FALKLAND ISLAND NEPATIGAE AND ANTHOCEROTAE: A TAXONOMIC AND PHYTOGEOGRAPHIC STUDY ' Thesis for the Degree of Ph. D. MICHIGAN STATE UNIVERSITY JOHN J. ENGEL 1 9 7 2 u‘ug‘a-as - Michigan State University This is to certify that the thesis entitled ILKLAND ISLAND HEPATICAE AND ANTHOCEROTAEt A TAXONOMIC AND PHYTOGEOGRAPHIC STUDY presented by JOHN J. ENGEL has been accepted towards fulfillment of the requirements for PH.D. demein BOTANY e PLANT PATHOLOGY ‘ /44 “1‘1" Major professor For H. A. lmshaug 2h-72 2:" outcome a? \ NUAB 8 SBNS' ‘ A 300K B‘NDERY INC. ? , LIBRARY amos as .£33 at". ABSTRACT FALKLAND ISLAND HEPATICAE AND ANTHOCEROTAE: A TAXONOMIC AND PHYTOGEOGRAPHIC STUDY By John J. Engel The Falkland Islands were visited in l968 and ca. 1000 collections of Hepaticae and Anthocerotae were made. These collec- tions, which represent the first made by an hepaticologist in the Falklands, as well as the collections of earlier visitors such as Sird. D. Hooker and Dr. Carl Skottsberg, were studied in detail and identified insofar as possible by comparison with pertinent type material. The results of this study are presented in the form of keys to all taxa, and a treatment of each of the l27 Species. These treatments include taxonomic and nomenclatural remarks where pertinent, statements of ecological tolerances. and requirements, phytogeographic remarks, lists of literature records and lists of material studied. Particular attention is given to the typification of Falkland Island and related taxa. Studies resulted in one new genus, Hygrobiellopsis (Schust.) Er‘99], and the following new species: Adelanthus tenuis, Andrew- Maustralis, A. planifolius, Chiloscyphus hookeri, Frullam‘a oseodolobulata and Metzgeria falklandica. Detailed plates of all new taxa have been prepared. In addition, the following nine new combinations were made: Cheilolejeunea savatieriana (Besch. 8. Mass.) 1 John J. Engel Engel, Clasmatocolea cookiana (Mass.) Engel, Clasmatocolea georgien- sismott.) Engel, Hygrobiellopsis isophyllum (Schust.) Engel, Meta- rygrobiella tubulata (Hook, f. & Tayl.) Engel, Microlepidozia cuculli- ir_‘2i_a_(Steph.) Engel, fl. _m_M_l_i_s_ (Steph.) Engel, fl. saddlensis (Besch. lilass.) Engel, and fl. setiformis (De Not.) Engel . The ecology of the Falkland Island Hepaticae and Anthocerotae is assessed by the arrangement of the taxa into the communities recog- nized by previous workers. The distribution of taxa within the Falkland Islands is dis- cussed, especially with reference to available rainfall data. An assessment of the phytogeographic affinities of the Falkland Island Hepaticae and Anthocerotae is made, with special reference to the characteristics and differentiation of south temperate and subant- artic distribution patterns. These patterns are historically based on the classification of austral regions which primarily utilize vegetational criteria. The relative merits of the several zonation schemes is discussed and sunmarized. FALKLAND ISLAND HEPATICAE AND ANTHOCEROTAE: A TAXONOMIC AND PHYTOGEOGRAPHIC STUDY By \ John J". 'FEngel A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Department of Botany and Plant Pathology 1972 w'F-l ACKNOWLEDGMENTS The entire project was carried out while I was a research assistant of Dr. Henry A. Imshaug. This support is acknowledged with thanks. I should like to extend special heartfelt thanks to Dr. Henry lmshaug for his interest, good humor, invaluable guidance and stimu- lating discussions throughout the project. Thanks also to Dr. Imshaug for providing me with the opportunity of collecting in the Falkland Islands. Ishould like to thank Mr. and Mrs. Les Gleadel of Port Stanley, Falkland Islands, in whose home I resided while in Port Stanley. Their hoSpitality and kindness will not soon be forgotten. Thanks also to the many other individuals of the Falklands who extended their hospi- tality. Iwould like to extend special thanks to my wife Karen, who has aided at many and various stages in preparation of the manuscript, and not least for her continued encouragement. I am indebted to Richard C. Harris for assistance with the Latin diagnoses. I am grateful to the following hepaticologists for identifying Falkland Island groups in which they specialize. The keys to these groups were also prepared by these specialists. These indi- viduals are: Dr. R. Grolle, Jena Germany (Cryptochila, Jamesoniella), with special thanks to Dr. Grolle who generously supplied unpublished i‘i information, Dr. Gabriela Hassel de Menendez, Buenos Ai res, Argentina (Aggraliggaflijg), Dr. H. Inoue, Tokyo, Japan (Plagiochila) and R. Jiri Vaia (Jungermannia). 1am indebted to Dr. H. Roivainen of Helsinki, Finland, who determinedmy Falkland Island moss collections. I have used his de- terminations in this treatment, especially in the ecology section. lwish to acknowledge the aid of the individuals and institu- tions listed below for the loan of specimens: Dr. Francesco Bianchini (IER), Dr. C. E. B. Bonner (G), Dr. A. Bresinsky (M), Dr. Suzanne Jovet-Ast (PC), Dr. I. Mackenzie Lamb (FH), Prof. Dr. J. Miege, (G), Dr. Elsa Nyholm (S-PA), Dr. Clark T. Rogerson (NY), R. Ross (BM), Dr. R. Santesson (UPS), Dr. R. M. Schuster (MASS) and Dr. Gary L. Smith (NY). .as TABLE OF CONTENTS Section Page I. THE FALKLAND ISLANDS ................. l A. Location ...... . .............. l B. Topographical Features .............. l C. Geological Features ............... 2 D. Soils ...................... 4 E. Climate ..................... 4 ll. COLLECTORS 0F HEPATICAE AND ANTHOCEROTAE ....... 7 III.ECOLDGY....................... 15 A. Introduction ................... 15 B. Distribution of Bryophytes in Plant Communities of the Falkland Islands ............ l7 1. Maritime Tussock Formation .......... l7 a. fgaflabellata Association ........ l7 2. Oceanic Heath Formation ........... l8 it -r—Si§§ad§iifib"iiiiial122e.-aaaoas: I I : : : ‘2? 3. Feldmark Formation .............. 27 4. Fen and Bog Formation ............ 31 a. Rostkovia Association .......... 3] b. Wssociation ........... 32 c. Juncus scheuzerioides Association . . . . 32 iv TABLE OF CONTENTS - continued Section Page 5. Bush Formation ................ 32 a. Chiliotrichum Association ........ 32 b. FleEe Association ............. 33 6. Littoral Vegetation ............. 33 7. Fresh-Hater Vegetation ............ 33 8. Sheltered High Altitude Cliff Vegetation. . . 34 9. Man Influenced Communities .......... 36 a. Pastureland ............... 36 b. Groves of Planted Trees ......... 36 c. Gorse Thickets .............. 37 IV. PHYTOGEOGRAPHY .................... 38 A. Austral Regions ................. 38 B. The Phytogeographic Categories .......... 43 1. Temperate .................. 45 a. American Temperate ............ 45 1) Endemic to Falkland Islands ..... 45 2) Fuegian-Falkland ........... 46 3) Magellanian-Falkland ......... 46 4) Valdivian-Falkland .......... 47 5) Valdivian + Magellanian-Falkland. . . 48 5) Andean-Falkland ........... 52 b- Amphipacific Temperate .......... 52 c. Mphiatlantic Temperate ......... 53 CL Pan-temperate .............. 54 2- Subantarctic ,,,,,,,,,,,,,,,,, 55 3- ADtarctic .................. 55 4' NidFSPread .................. 56 C. Distribution Within the Falklands ........ 56 v TABLE OF CONTENTS - continued Section Page v. SYSTEMATIC ACCOUNT .................. 61 A. Introduction ................... 6T l. Format .................... 6l 2. Nomenclature and Literature Citations . . . . 6l a. Author Citations ............. 6l b. Journal Citations ............ 6l c. Synonymy ................. 62 d. Typifications .............. 62 e. Herbarium Citations ........... 63 3. Ecology ................... 63 4. Distribution ................. 64 5. Literature Records .............. 64 6. Specimens Seen ................ 65 3~ Key to Classes and Orders of Hepaticae and Anthocerotae C. Order Jungermanniales: Key to the Genera of the Falkland Islands .............. 68 0- Order Metzgeriales: Key to the Genera of the Falkland Islands .............. 458 5- Order Marchantiales ............... 5T4 F- Order Anthocerotales ......... . . . . . . 519 V1. LITERATURE CITED ................... 523 “1° INDEX or TAXA .................... 543 “”- PHOTOGRAPHS ............... 561 Ix' "A” . 569 vi I Page 129 187 ; ‘ IN 189 i Wflanifoliu Engel ............ T96 5 WM subsp. hookeri Engel ....... 264 5' Whookeri subsp. hookeri Engel ....... 266 ‘ 7' Melanthus m Engel and Grolle ........... 425 5- MMEngel and Grolle ........... 427 ._ 9- lelania W Engel ............. 447 ‘A 10' :"llwamm Engel ............. 449 " _- _- - ria falklandica Kuw. late re ared b Hen“! Kuwa ara ..... (F3. . p .p. . . y ..... 506 —-—vr 3. 4 a“; . ti association at Kidney Island ..... '2‘.:‘iiroiriectng blades of Poa flabellata at Kidney Island: photo by H. Flmsfiaug ........... “maria association near Mt. Usborne caravan; PBS 5; R. C. Harris ............... Stream through Cortaderia association near Ht. Usborne caravan ................ . Hat depression in Cortaderia association in gap between French Peaks ................ Stonerun as seen from air over East Falkland ..... ' MW in stonerun ............. Dwarf shrub heath association near Mt. Adam ..... Stream in dwarf shrub heath association near Heddell Island settlement ............. ' @9111 tica component of dwarf shrub heath association in Waterfall Valley, Nestpoint Island . Dr: Iggmark on ridge of north slope of Mt. Fegen. ' Shzltered high altitude cliffs (arrow) of Mt. Adam Wt "ea; ““0 by R. c. Harris ......... firm of planted trees (mainly Po ulus, Nothofagus and conifers) at Hill Cove sett ement ....... Interior of planted re - t at Hi 1 Cove sett] :tgt‘ove (mainly Nothofagus) ooooooooooooo Page 562 562 562 562 564 564 564 564 566 566 566 566 568 LIST OF PHOTOGRAPHS - continued Photograph Pa ge l5. Extensive growth of archegoniphore-bearing plants of Marchantia berteroana at south shore of Cape Pembroke peninsula .............. 568 l6. Extensive growth of archegoniphore-bearing plants of Marchantia berteroana at south shore of Cape PemEroke peninsula .............. 568 ix 5‘ 4 .“ \ - N LIST or MAPS] Map Page l. Temnoma quadripartita (Hook.) Mitt ............ 570 2. Isotachis humectata (Hook. f. & Tayl.) Steph. Open chle indicates precise locality unknown ....... 570 . a 3. Lepicolea ochroleuca (Spreng.) Spruce. Plus Hondurus north to Mexico .................... 570 " eraw‘.) rite. 9"?".°T"f‘? refer? 5. Gackstroemia magellanica (Lam.) Trev ........... 572 6- Mepidozia bicuspidata (Mass.) 5. Arnell ...... 572 7~ WbleHODSis mphyllum (Schust.) Engel ....... 572 8' W162? Egilifgigcigy l ocZi TEyS::::ogen?r?1 T; . open . S72 9' anéfiflpietifieigehiiiflninown. 574 10' Mblephamstoma (Steph.) Fulf. ......... 574 “' Mm (Lehm. e Lindenb.) Fulf ....... 574 12' Mpseudozoopsis (Herz.) Fulf ........... 574 \ I We from; maps with the exception of numbers IS, 4T , 59 and SO were was adapteclmidr'gS made by the author. The south polar rojection detailed ma tour a Nafional Geographic Society map (1943). The non- here adapteii (if SOUthem South America and the world projection map wot e '0'“ the Goode base map series, University of Chicago. fTorn seveigisuog detaiied maps of southern South America were adapted l'ndly Sent t - . Naval Oceanographic Office charts. Map 59 was 0 me by Dr. Hugo ders. University of Uppsala. Maps 61 n were ada 6 Filled from a Di ma . Ma 3,,“ reproduced from Greenerfiggfite of Overseas Surveys p P L D 4 ‘5 v. \\ I.. ‘- \\ N. 4- .N LIST OF MAPS - continued hp Page 13. Anastrepia bifida (Steph.) Steph ............. 576 I4. Roivainenia jacquinotii (Mont.) Grolle. Plus South Georgia ..................... 576 l5. Cryptochila grandiflora (Lindenb. & Gott.) Grolle. After Grolle (l97l) .................. 