ econ UD b++4+4+ 11 b++++4+1 b+++4+ 11 b+++4+1 1 b+l14+4+1i Peon rd Tae SSSREERR C. R. HILL & J. M. CAMUS 270 I LULHIK * * * 4 * * * 1H * ru NNN Rr c za * 2g BA — T T T 6 ue be i rs Se 19 * * LZ a el * m@ x * x2 * <4 * er Z Te | J + * mo I I I I J Af ve * * _——— “ [pees eee ae 82 i =) E J [sss } x vy 5 I I I I LC I LT T I I se I ST: ED a a rT I a we e ae | = J E ] i esi | v2 r I I I : I I st I I LS ae I Do i I T I oa] €9 ee Ot —— a Cc = ae c 2 he es +0. EO (o> Ol SSS CSS SC OS eM St OS TE Zo Of NE eS 10 MO: 2. OC 20 (OG) FO, 200 ODP OD FD FD FD MD on O@ 2i2e 2 BSS ESS Oc z= O> 22> 22> NSF ES FS OF => mr aM ZO. KO CO) 20. OO (20° SO. 20 ZO “fo Mo. 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