576 It. Cryptochila paludosa (Steph.) Grolle. Plus Tristan da Cunha. Open cirfle indicates precise locality unknown. . . . .................... 578 ll. Jamesoniella colorata (Lehm.) Schiffn. After Grolle (T971)....... 578 l8. Herzogobryum erosum (Carring. & Pears.) Grolle ..... 580 ll. Herzogobryum teres (Carring. & Pears.) Grolle ...... 580 20. Herzogobryum vermiculare (Schiffn.) Grolle ....... 580 21° marldtmhyllum clandestinum (Mont. em Grolle) Grolle. 580 22- Marldophyllum densifolium (Hook.) Angstr. _e_)_c_ Mass. . 582 23' WIdOPhI/llum mtscheanum Grolle. Open circle indicates precise locality unknown .......... 582 24' Weancellata (Nees) Steph ........... 582 25' Baiéntlfim'ierinacea .................. 582 26' WMEngel subsp. hookeri. Plus 49° 02' ’ P- coAtantifolius - open circle ....... 584 27' masmat¢lfiftcookiana (Mass.) Engel . . . . . . . . . . 584 28' WINNIE (Hook. f. & Tayl.) Grolle . . . . 584 :: Wgeorgiensis (Gott.) Engel. . . ...... 586 WWW“. f. a Tayl.) Grolle. . . . . 586 31' Clasmatocolea humilis (Hook. f. & Ta 1. Grolle. Motor localities only. .y. ). ........ 535 Clasmat . Wéfipm (Gott.) Grolle. Plus 586 xi LIST OF MAPS - continued llap 33. 36. w SD J) .._a 42. w 46. 47. 48. 5). 52. 4 . Clasnatocolea vermicularis (Lehm.) Grolle. Plus Triathlon andTostfllica ................ Leptoscyphus abditus (Sull.) Dugas ........... . Leptoscyphus aequatus (Hook. f. & Tayl.) Mitt. Open circle indicates precise locality unknown ....... Leptoscyphus gpansus (Lehm.) Grolle. Open circle indicates precise locality in Mascarenes unknown . ' Q . Leptoscyphus patagonicus (Steph.) Grolle ........ . Lo hocolea austrigena (Hook. f. & Tayl.) G. L. & N. us ristan a unha. Open circle indicates precise locality unknown ................... . Lophocolea elata (Gott.) Steph .............. 40. Lghocolea leptantha (Hook. f. 8. Tayl.) G. L. & N. Opeanle indicates precise locality unknown . . mphocolea semiteres (Lehm.) Mitt. Open circle indicates precise locality unknown .......... Lophocolea sfivatica Mitt ................ Qphocolea textilis (Hook. f. It Tayl.) G. L. & N.. . . . . Eaghyglossa dissitifolia Herz. & Grolle ......... 45. B§c_hyglossa fissa (Mitt.) Herz. 8. Grolle ........ Pa_chyglossa spegazziniana (Mass.) Herz. & Grolle . . . Saccogynidium australe (Mitt.) Grolle .......... Mochila hirta Tayl. g_x_ Mitt ............. . Acrobolbus ochrophyllus (Hook. f. 8 Tayl.) Schust. . . . . Austrolophozia fuegiensis (Steph.) Schust ........ liarsu idium urvillianum (Mont.) Mitt. Open circle n cates precise locality unknown .......... _Cgmialoziella dusenii Steph ............... xii Page 588 588 588 588 590 590 590 590 592 592 592 594 594 594 594 596 596 596 596 598 LIST OF MAPS - continued Nap 53. Adelanthus lindentmgianus (Lehm.) Mitt. North to 55. 56. 57. Venezuela and Costa Rica; plus Ireland, after Grolle (l969). . ................... Nettsteinia densiretis (Herz.) Grolle .......... Aadula helix (Hook. f. & Tayl.) Grolle ......... Jensenia pisicolor (Hook. f. & Tayl.) Grolle ...... Pallavacinia xMoides (Hook. f. & Tayl.) Trev ...... . Sm h 0 na hochstetteri Nees & Mont. Open circle in icates precrse locality unknown .......... . hetzgeria decipiens (Mass.) Schiffn. Adapted and modified from Kuwahara (I966) ............. . Metzgeria leptoneura Spruce. Adapted and modified from Kuwahara (T966) .................... . Map showing mean annual rainfall at stations in the Falkland Islands for which records are available for six or more years (data extracted from Moore, l968). . Map of personal collection localities .......... . South polar projection showing delimitation of subantarctic region (after Greene, l964) ....... xiii Page 598 598 598 600 600 600 602 602 604 606 608 THE FALKLAND ISLANDS A. Location The Falkland Islands lie between 5l° 00' and 52° 30‘ S., and 57° 40' and 6l° 30' II. in the South Atlantic Ocean, 520 km. east of the Strait of Magellan. The Falklands are an archipelago of more than 230 islands, comprised of two principal islands, East Falkland with adjacent small islands making up ca. 5,000 sq. km. and West Falkland, with adjacent small islands, ca. 3,500 sq. km. The Falkland Islands were previously settled by the French, who referred to them as Iles Malouines. The islands are currently administered as a British crown colony, but claimed by Argentina and referred to as Islas Malvinas. 0n various Argentine maps, East Falk- land Island is known as "Isla Soledad" and West Falkland Island as "Isla Gran Mal vina." B. Topographical Features In general, the Falklands are made up of generally hilly ter- rain. The southern portion (Lafonia) of East Falkland is low and flat. The principal mountainous areas extend east-west in the north of the islands, with Mt. Kent (458 m.) and Mt. Usborne (705 m.) in East Falkland, and Mt. Adam (750 m.) , Mt. Fegan (360 m.) and Byron l Heights (520 m.) in West Falkland. The Hornby Mountains (including hit. Maria, 658 m., Muffler Jack Mt., 547 m. and Mt. Moody, 554 m.) lie in Ilest Falkland and roughly parallel Falkland Sound. The small islands occasionally reach quite high altitudes e.g., Keppel Island (ht. Keppel, 342 m.), Saunders Island (Mt. Harston, 433 m., Rookery lit, 422m. and Mt. Rees 372 m.) and Weddell Island (Mt. Weddell, 380 m.). Alarge number of valleys, small rivers, streams and boggy areas are present (an the islands. C. Geolggi cal Features The very brief and general account of the geological features provided here merely summarizes the major rock types occurring in the Falklands. Detailed accounts may be found in Adie (I952 a,b, I953, TISB), Andersson (T907), Baker (l922), Cawkell et al . (1960) Halle (l9l2) and Riggi (T934). The following review is based primarily on the summary in Cawkell et al. (T960). The Falklands are almost entirely composed of Palaeozoic and Mesozoic sedimentary rocks. Virtually the only metamorphic and ig- neous rocks are restricted to Cape Meredith, the southernmost tip of the Falklands. The Falkland Islands lie on the edge of the Patagonian continental shelf, but as Cawkell et al. (339. _c_i_t_.. p. 173) state. “there is no stratigraphical connection betweenthe I; Falklands and the nearer parts of the South American mainland." The Falkland rocks have strong lithological and structural affinities with those of South Africa, Uruguay and southern Brazil. There is a strik- ing similarity between the marine invertebrate and plant fossils of the Falklands and South Africa, and this evidence, along with studies on fold systems, was used by Adie (l952b) to suggest that during the late Palaeozoic-early Mesozoic, the Falklands were in an inverted position some 250 km. east of the Eastern Province, South Africa, and after fragmentation of Gondwanaland, the Falklands drifted to their present position. Today there are two main morphological elements which may be recognized within the Falklands. With the exception of the Cape Meredith region, the remainder of West Falkland and the northern half of East Falkland are composed of Devonian sandstones and quartz- ites. The quartzite is particularly evident as conspicuous parallel ridges which run east-west along the folding zone. The sandstone occurs principally in valley bottoms (Skottsberg, 19l3). The southern region (Lafonia) of East Falkland is built up of Permo-carbonian sand- and claystones. The stoneruns, which are so conspicuous in West and northern East Falkland are absent in Lafonia (Skottsberg, Log. 93.). The only limestone occurring in the Falklands is the raised beach at Shell Point, near Fitzroy, East Falkland. This region was not per- sonally visited. There is no evidence to suggest that the Falkland Islands SUAborted permanent glaciers during the Pleistocene. It is believed that even during the glacial maxima, periglacial conditions persisted IL in the islands. However, evidence has shown a Permo-Carboniferous ice sheet existed on the Falklands and this glacier deposited a consider- able thickness of tillite. D. Soils In the absence of any comprehensive studies on the soils of the Falklands, only a few general remarks will be made here. For further information see Davies (T939), Moore (T968) and Skottsberg (l9l3). Podsolization is probably favored by the generally acid condi— tions, combined with Teaching and lack of disturbance (Moore, T968). Peat accumulation is widespread, the extent depending largely on local drainage. As Moore (129. git”, p. 5) indicates, "undegraded plant remains normally constitute a high proportion of the top soil, which may vary from a shallow, hard, dry peat under gym on the quartz- ite ridges to a black, plastic humus under the Chiliotrichum scrub that fringes, rivers and creeks." E. Climate The Falkland Islands lie on the northern edge of the principal depression belt through Drake Passage (Cawkell et al., T960). These islands are consequently dominated by the westerly winds, fronts and airmasses moving across this region. Although the Falklands are influenced by their position on the lee side of the Andes the 520 km. of Open sea greatly tempers this effect. Precipitation.--Moore (T968) has assembled the mean annual precipitation figures for those localities with records available for six or more years. This data is presented in Map 61. It may be ob- served that the drier areas of the islands are in the south, and the wetter areas in the north. The stations with highest available fig- ures lie on the immediate lee sides of mountain ranges, and a gen- eral gradient may be observed from west to east in either island, with Port Howard the wettest station in West Falkland and Port Stanley one of the wettest in the East Falklands. The 609 mn. figure for Port Stanley is second only to the 636 mm. of Port San Carlos. The Port Stanley data in Moore (T968) may be conservative, as Davies (T939, p.2) records "in the order of" 635 mm., and Cawkell et al. 0960, p. l88) records "about" 686 mm. for the capital. Moore (T968) points out that the low rainfall of Pebble Island could result from its lying in the “rain shadow" of the Mount Adam complex. The dif- ference between San Carlos and Port San Carlos may be due to The Verde Mountains which lie between the two stations or to some other local factor. Cawkell et al. (T960) attempt to explain the slight summer maximum in rainfall figures for Port Stanley through the greater tend- ency for convection and showers in the northeast part of East Falk- lands. The air has become moist and unstable after it has crossed the Drake Passage, and it is pointed out that an unstable air mass approaching the Falkland Islands from the southwest will produce only cumulus cloud cover at Fox Bay and Goose Green. However, after this airmass has been sufficiently heated from the land, and perhaps been uplifted by high ground west of Port Stanley, it is able to produce showers in the northeast region of the islands. It is likely that this phenomenon of heating and uplifting is occurring on both island groups and accounts for the data obtained. The rainfall is distributed fairly even throughout the year, with a slight maximum in summer (December-January). The number of days with rainfall is high (ca. 250 days) with few days of excessive rainfall. The rainfall intensity is thus low, and daily falls ex— ceeding l3 mm. are rare (Cawkell et al., T960). Temperature.--The temperature data for Port Stanley is as follows (after Moore, T968): . . . Mean Mean Mean Minimum Maxamum Minimum Maximum January 8.8 O 23.9 5.4 l2.9 July 2.2 -7.8 8.9 O 4.3 Vhile temperature data is available from only Port Stanley, Fox Bay and Vestpoint Island, Moore (p. 7) states there is a "suggestion of increasing temperatures towards the west and south of the islands." Humidity, Cloud and Sunshine.--Cawkell et al . (T960) indicate that the relative humidity average values are high; the mean annual value is 82% at 0900 ZT, and varies from a mean of 75% in November to90i in July. The Falkland Islands witness a large number of over- cast days. The average annual duration of sunshine is less than 35% of the total possible for the latitude of Port Stanley (Cawkell et al., Lie 31L). , , " ' ficant collections of hepatica have been those y; '7' -' and Halle, while those of Lechler and Cunningham @pérous. Below is a chronological list of collectors V 3&1 Titanium data. b A m. -- a- .Auvrofa v.19»... In H 244)..) .0 .i. I I ll! AVll nan. .. u .. a .A 52 :85 c2283 :85 NomPIFomF capo; ucuv a .53 .95 $8: 2283 2 Egg to; mm "883.8% ‘28,; 88399...” . mom xom ”J3 mcnwgmocoe mzowcm> mucm_mH comma» cw magma uvmcmansncs Ear—Pr: pczoz 2m upon: me :ovuumFFoo mopcopm “so; new mmusom— gamma: Easmcrccao 2m czo=xcs cormmwp upena< mznmgmocos mzowem> cw magma mumgmwpnaqca anppw: peso: . Fum Laden» tang an. zm .zm Aeewpv capsup a Loxoo: mm smwppvz ago; new vaF a msaosu Ax=FFoumcu .pcoz «av .Anmm—V ocomvvow :oEwm peso: . Awwwpv uznzupvsnw p mwzoa ugom "mm NNm— uppvammu" o::o_o M Awmwpv usasoeuaeo m mesa; use; new owrupop a mfiumma secea< ocvsnppnaa wwwowmm F392 u:.uuo__ou mwhuwwwmoo eeeu.ueexu, . \ Nauk‘tu: KG “Kuw.—[03400 «50 L." H J 1—(0 HUOJOZOKIU .mco_pum__oo ucmpmH ucm_x_mu m nzmgopvsm . . w opmoop op muaEmuum :_ . _zemmmoozmca coma w>m 3 mm .Ammmpv meooz seem xppmapocwga umpomguxm mama _ zmHm .Apom_ .~oucmucoz mu _mmmm: «new .4H4 5 52333 ozzo cw zoom :owuum—Foo G .ma: .(mw appopv econooum omen »m_cmpm pee; "do czocxc: .A__mp .meanmsuoxm mom maze; Loev mucmFmH 3oz a Lm>mwm .ppouuwz .mmogm _msop_;a pass a m>ou sax .ucumH ucvoa “mm: ou o>oo ~_F: a sum zopfiagm mw> yum: .ucmzo: ugoa o» hum xom “m3 tampon ppozuowam one EL< cugoz 0» :930m ocm ovcoenJ CLocuxoc mopgou com Ocsonma 9:302 Lovo>~om HLOQ mm Hopcoum uLoa "mm flIIIIIIIIIIII . ommF Amvmgomm nuaom firrrrlrrrrr _Lwasoooo . ow wusogcmv elm xu>ezm Adcmucmamo mvcwpmm acupxFuJ mcomeu czocxc: mcwzuuwx MNmF COHFwEMI copuwvuaxw m0-uI Owcmpnwmmz Ucuv momprnom~ zwroozm mLoonuuoxm TO Below is a list of my collection numbers together with collec- tion data. A statement of the ecology is made for nearly every local- ity since ecological data has been omitted under the specimen citation portion of each series. The localities are indicated on Map 62. All localities are cited with reference to Universal Transverse Mercater Grid (Zone 21) co-ordinates as shown on maps of Falkland Islands l:50,000 (Directorate of Overseas Surveys 453, Series H79T , T96l). This map is included in Moore (T968) . 2369-239l 2392-2430 243l-2440 244l-2442 2443-245l 2452-2455 2456-2478 2479-25l3 25l4-2556 EAST FALKLANDS: UTM Grid 2lF VC 3472. Stanley: Dwarf shrub heath & outcrops on Tumbledown Mt. , l55-230 m. - 3 January T968 EAST FALKLANDS: UTM Grid 2TF VC 387T. Stanley: Outcrop on Sapper Hill, T35 m. - 4 January I968 EAST FALKLANDS: UTM Grid 21F VC 4772. Port William: Dwarf shrub heath on S shore of Cape Pembroke peninsula, near Surf Bay. 5 January T968 EAST FALKLANDS: UTM Grid ZTF VC 4773. Port William: Outcrop in dwarf shrub heath on central part of Cape Pembroke peninsula between Surf Bay & Kelly Rocks, c. l5 m. - 5 January T968 EAST FALKLANDS: UTM Grid 2lF VC 4674. Port William: Dwarf shrub heath on N shore of Cape Pembroke peninsula E of Yorke Pt. 5 January T968 EAST FALKLANDS: UTM Grid 21F UC 7l7l. Mt. Usborne: Cortaderia heath on S side of The Gap, 260-275 m. - 6 January T968 EAST FALKLANDS: UTM Grid 21F UC 7l7l. Mt. Usborne: Cortaderia heath at The Gap, 275-290 m. - 6 January T968 EAST FALKLANDS: UTM Grid 2lF UC 737T. Mt. Usborne: Feldmark on summit of Mt. Usborne l, c.7OO m. - 7 Janu- ary T968 EAST FALKLANDS: UTM Grid 2lF UC 737T. Mt. Usborne: Sheltered cliffs with seepage on ridge between Mt. Usbornes l & 2, 685 m. 7 January T968 2557 2558-2579 2580-2592 2593 2594-262l 2622-2640 264l-2646 2647-2675 2676-2679 2680-2687 2688-2699 2700-2738 ll EAST FALKLANDS: UTM Grid 21F UC 7370. Mt. Usborne: Stonerun in Cortaderia heath on S slope of Mt. Usborne 1, 550 m. 7 January 1968 EAST FALKLANDS: UTM Grid 21F UC 7069. Mt. Usborne: Cortaderia heath below The Gap, c. 90 m. - 8 January I968 EAST FALKLANDS: UTM Grid 21F UC 7069. Mt. Usborne: Stonerun in Cortaderia heath below The Gap, c. 90 m. - 8 January 19 EAST FALKLANDS: UTM Grid 21F UC 7070. Mt. Usborne: Stonerun at base of Cantera Mt. , c. 90 m. - 8 January 1968 EAST FALKLANDS: UTM Grid 21F UC 7569. Mt. Usborne: Seepage area in Cortaderia heath on SE slope of Mt. Us- borne 2, c. 455 m. 9 January 1968 EAST FALKLANDS: UTM Grid 21F UC 7669. Mt. Usborne: Marsippospermum-mire in Cortaderia heath on gap between Mt. Usborne 2 81 Table Rock, c. 440 m. - 9 January 1968 EAST FALKLANDS: UTM Grid 21F UC 7468. Mt. Usborne: Cortaderia heath on gap between Mt. Usborne 2 & Ceritos Rocks, 475 m. 9 January 1968 EAST FALKLANDS: UTM Grid 21F UC 7068. Mt. Usborne: Cortaderia heath with stream and occasional sandstone outcrops in valley SW of Mt. Usborne, c. 60 m. - 10 January 1968 EAST FALKLANDS: UTM Grid 21F UC 6662. Darwin Settle- ment: Cortaderia heath near mouth of Ceritos Arroyo. 10 January 1968 EAST FALKLANDS: UTM Grid 21F UC 6457. Darwin Settle- ment: Coastal cliffs near dwarf shrub heath on S side of Carcass Bay, Darwin Harbour. 11 January 1968 EAST FALKLANDS: UTM Grid 21F UC 6359. Darwin Settle- ment: Dwarf shrub heath at Boca House on Brenton Lock. 11 January 1968 EAST FALKLANDS: UTM Grid 21F VC 3073. Stanley: Out- crops in dwarf shrub heath on summit ridge of N peak of Two Sisters, 245-290 m. - 13 January 1968 12 2739-2774 EAST FALKLANDS: UTM Grid 21F VC 2374. Stanley: Cliffs on rock dome at summit of Mt. Kent, 455 m.; in dwarf shrub heath - 14 January 1968 2775-2780 EAST FALKLANDS: UTM Grid 21F VC 3074. Stanley: Peat " bog in valley N of Two Sisters, 75-90 m. - 14 January 1968 2781-2803 EAST FALKLANDS: UTM Grid 21F VC 4880. Kidney Island: Tussock grassland N of Shanty, c. 12 m. - 16 January 1968 2804-2812 EAST FALKLANDS: UTM Grid 21F VC 4880. Kidney Island: Coastal rocks at margin of tussock grassland on SE shore between landing bay 8 SE Pt. - 17 January 1968 2813-2840 EAST FALKLANDS: UTM Grid 21F VC 4373. Port William: Dwarf shrub heath on ridge on Engineer Point peninsula, 18-25 m. 19 January 1968 2841-2843 WEST FALKLANDS: UTM Grid 21F TC 4202. Westpoint Island: Pastureland in Misery Valley, 215-230 m. - 20 January 1968 2844-2878 WEST FALKLANDS: UTM Grid 21F TD 4302. Westpoint Island: Hebe-scrub in dwarf shrub heath near The Waterfall, 30-90 m. 720—January 1968 2879-2880 WEST FALKLANDS: UTM Grid 21F TD 4006. Westpoint Island: Coastal rocks at margin of tussock grassland along NE shore adjacent to Cape Terrible. - 21 January 1968 2881-2895 WEST FALKLANDS: UTM Grid 21F TD 4104. Westpoint Island: Dwarf shrub heath at W base of Black Hog Hill, adjacent to Devils Nose, c. 18 m. - 21 January 1968 2896 WEST FALKLANDS: UTM Grid 21F TD 4204. Westpoint Island: Pastureland on divide between Devils Nose & Cat Cove, 60 m. 21 January 1968 2897-2940 WEST FALKLANDS: UTM Grid 21F TD 4404. Westpoint Island: Hebe-scrub in dwarf shrub heath on steep slope & cliffs TEE-ing The Woolly Gut. - 22 January 1968 2941-2942 WEST FALKLANDS: UTM Grid 21F TD 4404. Westpoint Island: Pastureland on SE slope of settlement bay, c. 60 m. - 22 January 1968 2943- 2963 WEST FALKLANDS: UTM Grid 21F TC 8087. Hill Cove: Out- crops on summit of West French Peak, 290 m.; in dwarf shrub heath - 24 January 1968 L 2964-2973 2974-2984 2985-2992 2993-3002 3003-3006 3007-3027 3022-3047 3048 049-3056 3057-3062 3063-307 9 soap-3735 13 WEST FALKLANDS: UTM Grid 21F TC 8187. Hill Cove: Cliffs on summit of East French Peak, 305 m. , in dwarf shrub heath. 24 January 1968 WEST FALKLANDS: UTM Grid 21F TC 8087. Hill Cove: Cortaderia-heath with stream, gap between French Peaks, 24 January 1968 WEST FALKLANDS: UTM Grid 21F TC 8781. Mt. Adam: Out- crops on summit of southermost peak, 685 m. - 25 January - me 1968 WEST FALKLANDS: UTM Grid 21F TC 8781. Mt. Adam: Corta- deria -heath with stream in basin E of summit, 580-595 m. 25 January 1968 UTM Grid 21F TC 8781. Mt. Adam: Corta- WEST FALKLANDS: deria-heath with outcrops on E slope of main peak, c. 610 m. - 25 January 1968 WEST FALKLANDS: UTM Grid 21F TC 8781. Mt. Adam: Shel- tered cliffs on E side of summit ridge, 670-700 m. - 25 January 1968 WEST FALKLANDS: UTM Grid 21F TC 8782. Mt. Adam: Shel- tered cliffs on ridge S of northern Take, 610 m. - 25 January 7968 WEST FALKLANDS: UTM Grid 21F TC 8686. Hill Cove: Creek in dwarf shrub heath, valley between Mt. Donald & Mt. 25 January 1968 Adam, 180 m. WEST FALKLANDS: UTM Grid 27F TC 8l88. Hill Cove: Grove of planted trees (mainly hardwoods) in settlement, 15 m. 26 January 1968 UTM Grid 21F TC 7289. Hill Cove: Out- WEST FALKLANDS: crops 8 dry feldmark on ridge of N slope of Mt. Fegen, c. 275 m. 26 January 1968 Hill Cove: Cliffs WEST FALKLANDS: UTM Grid 21F TC 7289. in dwarf shrub heath on summit of Mt. Fegen, 335-360 m. 26 January 1968 WEST FALKLANDS: UTM Grid 21F UC 2078. Port Howard: Sheltered cliffs on pass SW of Mt. Maria summit, c. 610 m. 28 January 1968 WEST FALKLANDS: UTM Grid 21F UC 2079. Port Howard: Feldmark on summit of Mt. Maria, 658 m. - 28 January 1968 ‘9‘ 14 131-3154 WEST FALKLANDS: UTM Grid 21F 00 2279. Port Howard: Large outcrops in dwarf shrub heath, Freezer Rocks, 320 m. 28 January 1968 3155-3186 EAST FALKLANDS: UTM Grid 21F WC 3270. Stanley: Mosaic of dwarf shrub heath 8. peat bogs, headwaters of Mullet Creek Stream, c. 60 m. - 30 January 1968 3187-3217 EAST FALKLANDS: UTM Grid 21F WC 2971. Stanley: Out- crops in dwarf shrub heath along Goat Ridge, c. 180 m. 30 January 1968 3218-3239 EAST FALKLANDS: UTM Grid 21F VC 4379. Port William: Outcrops in dwarf shrub heath on summit ridge of Mt. Low, c. 245 m. 31 January 1968 3240-3262 EAST FALKLANDS: UTM Grid 21F WC 4474. Port William: Outcrops in dwarf shrub heath on N side of Gypsy Cove. \ 1 February 1968 3263 WEST FALKLANDS: UTM Grid 21F TC 0164. New Island: Dry feldmark on summit of cliffs at NW tip opposite Landsend Bluff, 90 m. 2 February 1968 1264-3218 WEST FALKLANDS: UTM Grid 21F TC 0863. New Island: Scattered outcrops in dwarf shrub heath along ridge be- tween Bold Hill 8 Bold Pt., 105 m. - 3 February 1968 3219-3280 WEST FALKLANDS: UTM Grid 21F TC 0864. New Island: Pastureland S of Bold PT., 30 m. - 3 February 1968 3281-3286 WEST FALKLANDS: UTM Grid 21F TC 0258. New Island: Sandstone bluffs (seal caves) at South End. - 4 February 1968 3281-3300 WEST FALKLANDS: UTM Grid 21F TC 0362. New Island: Seep- age cliffs at margin of dwarf shrub heath along shore of Ship Harbour N of $th Island. - 4 February 1968 3301-3332 WEST FALKLANDS: UTM Grid 21F TC 2842. Weddell Island: Dwarf shrub heath in Waterfall Valley, west of settle- ment, c. 125 m. 5 February 1968 3133-3353 WEST FALKLANDS: UTM Grid 21F TC 2839-2840. Weddell Island: Dwarf shrub heath at summit of Mt. Weddell, 380 m. - 6 February 1968 3354-3400 WEST FALKLANDS: UTM Grid 21F TC 2941. Weddell Island: Rock done on summit of peak NE of Mt. Weddell, 335 m.; dwarf shrub heath. - 6 February 1968 3402-3415 3416-3443 3444-3456 3457-3476 3477-3500 3501-3521 3522-3538 15 WEST FALKLANDS: UTM Grid 21F TC 2842. Weddell Island- Dwarf shrub heath on slope SW of settlement, 180 m. - 6 February 1968 WEST FALKLANDS: UTM Grid 21F TC 3043. Weddell Island: Dwarf shrub heath near headwaters of House Creek, c. 15 m. - 7 February 1968 WEST FALKLANDS: UTM Grid 21F TC 9334. Fox Bay: Dwarf shrub heath at summit of East Head, 180 m. - 7 February 1868 WEST FALKLANDS: UTM Grid 21F TC 8536. Fox Bay: Out- crops at summit of Fox Bay Mt. , 307 m.', dwarf shrub heath. 8 February 1968 WEST FALKLANDS: UTM Grid 21F TC 8537. Fox Ray: Dwarf shrub heath on N base of Fox Bay Mt. , 75 m. 8 February 1968 WEST FALKLANDS: UTM Grid 21F TC 9039. Fox Bay: ture near mouth of Cheeks' Creek, c. 12 m. - 9 February 1968 Pas— WEST FALKLANDS: UTM Grid 21F TC 9341-9441. Fox Bay: Mosaic of dwarf shrub heath 8. peat bogs near peat cut- tings in Ram Paddock, 45-60 m. - 10 February 1968 WEST FALKLANDS: UTM Grid 21F TC 8950. Fox Bay: Dwarf shrub heath in sandy area in stream valley E of Sullivan House, c. 75 m. - 11 February 1968 III . ECOLOGY A. Introduction Skottsberg (1913) made a vegetational analysis of the vascular plants of the Falkland Islands, and Moore (1968) based his account of the Falkland vegetation primarily on that of Skottsberg. Brief accounts of the natural vegetation are also provided in Davies (1939) and Cawkell et a1. (1960). In summarizing the bryophyte ecology in the Falkland Islands, I have utilized the community group- ings and terminology of Moore (Log. 5131.). I have, however, added one enormity grouping, i.e., the sheltered high altitude cliff vege- tation, which is primarily cryptogamic. As emphasized by Moore the predominant Falkland plant associations are closely related to one another. The associations and formations, especially those of lowland areas. frequently are not arranged as discreet units, but a kind of mosaic of community groupings is rather the pattern. Moore (1968) has recognized five formations. In addition, he has recognized four associations (Senecio candicans. Ammophila- ilmg. Eleocharis and Mfliophyllum) which do not merit formation Webbing. These are arranged according to types of vegetation. l6 17 8. Distribution of Bryophytes in Plant Communities of the Falkland IsTands l. MARITIME TUSSOCK FORMATION a. POA FLABELLATA ASSOCIATION This primarily coastal association was previously far more prevalent than observed today (see Hooker 1847, Moore 1968). The un- restricted, extensive grazing of sheep has been primarily responsible for the depletion of this formation. Pigs, which destroy plants by digging up the roots, are of lesser significance. Tussock grass to— day is confined to smaller offshore islets (e.g., Kidney Island) which have not been stocked with sheep, and a few fenced off areas on the larger islands. On Kidney Island, where there are extensive areas of almost pure stands of Pia flabellata, this association has reached its maximum development (see Photo 1). ' As discussed in Moore 119681.103 flabellata tussocks can attain a height of 2-3 m. with each tussock composed of a fibrous stock (1.5 - 2.0 m. high and 1.0 - 1.5 m. in diameter) surmounted by a dense crown of leaves. In areas where the community is uninterrupted, the dense canopy of interlaced leaves excludes all associated species except Carex trifida (see Photo 2). On the tussock bases L_ophocolea Tenta and Telaranea pseudozoopsis were particularly common, and Cephaloziella dusenii, Wdivaricata (very rare), L. leptantha, 1:. sabuletorum, l141'_C_hg_gt_i_a_berteroana var. polylepida and Campylium so. were also present. Lgphocolea lgatantha, L. sabuletorum and Hygroamblystegium 18 WW. skottsbergii were found on soil between the tussock bases. On the coastal cliffs of Kidney and Westpoint Islands, at the margin of the tussock community, Lophocolea sabuletorum was common on rock and in rock crevices. but Muelleriella crassifolia was also pres- ent. Luphocolea divaricata is the only hepatic restricted to this association. 2. OCEANIC HEATH FORMATION This formation has been subdivided into two groups. Cortaderia pilosa is dominant or co-dominant in one and dwarf shrubs are prominent in the other. Most of the Falkland Islands are covered by one facies or another of these associations. a. CORTADERIA ASSOCIATION This association, as Moore (1968, p. 15) states, "can be de- veloped on most non-swampy ground having indifferent drainage. It is widesoread on level or undulating country below 100 m. . . . but it is also very common on gentler slopes up to an altitude of ca. 180- 200 m." (see Photo 3). I have found this association, which is dom- inated by Cortaderia pilosa, occurring at various altitudes up to ca. 610m. The grass blades form an excellent protective cover against the sun and especially the winds, while the coarse fibrous peaty humus which develops under the tussocks retains considerable moisture. These factors contribute to a protected environment in which a surprisingly large number of bryophyte species may be found. This association includes several facies, each differing in the means of providing 19 protection and moisture conservation. These facies will be discussed separately. Bryophytes directly associated with Cortaderia, in depressions, pockets or holes, and deriving protection and moisture from the asso- ciation are as follows: Adelanthus lindenbergianus, _A_. Legals (only above 245 m.), Anastrepta bifida, Balantigfiis cancellata, Blepharido- phyllumclandestinum, B. gottscheanum, Cheilolejeunea savatieriana (c. 610m), Chiloscyphus hookeri subsp. hookeri (275-290 m.), Clasmatocolea humilis, Jamesoniella colorata (275-290 m.), Lepidozia Mme-290 m.), Leptoscthus patagonicus, Microlepidozia mollis (275-290 m.), Riccardia georgiensis, Saccogynidium australe, _S_. gaggi- l_o_§u_m, Telaranea plumulosa, l. pseudozoopsis (275-290 m.), Wettsteinia densiretis, Calliergon sarmentosum, Conostomum australe, Dicranoloma robustum, D. setosum, Racomitrium lanuginosum, Sphagnum fimbriatum, S. magellanicum and Webera albicans. The stream facies.-—Streams are frequently present in the Massociation (see Photo 4), and often the bank vegetation (which frequently is Gunnera magellanica) or the stream itself offers protective niches for bryophytes. Bryophytes of the stream or stream bank habitat are Balantiopsis bisbifida, B. cancellata, 8. erinacea, _ClgsgiLtocolea humilis, C. vermicularis, Isotachis humectata, Waustrigena (very common), _L_. M, _L_. sabuletorum, _l_._. Mg, Metahygrobiella tubulata, Noteroclada confluens, Palla- 1.451313 xiphoides, Pseudocephalozia tristaniana, Riccardia fuscobrunnea, 9m. 5. gpgctabilis, Schistochila subintegerrima, Telaranea plumulosa, 1. pgudozoopsis, Blindia torrentium, Drepanocladus fluitafi, L'\ 20 Philonotis vagens 9.0.9. Sciaromium conspissatum. Lgphocolea austrigena and L. elata are the only Hepaticae which are nearly always submerged in these streams. The wet depression facies.--Wet depressions, where the peat is saturated and small pools may occur, provide another means of protection (see Photo 5). Here Adelanthus tenuis, Clasmatocolea humilis, Jensenia pisicolor, Paracromastigum subsimpl ex, Riccardia oountiformis, R. pailidevirens, R. Lapillosa, Telaranea flumulosa, Temnoma quadripartitum and Sphagnum magellanicum may be found. The stonerun facies.--Stoneruns are of frequent occurrence in the Cortaderia association (see Photos 6-7). Often cool, damp, shaded crevices are present and here occur Clasmatocolea fulvella, Frullania gagellanica, Jamesoniella colorata, Lejeunea corralensis, Lophocolea sabuletorum, Metzgeria decipiens, Plagiochila ansata, Andreaea verru- c_ul_o§g and Disticflphyllum cavifolium. The rock outcrgp facies.--On rock outcrops (particularly of exposures protected from prevailing winds) with their crevices, over- hangs and small niches as well as on soil immediately below the out- crops occur Chiloscyphus hookeri subsp. hookeri, Clasmatocolea firi- yellflabove 580 m.), C. humilis, _C_. koeppensis (475 m.), Leptoscyphus whicus, gaphocolea austrigena, Pachyglossa spegazziniana var. @1115, Riccardia pgllidevirens, Telaranea plumulosa, Andreaea 0952115 A. parallela and Racomitrium Tanuginosum. The Cortaderia association, while not possessing the bryophyte Species diversity of the closely related dwarf shrub heath, possesses the greatest assemblage of rare Falkland species, several of which are 21 also rare in southern South America. Hepaticae unique to this asso- ciation are: Anastrepta bifida, Blepharidophyllum gottscheanum, £53.. tachis humectata, Lejeunea corralensis, Microlepidozia mollis, Daracromastigum subsimplex, Pseudocgahalozia tristaniana, Riccardia pallidevirens and Saccogynidium australe. b. DWARF SHRUB HEATH ASSOCIATIONS Moore (12;. _c_i_t., p. 15) states that The communities included here are usually found on com- paratively dry ground, being best developed on rocky ridges, stony areas or places where the immediate sub- soil is of relatively coarse material so that drainage is good. A hard, dry peat underlies well-developed heath. The principal species in the dwarf shrub heath are metrum rubrum, Pernettyg pgmila, Blechnum magellanicum, _B_. penna-marina, Baccaris magellanica and Bolax wifera. Various combinations of these may become co-dominant, and Moore (1968) has listed the following asso- ciations: E_mpetrum rubrum-Blechnum penna-marina, Empetrum rubrum- Blechnum m_agellanicum, Empetrum rubrum-Pernettya pumila, Bolax gmmifera-Eflietrum rubrum and Blechnum magellanicum-B. Ema-marina. Moore (l_9_c_.fl., p. 15) states "The Empetrum rubrum association is much the most common" and the discussion below pertains to this asso- ciation. As in the Cortaderia association, there are several means of protection against the sun and winds as well as the conservation of adequate moisture so that this association has a fairly rich bryophyte flora. Each local facies will be discussed separately. 22 Bryophytes commonly grow on the bases of Blechnum magellanicum, the fronds of which provide an excellent protective cover, particu- larly when interlaced with fronds of other Blechnum plants or branches oiE_np_et__rp_lg(see Photo 8). The soil below such situations often has aconsiderable bryophyte cover. Frequently Bglax cushions, particu- larly old, decayed portions, have a bryophyte (mostly hepatic) cover. In these situations occur Adelanthus lindenbergi anus , Balantigasis bisbifida, B. cancellata, Blepharidophyllum cl andestinum, B. densi— folium, Qphaloziella dusenii, Chilosgyphus hookeri subsp. hookeri, Clasmatocolea humilis, C. vermicularis, Frullania boveana, Gackstro- egigmgellanica, Hyalolepidozia bicuspidata, Jamesoniella colorata, L_egidozia chordulifera, L_. laevifolia, Leptoscyphus aequatus, I_._. expansus, _L_. patagonicus, Lgahocolea Tenta, L_. leptantha, 1;. sabuletorum, _L_. sylvatica, Marchantia berteroana var. p_ol yl epida, Metzgeria decipiens, M. I_eptoneura, Plagiochila ansata, _P__. el ata, _P. hirta, Riccardia ggorgiensis, R_. opuntiformis, R. papillosa, _R_. §pe£tabilis, Roivainenia ja_cquinotii , Telaranea blepharostoma, L plumulosa,T. pseudozogpsis, Wettsteinia densiretis, Anisothecium hookeri, Blindia curviseta, mum macrophyllum. Bryum (subsect. Doliolidium) sp., CampyIOpus introflexus, Ceratodon purpureus, Chgrisodontium aciphyllum, Distichophyllum cavifolium, p_. dicksoni, D. flaccidum, Dicranoloma fal kl andicum, D_. robustum, Fissidens Ligidulus, Polytrichadelphus robustus, Polytrichum strictum, figgdodistrichium austrogeoLgicum, Racomitri um lanuginosum, I}; rupestre and Tortula antarctica. 23 The stream facies.—-Streams frequently occur in the Empetrum mbrgg association. Vascular plants such as Blechnum magellanicum, B. penna-marina. E_mpetrum rubrum and Gunnera magellanica frequently offer a protective cover, which may be very dense, and at the same time provide cool, moist niches for bryphytes (see Photo 9). Several Species grow partially or wholly submerged in the streams, Lophocolea Wand I_._. wanearly exclusively so. This facies, when compared to the stream facies of the Cortaderia association, while having the same Lophocolea species, has quite a different species compliment as well as greater species diversity. The bryophytes of this habitat include Adelanthus tenuis, Balantiogsis bisbifida, B. M, B. erinacea, Chiloscyphus hookeri subsp. hookeri, Clasmatocolea humilis, C. vermicularis, Jamesoniella colorata, Jensenia Eicolor, Jungermannia crassula, Lepidozia chordulifera, L. laevifolia, Leptoscyphus gpansus, Lephocolea austrigena, L_. 91.9.5.9..- L. sabuletorum, L_. sylvatica, Marchantia berteroana var. polylepida, hgtgroclada confluens, Pallavacinia xiphoides, Riccardia ggorgiensis, R. 29.217151- 3. spectabilis, Saccogynidium vasculosum, Smphyogyna hygenOphyllum, Telaranea Llumulosa, L pseudozoopsis, Bartramia patens, Mlaevigatum, Campylcgaus introflexus, Fissidens rigidulum, Lgpgotheca ggudichaudi , Polytrichum strictum, Racomitrium subnigritum and Sphagnum fimbriatum. The wet depression facies.--Like the Cortaderia association the Mm association has shallow depressions where the peat is saturated and small pools may occur. The habitat has a characteristic 24 bryophyte flora with Balantiopsis erinacea, Clasmatocolea humilis, Metahygrobiella tubulata, Saccogynidium vasculosum (rare), 9311.44.42 macrophyllum, Drepanocladus longifolius, Sphagnum engelii and _S_. fimbriatum. Sphagnum falcatum was present in depressions where free standing water occurred. The rock outcr%facies.--Rock outcrops of varying sizes are frequently present in this association, with protection provided in amanner similar to that in the rock outcrop facies of the Cortaderia association. Bryophytes occurring in the various niches associated with the outcrops, as well as on soil at the base of the outcrOps, are Adelanthus lindenbergianus, A. tenuis, Andrewsianthis australis, A. Llanifolius, Austrolophozia fuegiensis, Cephalolobus scabrellus, gphaloziella dusenii, Cheilolejeunea savatieriana, Chiloscyphus Msubsp. hookeri, Clasmatocolea fulvella, C. humilis, _C_. heppensis, Frullania boveana, 5. magellanica, _F_. gaudolobulata, figckstroemia magllanica, Harpalejeunea marginalis, Jamesoniella colorata, L_epicolea ochroleuca, Lepidozia chordulifera, I_._. laevifolia, Laptoscyphus abditus, I_._. expansus, L_caphocolea austrigena, _L_._. Lent-a, Lophozia hatcheri, Marchantia berteroana var. polylepida, Marsupidium milleanum, Metzgeria decijiens, M. falklandica. I_Vl_. multiformis, Eaghyglossa gpegazziniana var. exilis, Plagiochila ansata, _P_. elata, E. gayana, P. hirta, P_. obovata, Riccardia opuntiformis, 3. papillosa, B. pgehensilis, Schistochila pgghyla, _S_. subintegrrima, Symphyogyna Windphyllum, Telaranea p1 umulosa, '_I'_. pseudozoopsis, Temnoma ggadwi- partitum, Wettsteinia densiretus, Acrocladium auriculatum, Andreaea 25 subulata (exposed rock surfaces), Bartramia patens, Bryum cf. Hatcheri, Campylogus introflexus, Catagonium nitidum, Ceratodon purpureus, Chorisodontium aciphyllum, Conostomum australe, Dicranoloma imponens, D. robustum, g. setosum, Dicranoweisia austrocrispula, Leptotheca gaudichaudi, Muelleriella crassifolia .Philonotis scabrifolia, Poly- trichadelphus robustus, Polytrichum strictum, Pseudodistrichium austrogeorgicum, Psilopilum antarcticum, Racomitrium stenocladium and Tortula anderssonii . The Hebe Facies.--Hebe elliptica, when occurring as scattered bushes, may be regarded as a relatively minor component of a dwarf shrub heath (Moore, l968). A heath of this type was visited in Waterfall Valley, Westpoint Island. The 53% shrubs occurred on steep sl0pes on the valley sides (see Photo l0), while through the valley bottom flowed a rather fast moving stream, with Gunnera magel- mforming a dense carpet along the watercourse. The sides of the valley were often cliff like, with numerous damp, shaded, shel- tered ledges and overhangs, where Leptoscyphus gpansus, LOphocolea sabuletorum, L. sylvatica, Marchantia bertercoana var. polylepida, fiajlavacinia x_iphoides, Riccardia gorgiensis, _R_. spectabilis, Symphyogyna hymenophyllum, Muelleriella crassifolia and Bartramia patens occurred, with Megaceros fuegiensis, Riccardia georgiensis, Maranea pseudozoopsis, and Webera albicans occurring under the water- fall or in the spray zone. The bryOphytes associated with the stream bed were typical of this habitat with Lophocolea austrigena, l_._. elata, Riccardia spectabilis, Telaranea pl umulosa, L Eeudozoqpsis and Momium conspissatum present. 26 Hebe elliptica also occurred on cliffs facing The Woolly Gut on Westpoint Island. The cliffs were rather moist, where numerous crevices, ledges, and overhangs were present. The bryophyte vege- tation included Clasmatocolea vermicularis, Hyalolepidozia bicuspidata, Leptoscyphus expanSUS. LOJhocolea lenta, L_. sabuletorum, I_._. semiteres, Marchantia berteroana var. polylepida, Metderia decipiens, fl. vio- lacea, Telaranea pseudozoopsis, Bartramia patens, Distichophyllum dicksoni, Eriow aLiculatus, Fissidens rigidulus, fiygroamblystegium fuegianum, Muelleriella crassifolia and Webera sp. gophocolea semi- teres, Megaceras fuegiensis, and Metzgeria violacea are the only Hepaticae or Anthocerotae found exclusively in this association. Summary of dwarf shrub heath Hepaticae.--The dwarf shrub heath associations have the highest number of hepatic species of any of the Falkland associations (75), and of these the following l7 are unique to it: Andrewsianthus pjanifolius, Cephalolobus scabrellus, Frullania boveana, fl. pseudolobulata, Hagialejeunea marginalis, hingennannia crassula, L_epicolea ochroleuca, Lepidozia laevifolia, geptoscyphus aequatus, Metzgeria leptoneura, fl. multiformis, 53910- mgayana, E. obovata, Riccardia alcicornis, _R_. prehensilis, S_chistochila pachyla and Telaranea blepharostoma. Of those that are not restricted to this association in the Falkland Islands, 49% also occur in the Cortaderia heaths, which is indicative of the rather close relationship between the associations. However, I regard the bryofloras of the Cortaderia association and dwarf shrub heath asso- ciations as sufficiently distinct, and the differences aid in dis- tinguishing the two associations. mm mg: of introduced grasses (commonly H____olcus l___anatus) (see Mics, 1939); (Evenin the most severely altered state of the oceanic heaths, I have found it possible to determine the natural vegetation type through examination of the bryoflora of stream banks. The stream banks are smat less disturbed and here the natural vegetation is least altered. See further comnents under Man Influenced Communities at the end of the ecology section. 3. FELDMRK FORMATION The feldmark formation has been characterized by the following four features: i. Considerable areas of bare, exposed, usually mineral soils with scattered small rocks or boulders. According to Greene (l964), feldmarks have a vegetation cover of one half or less the total area. ii. The tendency for the cushion habit to predominate. The feld- mark cushions do not form a continuous layer of peat (Mace, l955), and do not have among them the cushion-bog1 dominants of other associations (in the Falklands the Astglia associa- tion). Likewise, Azorella, the principal feldmark cushion _k 5.4 ‘l.‘ The term cushion bag is used to indicate cushion plant com- m peat; see Mace (1965) for a discussion. ii. _n. 28 ‘, *Mvotcur in cushion bogs (Wace, l960) . Hace 'fifiuifi) lists the principal species for feldmarks Minimally islands: '“mtand Islands--Azorella selago, Bolax gumifera ' nRerguelenquorella selago, Colobanthus kerguelensis, Ly. allia Eerguelensn Macquarie--Azorella selago, Racomitrium crispul um, Dicranowei ss 1' a antarcti ca Tristan da Cunha-- mpetrum rubrum, Racomitri um crispul um, _il. lanuginosum Gough Island--Racomitrium crispulum, Thysanomitrium richardii, Jamesoniella spp. The cushion habit is predominant in all of the above flowering plants. Bryophytes commonly assume either a cushion or com- pact habit. An often conspicuous cryptogamic component of the vegetation. 0n the Falkland Islands Neuropogon is a conspicuous element of the feldmark vegetation. Mosses are conspicuous in Mac- quarie Island feldmarks, and Ashton and Gill (1965) prepose that the feldmark comunity be termed the Azorella-Ditrichum alliance. . A preference for higher altitudes. Wace (l960) lists feldmark altitudinal ranges for the following islands: Falkland Is- lands (600-700 m.), Macquarie Island (200-450 m.), Kerguelen Islands (mo-300m.) and Tristan da Cunha and Gough Islands Race (1960) and "ace and Dickson (l965) (@4500 It.) . a F giantempart of the archipelago. Moore (1968, p. 16) masons "Apparently the formation is not solely a response to high altitude, because a comparable community type occurs along rock detritus along exposed cliff tops on the west coast just north of Cape Meredith . . ." (southernmost point of West Falkland Island). On Westpoint Island it is presumably expo- sure which results in a feldmark at the summit of Mt. Misery (335 m.). Feldmarks may also occur at rather low altitudes on Macquarie Island, where they occur above 183 m. (Ashton and Gill, 1965). The general aspect of Falkland feldmarks differs considerably. lihile there may be a continuum present, there appear to be two general types of feldmarks. i. The mesic feldmark. This type is characterized by a rather conspicuous fruticose lichen flora (principally Neuropogon). The lichen flora is apparently a result of the presence of mists and frequent enshrouding clouds. The feldmark community on the sumnit of Mt. Usborne 1 (ca. 700 m.) had a rather lush bryophyte vegetation due presumably to the numerous, rather small, scattered pools which were present. Associated solely with the pools, either submerged, at the edge of, or on rock News, ' - ‘a in.‘ m in. 2;? it 30 in the pools were Acrobolbus ochrophyllus, Adelanthus integer- rimus, A. lindenbergianus, Anastrophyllum ciliatum, Archeochaete kuehnemannii, (_Igphalozia badia, Clasmatocolea fulvella, Her- zogobryum vermiculare, tgphozia hatcheri, Pachyglossa fissa, and Plagiothecium laetum var. usbornense. Species not exclu- sively associated with the pools were Blepharidgahyllum densi- folium, Clasmatocolea koeppensis, Gackstroemia magellanica, Lepidozia chordulifera, i_._. fuegiensis, Leptoscyphus abditus, Leptoscyphus expansus, Pachyglossa dissitifolia, Plagiochila hirta, Riccardia granulata, B_. opuntiformis, Chorisodontium sp., Conostomum australe, Dicranoloma nigricaule, Polytrichum strictum and Racomitrium microcarpon f. papillosum. The feld- mark community on the summit of Mt. Maria (658 m.) was rather xeric, had a conspicuous lichen cover, but only a single bryo- phyte (Riccardia flintifomis). Nhile Adelanthus integerrimus is the only hepatic found exclusively in this formation, it is of interest to note that the following hepatics are found exclusively in the Mt. Us- borne feldmark and sheltered high altitude cliff vegetation: A_cg)bolbus ochrOphyllus, Anastrophyllum ciliatum, Archeochaete kuehnemannii, Herzogobryum vermiculare, Pachyglossa dissiti- folia and E. fissa. '. flexeric feldmark. This type is characterized by a compara- tively poor lichen cover (Neumpogon is absent). The moisture available to this feldmark type is derived solely from rain 31 and not from enshrouding clouds, and exposure is likely the factor which governs the comparative lack of moisture. Two feldmarks of this type were visited: the northwest tip of New Island Opposite Landsend Bluff (90 m.), and the north slope of Mt. Fegen (c. 275 m.). Very characteristic of the bryOphytes of this formation are the following three mosses, all of which grew on exposed rock surfaces, Andreaea pseudomutabilis, Racomitrium lanugin- mend g. rupestre. On soil in crevices under rock over- hangs are L_epidozia chordulifera and Ditrichum sp.; _Brflm argenteum may be found on B_o_l__a_x_ cushions. This feldmark type is quite xeric and exposed to the winds, and conse- quently supports only a very small bryophyte vegetation. Even the rock crevices, which often are relatively moist in the Falkland Islands, were dry and only L_epidozia chordulifera, which is able to grow in a wide range of habitats, was appar- ently able to tolerate the dryness. 4_.__ FEN AND BOG FORMATION This formation occurs where the water table is just below, at, or above the ground surface. a. ROSTKOVIA ASSOCIATION Moore (1968, p. 17) states, "This [association] is normally confined to the wettest depressions, where drainage is impeded, and 32 standing water is often present for long periods." In this associa- tionSphagmum falcatum grew submerged or nearly so, with _S_. fimbriatum also present. No hepatics were observed. b. ASTELIA ASSOCIATION Moore (19g. c_i_t_., p. 17) states, "This [association] is usu- ally found over deep peat and consists of a series of low flat cushions ora dense carpet composed of a few species which all are able to re- tain water in the dense covering of old leaves and branches surrounding their stems." These cushion bogs grade into the Rostkovia association and the wetter facies of the Cortaderia heath. The observed Astelia associations which did not grade into the above associations were dense, compact, and very hard, and no bryophytes were observed growing here. c. JUNCUS SCHEUZERIOIDES ASSOCIATION No bryophyte collections were made in this association. L BUSH FORMATION This formation is of restricted occurrence in the Falkland Islands, and only two native shrubs contribute to associations of this formation. a. CHILIOTRICHUM ASSOCIATION This association was not visited in the Falkland Islands. According to Moore (1968) and Skottsberg (1913) it is best developed 33 along the fringes of streams and rivers. Skottsberg (10c. cit.) lists two localities for this association, both in western Nest Falkland. b. HEBE ASSOCIATION Hebe elliptica, which is restricted to the West Falkland Is- lands, was Observed at Westpoint Island, but the plants were present only as scattered individuals or small thickets in a dwarf shrub heath (q. v.). When a dense stand of Hebe elliptica occurs, a true he_be_association is said to be present. Such an association is very rare in the Falkland Islands. A photograph of an example on Fox Is- land is in Skottsberg (l9l3, pl. 13). 6. LITTORAL VEGETATION The Senecio candicans and Ammophila-Elymus associations belong here, and both occur in sandy areas. The former association Was visited on the south shore of Cape Pembroke peninsula, near Surf Bay (East Falklands), and the latter at Gypsy Cove, Port William region (East Falklands). No bryophytes were found in either of these asso- ciations. It is likely that the dune and sandy areas are too unstable for bryOphyte survival . L_fliESH-NATER VEGETATION Moore (1968) includes the Eleocharis and Myriophyllum associa- tions in this category. No bryophytes were observed in either of these associations. The bryophytecollections which were frequently 34 gathered on the banks of shallow lagoons and ponds are included in the association of the surrounding area. 8. SHELTERED HIGH ALTITUDE CLIFF VEGETATION These communities consist principally of bryophytes, and occur above 6l0 m. in the Falklands (see Photo 12). They possess a highly characteristic assemblage of hepatics, and because of their restric- tion to high altitudes, I believe they deserve separate recognition. Cliffs on ridge tops have numerous protective niches such as ledges and rock crevices, the latter frequently beneath overhangs of varying sizes. The crevices, especially when beneath ledges, are commonly hoist, cool, and well shaded, a situation highly favorable to bryo- phyte growth. When this is accompanied by water seepage, the bryo- phytes are particularly luxurient. The variation in exposures, mois- ture, light, and the consistently rather cool temperatures, etc. , probably accounts for the considerable diversity of these communities. The following species occur in this community; those indicated by an asterisk are restricted to this community: Acrobolbus ochrophyllus, *flgplophozia fuegiana, Adelanthus lindenbergianus, A. tenuis, I_ndrgwsianthus australis, Anastrop_hyllum ciliatum, Archeochaete @nemannii, Austrolophozia fuegiensis, Bal antiopsis bisbifida, Mharidophyllum clandestinum, g. densifolium, Cephalozia badia, fligjplejeunea savatieriana, Cl asmatocolea fulvella, g. ggorgiensis, Q. mappensis, Cryptochila ggandiflora. 93.391591, *DJ'BIOthllum 35 acutilobum, Gackstroemia magellaniga *Herzogobryum erosum, *fi. teres, H. vermiculare, *iiygrobieljgasis iSOphyllum, Jamesoniella colorata, Jensenia pisicolor, L_epidozia chordulifera, I_._. fuegiensis, Leptoscyphus abditus, L_. mansus, Marsupidium urvilleanum, Metzgeria falklandica, Pachyglossa dissitifolia, _P_. fissa, P. sgegazzininana var. exilis, *Plagiochila atroviridis, _P_. elata, P. hirta, *Radula helix, 3. papil- l_o_s_g,3. saxicola, 3. tenax, Roivainenia jacquinotii, Schistochila carnosa, S. subintegerrima, Symphyogyna Mengphylj um, Telaranea flunulosa, TriandrOphyllum subtrifidum, Andreaea a_lpina, Bartramia hptophylla, Dendroligotrichum sguamosum, Dicranoloma harioti, Philon- msscabrifolia, Polytrichadelphus robustus, and Polytrichum strictum. The following combination of hepatics was very frequently encountered and were often intermixed: AustrOTOphozia fuegiensis, Balantiopsis bribifida, Q‘yptochila spp., Herzogobryum teres and Pachyglossa spp. In these sheltered high altitude cliffs there are an amazingly high percentage of Hepaticae which either (a) have a subantarctic distribution, (b) also occur in New Zealand, or (c) have very close relatives in New Zealand or Tasmania. The hepatic flora of this asso- ciation appears to be in large part an old, relic one. A large number of taxa (especially the genera Acrolpphozia, Austrolophozia, Her:z_o_— gobryum and Pachyglossa) may represent remnants of an early, wide- Spread flora involving the Antarctic continent. It is possible that the high altitude cliffs served as nunatak areas. This is in marked contrast to the lowland associations where the hepatics are charac- teristic of a flora that has recently (i .e.. post Pleistocene) invaded 36 the Falkland Islands. Exemplifying this is the complete absence of the genus Lophocolea fromthe sheltered high altitude cliffs. This genus is not characteristic of austral relic floras, but as many members of the genus are highly plastic (often strikingly so), and tend to be weedy, the taxa of the genus are more characteristic of areas that have been recently invaded. 9. NAN INFLUENCED COMMUNITIES a. PASTURELANO In the Falkland Islands there are wide expanses of pasture- land (see Davies, 1939) which are regularly grazed by sheep. The bryophytes occurring here are Bryum (sect. Rosulata) sp., Campylopus introflexus, Ceratodon purpureus and Polytrichum piliferum. It is of interest to note that all of these mosses are quite widespread and weedy, and it is not surprising that these three rather plastic species were able to tolerate highly disturbed situations. Near the mouth of Cheeks' Creek, Fox Bay region, a collection was made in a stream cut through a pasture. It is probable that before conversion to pastureland the vegetation of the area was that Of a dwarf shrub heath association, as the bryoflora was similar to the stream in that association, with the exception of the presence here of Lophocol ea irregul ari s. b. GROVES 0F PLANTED TREES At the Hill Cove settlement, a mature grove exists, which, according to Dallimore (l9l9) was introduced in the period 1889-1894. 37 The hardwood grove, which is made up of Nothofagus sp. and Populus sp., has a rather dense canopy with little woody undergrowth (see Photos l3-l4). The hepatics found here were gptoscyphus expansus, Loghocolea Tenn, L. leptantha and Marchantia berteroana var. polylepida; all occur in a variety of natural communities. Adjacent to the hardwood grove was a stand of conifers. No bryophytes were observed in this stand, or the coniferous stand on Weddell Island. C. GORSE THICKETS Gorse (Ulex eurgpaea), a native of western Europe, is widely utilized in the Falkland Islands as sheep fencing. A gorse thicket was visited at Boca House near Brenton Lock, near Darwin Settlement, East Falklands. 0n soil beneath the gorse occurred Leptoscyphus gxpansus, Brachythecium austrosalebrosum, Bryum argenteum, _S_. (sect. Rosulata) sp. and Tortula robusta. Tortula monoica was the only bryophyte actually growing on the gorse and it occurred on the ex- posed roots. IV . PHYTOGEOGRAPHY A. Austral Regions Greene (1964) notes that the greatest barrier against arrival atasatisfactory botanical criterion for recognizing and subdividing the various zones in the austral regions is the critical lack of know- ledge about the plants of these lands. In view of this, most recent attempts at classification schemes are primarily vegetational and geographical . Wace (l960) devised a system based primarily on lowland vege- tation criteria. He combined all those islands bearing oceanic heath mm or flowering plant cushion bogs into a temperate category, and further divided this group into warm and cool temperate respec- tively according to the presence or absence of fern-bush communities. He limited usage of the subantarctic to include only those islands with a closed herbfield vegetation and in which Sphagnum and cushion bogs are absent. The islands in which there are no flowering plants (or only scattered individuals), be considered as Antarctic.1 Wace compiled the available temperature data and noted that the groupings based on lowland vegetation types correlated well with groupings M ————_. 1It must be understOod that islands, like Bounty Island off New Zealand, may be without vascular plants as a result of physio- graphic and biotic factors. 38 39 based on mean annual temperature. The following is his classification unmn SUBTROPICAL TEMPERATE SUBANTARCTIC ANTARCTIC Kermadec Islands Juan Fernandez Islands Chatham Island Tristan da Cunha. Inaccessible, Nightingale and Gough Islands New Amsterdam and St. Paul Islands New Zealand shelf islands Falkland Islands Marion and Prince Edward Islands Macquarie Island Crozet Islands Kerguelen and off-lying islands South Georgia Heard and Macdonald Islands Bouvet Island South Sandwich, South Orkney and South Shetland Islands Palmer Archipelago (Graham Land) Greene and Greene (1963) and Greene (1954) adopted Wace's (196 0) co”C‘iPt of the term subantarctic, but included Marion and Pfic . e Edward 15131195. and used the term Antarctic region to enbrace He ° Antarctic and subantarctic zones (see Map 53)° 40 Wace (1965, p. 239) extended his earlier scheme, stating "In order to arrive at a vegetational classification, the major features of the structure of the plant community should be used exclusively to delimit the different zones." He then proposed to use the limit of tree or woody shrub growth to separate the temperate from the sub- antarctic and the limit of closed phanerogamic communities to separ- ate the subantarctic from the antarctic vegetation. He characterized the subantarctic vegetation zone as having terrestrial plant communi- ties with the following four features: 1. Soligenous mires in which the important peat-forming plants are Bryales and Juncaceae (not Sphagna). 2. Feldmark communities composed of flowering plants with very close-growing and compact mat and cushion forms. 3. Closed herbfield communities in which large-leaved perennial herbs are conspicuous. 4. Communities of large trunk-forming tussock grasses around the coasts. H ace excluded all continental areas and restricted the usage 0f the te rmsubantarctic to remote islands. This zone included South Ge ' "9““ Marion, Prince Edward, Crozet, Kerguelen. Heard 3"“ Mac- quarie Islands. It is evident from the criteria outlined in Wace (I950. 1955) and Greene (1964) that the Falkland Islands are south temPf-‘Y‘ate ”the" 41 than subantarctic. The characters which place them in the south tem- perate region are the following: a) a widespread oceanic heath forma- tion, b) the presence of flowering plant cushion bogs, c) the presence ofgphggnuflbogs, d) the presence of woody shrub growth and e) the ab- sence of a closed herbfield vegetation. The presence of a feldmark comunity has been used as a characteristic of the subantarctic re- gion. In the Falkland Islands, this community is present, but cer- tainly does not reach the significant proportions and role that it does in the Kerguelen Islands. Skottsberg (1960) based his austral phytogeographic analysis on the following floristic regions. Five further regions are included, some including "outposts“ for "antarctic types." I. The antarctic zone, south of 60° 5., but including the South Sandwich Islands and Bouvet Island. 11. The subantarctic zone, between approximately 48° S. and 60° 5.. but including Marion, Prince Edward and Crozet Islands. A. Magellanian province. l. West Patagonian-Fuegian district. 2. Andean Patagonian-Fuegian district. 3. Falkland Islands and South Georgia district. 8' Kerguelen province. Marion, Prince Edward. Crozet, Ker- guelen and Heard Islands. C. Province of the subantarctic islands of New Zealand. no 42 III. The Austral zone, between approximately 40° S. and 48° S. A conspicuous antarctic floristic element is present, often forming distinct communities in a flora and vegetation of a different origin. A. Valdivian province. B. Province of the South Island of New Zealand, Chatham Islands and the Tasmanian tableland. Godley (1960) adopted the subantarctic zone of Skottsberg, but remarked (p. 472), "The presence or absence of Sphagnum bogs appears to me to be one of the significant features to be taken into consideration when delimiting a homogeneous subantarctic region." He regarded peat formation as a common feature of the region, and the bogs formed by cushion plants rather than by Sphagna. The Antarctic region includes regions south of the subantarc- tic and is characterized by a vegetation which ". . . is at best a poorly developed tundra" (Rudolph, 1971, p. 192). Soligenous mires. herbfields, tussock forming grasses and cushion plants are absent. and the vegetation is a tundra type dominated by mosses and lichens. Two flowering plants, Deschampsia antarctica and Colobanthgs grjssi- M. occur in Antarctica. 43 B. The Phytogeogrgihic Categories There are a large number of monotypic genera and stenotypic1 species which possess a range centering about the cool, moist south temperate and subantarctic regions. There is also a high proportion of taxa which Schuster (1969c, p. 82) states "can be interpreted as 'relict' groups, rather than as relatively new and as yet undiversi- fied groups." The chief difficulty in the assessment of the phyto- geographical relationships of the Falkland Island Hepaticae and An- thocerotae is the delimitation of the south temperate and subantarc- tic patterns of distribution. I have delimited these patterns after the concept of the zones or regions developed by Skottsberg, Godley, Greene and Wace. Ihave recognized 12 categories of distribution patterns of Falkland Island Hepaticae. The Anthocerotae fall within only one of these categories. The data for these distributions was gathered from Specimens personally examined, and reports extracted from the litera- tu"e- If the report is regarded as questionable, it 1'5 not lnClUded here. To my knowledge, references to Macquarie Island hepatics are restricted to the following: Ashton and Gill (1955). Gillham “967% G”11620966. 1957), and Hodgson (1953, 1961, 1962). The paucity of Macquarie Island reports undoubtedly affects the data in the amphi- pacific, Pantemperate and subantarctic groups. The only recorded he W1C Species shared by the Falkland and Macquarie Islands are Jame wm and gpidozia laevifolia (fide Hodgson in Ashton and Gill 1965). \ l . Taxa with very narrow habitat requirements. 44 CONSPECTUS 0F PHYTOGEOGRAPHICAL CATEGORIES Category Number of Percent of Species Total Flora Endemic 3 2.3 Fuegian - Falkland 10 7.8 Magellanian - Falkland 19 14.9 :3 on: US.— ‘g‘g Valdivian - Falkland 7 5.5 so an- Magellanian + Valdivian - 46 36.2 Falkland Andean - Falkland 2 1.5 Amphipacific temperate 12 9.4 \ .=:: 533____ Amphiatlantic temperate 10 7.8 1 SEE? E Pantemperate 5 3.9 \ Subantarctic 10 7.8 ~\ Antarctic 1 .7 \ Widespread 2 1.5 *1\ i . . I .e I .. a L . . r .0. . u pp. 9 . .- I o u a o. ..u .. . . . . a u .a. u 5. . . . .1. . .n. a a a I. a . s v . s a u I v .I. all nae. . . . . . .. ca. 3. . . v . - C- o. ... . .a. .. .. '5‘ i v in \Is \0. 1‘. \‘u\ s. \ ‘l! \ . \r .1. 45 l. TEMPERATE Species occurring within the south temperate regions of the world. a. AMERICAN TEMPERATE Species occurring within the south temperate regions of the American sector. There are three species which occur in southern South America as well as South Georgia (see S-f). Their distribution is regarded as temperate rather than subantarctic, as northward exten- sion from the subantarctic is not confined to higher altitudes. It has frequently been remarked that the Falkland vascular plant flora is an extension of the southern South American flora (Hooker, 1847; Skottsberg, 1909, 1913; Moore, 1968), a statement to be expected with regard to a continental shelf island. Moore states that of the 163 native species of vascular plants of the Falkland Islands, 89% occur on the mainland south of 40° S. All except three of the remainder are endemic to the Falkland Isl ands and are in large part related to mainland taxa. Like the vascular flora, the mainland hepatic representatives extend northward to varying degrees. Commenc- ing with the endemic category, the remainder of groups are arranged principally according to the degree of northward extension. Djidemic to Falkland Isl ands Andrewsi anthus pl ani fol i us Metzgeria falklandica Riccardia regularis 46 2) Fuegian-Fal kl and . Taxa occurring in the Falkland Islands and Tierra del Fuego. Acrolophozia fuegiana Lophocolea elata Adelanthus integerrimus Riccardia muntiformis Archeochaete kuehnemannii Riccardi a papillosa Balantiopsis bisbifida Riccardia saxicola Harpalejeunea marginalis Schistochila subintegerri ma Q Magellanian-Falkland. Taxa occuring in southern Patagonia north to 48° 3. or Fuegia and southern Patagonia north to 48° 3. The northern boundary was affixed by Skottsberg (1916) to delimit the Magellanian and Valdivian regions, and has been widely followed by various authors. Of the 19 taxa in this group, 74% have been found to occur in the magellanian moorland. Their occurrence in the moorland is indicated by an aster- isk, and those taxa restricted to the moorland are indicated by a double asterisk. This region occurs south of 48° 5. to Cabe de Hornos, and is limited by the magellanian rain forest to the east and the Pacific Ocean to the west (see discussion and map in Godley, 1960). The moorland is extremely wet, exposed to violent winds and has a permanently saturated peaty soil. 0f the remaining four taxa, Melanthus tenuis and Hygrobiellopsis isophyllum are known from deciduous Nothofagus forests, and Anastropmfllum ciliatum and Egg: Mpseudolobulata from evergreen Nothofagnas forests. ._.I g L.' '9- ‘. I n, ‘0 T a a ‘a , \,' . \T - . . ‘I v n. ’v . . l . e . ‘- I U ‘. D ‘v. ‘s 1‘ I \e. 47 Adelanthus tenui 5 *Mi crol epjdozi a seti formi s Anastrophyllum ciliatum *Neolepidozia ol igOphyl la *Andrewsianthus australis (”Plagiochila atroviridus *Cephalolobus scabrellus *Plagiochila obovata *Chiloscyphus hookeri *Riccardia manulata Frullania pseudolobulata *Riccardia spectabilis *Gackstroemia patagonica *Riccardia tenax Hygrobiellopsis isophyllum **Saccogynidium vasculosum *Leptoscyphus aequatus *Schistochi la [Echyl a **Microlefldozi a mol l is 4) Valdi vian-Fal k1 and. Occuring in Chile and/or Andean Patagonia (see group 5 for definition) north of 48° S. and south of 36° S. , or occasionally ab- sent from mainland and only on Juan Fernandez. Skottsberg (1916, p. 13) places the northern boundary of the Valdivian region at 40° S. , while Kuschel (1960) states that the zone occurs from 36° S. in the Andes,~37° S. in the central valley. As stated above, the southern boundary of the region has been placed at 48° S. This forest region is characterized by possessing a "species rich" vegetation, dominated by Eucryphia cordifolia, Laurelia serrata, Weinmannia trichosperma and Anomyrtus spp. (Holdgate, 1961). a) The following species are known to occur on the mainland and in all cases, except Lejeunea corralensis, on Juan Fernandez: 48 Jungermannia crassula Plagiochila gayana Lejeunea corralensis Wettsteinia densiretis Lophocolea sylvatica b) The following are known to occur on Juan Fernandez, but have not been reported from the mainland: Metzgeri a mul ti formi s Symphyogyna hochstetteri S) Valdivian + Magellanian - Falkland. Species which are essentially widespread in the American temperate zone. Of the 46 taxa in this group, 72% are known to occur in the magellanian moorland. Their occurrence in the moor- land is indicated by an asterisk. It should be repeated here that the moorland occurs south of 48° S. and thus is restricted to the Magellanian floristic region. With regard to the magellanian region, of the remaining 28%, Cheilolejeunea savatieriana, Clasmatocolea figppensis, inlophyllum acutilobum, Lophocolea irregularis, Mega- §g_r_g§_fuegiensis, Paracromastiggm subsimplex and Riccardia fusco- Pfllmoccur in the deciduous Nothofags zone, PlaLiochila ansata and g. m in the evergreen Nothofagus region and Metzgeria vio- lggggand Teleranea pseudozoopsis in the deciduous and evergreen Nothofagus regions. The taxa, with the exception of group f), are arranged accord- ing to their latitudinal range. 49 a) South from 45° 5.: Cheilolejeunea savatieriana *Lophocolea divaricata *Lepicolea rigida *Pigafettoa crenulata b) South from 43° 30' S. (line from north side of I. Guafo): *Blepharidophyllum gottscheanum *Microlepidozia saddlensis *Clasmatocolea cookiana Plagiochila ansata *Leptoscyphus mtagonicus Plagiochila hirta *Lcmhocolea leptantha *Riccardia pallidevirens *Lophocolea textilis *Teleranea plumulosa c) South from 40° 5.: D_lPlophyllum acutilobum *Riccardia alcicornis Maceros fuegiensis Telaranea pseudozoopsis *Plagiochila obcuneata dl South from 36° 3.; from latitude indicated: *Mastrepta blf'ida (39° 52' S.) *Bilantiopsis cancellata (39° 48' S.) *Bibntiopsis erinacea (36° 50' S.) *C1_asmatocolea fulvella (36° 50' S.) *CJXPtOChiLa paludosa (39° 52' s.) *Mm'ebma (39° 53' s.) *Mstroemia m_agellanica (39° 52' s.) *flfihocolgg austrigena (39° 52' S.) 50 *Marsupidium urvilleanum (39° 38' S.) Metzgeria violacea (39° 46' S.) *Plagiochila elata (39° 36' S.) *Radula helix (39° 38' S.) *Schistochila laminigera (39° 38' S.) *Telaranea blepharostoma (39° 48' S.) e) Species about which insufficient knowledge is known concerning range: *Frullania magellanica Lophocolea sabuletorum *gpjdozia chordulifera *Metahygrobiella tubulata *gpidozia fuegiensis Paracromastigum subsimplex Lethocolea radicosa Riccardia fuscobrunnea thCOlea irregularis *Riccardia prehensilis 1‘) Valdivian + Magellanian--Falkland Islands--South Georgia The species listed here are considered as south temperate rather than subantarctic, as northward extension from the subantarctic is not confined to higher altitudes. *Ephaloziena dusenii *Roivainenia jacquinotii Qgsmatocol ea koeppensi s SkOAT-Shem (1910, 1916) and Moore (1968) recognize three V999“ W0” zones in the region south of 40° S. 1. West Patagonia. The region including the west slope of the Andes to the Pacific Ocean, characterized by hi9h ii. 1.11. I 51 precipitation and supportive of lush rain forests and magel- lanian moorland. Andean Patagonia. The eastern slope of the Andes from base to snowline. This region experiences moderate rainfall and supports a deciduous, comparatively dry forest. Hueck (1966) recognizes two forested regions which extend to Andean Pata- gonia, the Northern Nothofagus forests which are composed predominantly of two species of deciduous Nothofagus, N. Wand N. procera, and the Araucaria - Libocedrus zone. East Patagonia. The steppe region, ranging east from the Andean foothills, characterized by grassland and xerophytic shrubs. As is to be expected, the vast majority of Hepaticae in groups 3, - . 4. and 5 occur predominantly 1n the West Patagonian zone. The fol ' ' ' ' low1ng Vald1v1an-Magellan1an Hepaticae and Anthocerotae occur with' . 1n the Andean Patagonia zone, and nearly all in the Lago Nahuel Hua ‘ ' Pl reg1on. None are restricted here. QLasmatocolea fulvella (31_asmatocolea humilis F_rullania magellanica 93m pisicolor L_epidozia chordul i fera Lophocol ea texti 1 is Marsupidium urvilleangfl Megaceros 1115916051.. Metzgeria vi olacea Plagiochilg elata Riccardia prehensilis Telaranea pseudozoopsis 52 L_eptoscyphus expansus, Megaceros fuegiensis, and Symphyogyna hymenophyllum are the only three hepatics occurring in east _f Patagonia. 6) Andean-Falkland. Species extending north of 36° S. in the Andes. 0f the American temperate species, there are two taxa which have utilized the Andes as a route for northward migration to lower latitudes. It is evident that migration has been northward as both species are rather common in southern South America, and occur in the Andes only at higher altitudes. Isotachis humectata Noterocl ada confl uens b. AMPHIPACIFIC TEMPERATE Distribution mainly in temperate parts of the South Pacific in the southern hemisphere, i.e., temperate South America (occasion- ally Juan Fernandez), New Zealand, Tasmania and southeast Australia. The relatively large number of taxa (12) in this group is illustra- tive of the rather close relationship between the New Zealand and Southern South American hepatic floras. There are varying degrees 01 Penetration northward into the New Zealand sector, and the taxa have been arranged accordingly. One asterisk indicates occurrence 0" New Zealand shelf islands, two indicates occurrence in New Zea- 1arid and three indicates occurrence on New Zealand shelf islands and New Zealand. Only two taxa also occur in the Northern Hemi- s . . . phere, these are Metzgeria decipiens and Triandrophylhafll ilfl’tr‘f‘dum' 53 1) South Island: Austrolqfiiozia ftggiensis 2) To North Island: *Chiloscyphus triacanthus Pallavacinia xiphoides Saccogynidium australe 3) To Tasmania: **Herzogobryum teres ***Lepidozi a laevi fol i a 4) To Australia: ***LoLhocolea len ta 5) To Japan: **Metzgeria decipi ens c. AMPHIATLANTIC TEMPERATE Symphyogyna hymenophyllum *Telaranea tetradactyla *Temnoma quadripartita *Triandrophyllum subtrifidum Distribution mainly in temperate parts of the south Atlantic, 1'.e., temperate South America, South Africa (and occasionally Tristan da Cunha). Species whose sole occurrence in the Indian Ocean sector is on subantarctic islands are included here and indicated by an asterisk. They are not considered as subantarctic species, as north- ward extensions from the subantarctic are not restricted to higher altitudes. 54 Adelanthus lindenbergianus Hyalolepidozia bicuSpidata *Blepharidophyllum clandestinum *Jensenia pisicolor *BlepharidophLllum densifolium Lepicolea ochroleuca *Clasmatocolea humilis Leptoscyghus expansus Clasmatocolea vermi culari s Pseudocepha l ozia tri staniana All of these taxa, with the exception of Hyalolepidozia bicuspidata,1 penetrate north via the Andes to lower latitudes (Adelanthus linden- bergianus, Clasmatocolea vermicularis, and Lepicolea ochroleuca reaching into the Northern Hemisphere), and are known from relatively high altitudes north of 36° 5. This is an ensemble of rather plas- tic, polystenic species which are able to adapt to a variety of eco- logical niches. Legtosgyphus expansus is particularly xeric tolerant as it is one of the two Falkland taxa known from the east Patagonia zone. Among them, Lepicolea ochroleuca perhaps has the narrowest ecological requirements, especially moisture requirements. d. PAN-TEMPERATE Species occurring in temperate regions of South America, Australia, and South Africa. Acrobol bus ochrophyl 1 us Lophocol ea semi teres Cryptochila gandi flora Marchantia berteroana Jamesoniel la colorata lhyalolepidozia bicugaidata has been reported from "Frai Jorge," Prov. Coquinhio, ChTTe. an. I, 55 2. SUBANTARCTIC Species occurring on one or more subantarctic islands (as defined by Greene, 1964) of at least one sector (e.g. , American, Indian Ocean, or New Zealand) with northward extensions only at higher altitudes. If one includes cool, south temperate taxa in this group the number of species becomes drastically inflated, and the south temperate and subantarctic elements merge and become confused. Several authors have not recognized the south temperate and subant- arctic regions (as defined here) and have included the patterns under the single category of "antipodal" (Schuster 1963a, 1969c, etc.), ”subantarctic" (Grolle, 1969, and others) or "antarctic" (Fulford, 1963b, 1966). There are only ten subantarctic species in the Falk- land flora. None of the Species occurs on Juan Fernandez, or pene- trates northward beyond the Valdivian region. The sectors in which the taxa occur are indicated after each species. Cephaloz‘ia badia (A) Pachyglossa dissitifolia (A) Clasmatocolea gemiensis (A) Pachyglossa fissa (I) Herzogobgyum erosum (A) Pachyglossa spegazziniana (A) Herzogobryum vermiculare (A,I) Riccardia georgiensis (A) Leptoscyphus abditus (I) Schistochila carnosa (A,I) LANTARCTIC Species occurring in the Antarctic Zone (as defined by Greene, ‘9641. with northward extensions into the subantarctic or temperate .tn '0‘ c \ ..........\ 56 zones only at higher altitudes. The only species which may be in- cluded here is Lgahozia hatcheri, which is restricted in the Falk- lands to the higher elevations. Unfortunately its precide distribu- tion as regards altitude in South Georgia is unknown. The worldwide distribution of L_. hatcheri can be termed bipolar. 4. NIDESPREAD Species not in any of the above categories and not pan- tropical. Aneura pi nguis Metzgeria legioneura C. Distribution within the Falklands The taxa below are arranged according to their occurrence in either the East Falklands or the West Falklands. Taxa known only from East Falklands; of these, 39% are known only from the Mt. Usborne region, and these are indicated by an as- terisk. *Acrolophozia fuegiana *Microlepidozia saddlensis *Anastrepta bifida *Microlepidozia setiformis Andrewsianthus planifolius *Neolepidozia 01 igophylla Blepharidophyllum gottscheanum Pachyglossa fissa Cephalolobus scabrellus *Paracromastigum subsimplex 57 Clasmatocolea cookiana *Gackstroemi a patagoni ca Junjermannia crassula Lejeunea corralensis Lepicolea ochroleuca Lepicolea rigida Leptoscyphus aequatus Mahocolea divaricata Lophocolea textilis Metzgeria multiformis *Microlepidozia mol l is Plagichila obcuneata Riccardia alcicornis *Riccardia granulata Riccardia prehensilis *Riccardia saxicola *Saccogynidium australe Schistochila laminigera Schistochila pachyla Symphyogyna hochstetteri Telaranea blepharostoma Telaranea tetradactyla Taxa known only from West Falklands: Diplophyllum acutilobum Herzogobgyum erosum Lethocolea radicosa Lophocolea irregul ari s tgphocolea semiteres Megaceros fuegiensis Metzgeria violacea Pigafettoa crenulata Plagiochila atroviridis Plagiochila obovata Radula helix Riccardia regularis Schistochila carnosa While I regard the highly local occurrence of many Falkland Island taxa to be a reflection of the availability of microhabitats and narrow ecological requirements, the comparison of hepatics re- stricted to the East Falklands (29) compared to the West Falklands (”lis hmnessive, and may well reflect differing climatic conditions 58 within each of the respective island groups. Alternatively, the dif- ference in species diversity between East and West Falklands may be due to critical local levels of temperature and rainfall which are present in various parts of the Falkland Islands (see Map 61 with rainfall indicated, compare especially San Carlos with Port San Carlos). Port Stanley is one of the wettest areas in the entire archipelago, and this feature coupled with the cool temperatures known for the region may account for the ten taxa which were found to occur only in the Port Stanley-Port William regions. These species, which fall within a variety of elements, were: Andrewsianthus planifolius Metzgeria multiformis Qphalolobus scabrellus Riccardia prehensilis Jungermannia crassula Schistochila pachyla L_epicolea ochroleuca Telaranea blepharostoma LOW divaricata Telaranea tetradactyla The northwesternmost portion of the Falkland Islands has higher temperatures and within this region are confined or nearly so. thl'ee vascular plants (Blechnum chilense, Gavilea macroptera, and WM), of the Valdivian element. Lophocolea semiteres ‘5 known in the Falkland Islands only from Westpoint Island. The SIii-Cites seems to be restricted to rather warm temperature regionS. mam”)! in the American sector (i.e. , the species is unknown for Ill e Ma9ellanian region), where it may be considered Valdivian. !3r..., 59 Of the taxa occurring on both East and West Falklands, three were confined to the wettest known stations on record. Frullania pseudolobulata, and Metzgeria falklandicg were collected at Port Stanley and Port Howard. Jensenia pisicolor is known from Port Stanley, Port Howard, and Mt. Adam. Rainfall figures for the Mt. Usborne region would be of great interest. A large number of the Hepaticae of this region are also known only from the wettest known stations on record, as the lists below illustrate. It is likely that the area receives as much rain- fall as that of Port Stanley, Port Howard, or Mt. Adam. Hepaticae known only from the regions of Mt. Usborne, Port Howard, and Mt. Adam: Anastrophyllum ciliatum Herzogobryum teres Blepharidophyllum gottscheanum Herzogobryum vermiculare Clasmatocolea cookiana Hygrobiellopsis isophyllum Motochila grandiflora Pachyglossa fissa ijptochila paludosa Hepaticae known only from the regions of Port Stanley, Mt. USPPI‘PEi Port Howard. and Mt. Adam: fltrolophozia fuegiensis Pachyglossa dissitifolia CI_Elsmatocolea fulvella Pachyglossa spegazziniana L . var. exilis . 3PM fuegiensis Schistochi Ia subintegerrima "' 1:? .. .. . h m . . .. In ,.. ." -- . .- ‘O‘ V.‘ y. n a, . .0. I I 60 Hepaticae known only from the regions of Mt. Osborne and Port Howard: Adel anthus integerrimus Cl asmatoeol ea' georgiensi s Cephalozia badia Riccardia fuscobrunnea Hepaticae known only from the regions of Mt. Usborne and Mt. Adam: Acrobolbus ochrophyllus Riccardia pallidevirens Archeochaete kuehnemannii Hepaticae known only from the regions of Mt. Usborne and Port Stanley: L—QQWSCYPPUS aeguatus Symphyogyna hochstetteri Hepaticae known only from the regions of Port Stanley, Mt. Usborne and Mt. Adam: flephari dophyl 1 um densi fol i um V . SYSTEMATIC ACCOUNT A. Introduction l. FORMAT The sequence of orders and families follows the system of classification outlined in Schuster (1966b). The austral genera not placed in families in Schuster (loc. cit.) are placed where appropriate. The genera are arranged in alphabetical order within families, and species in alphabetical order within genera. 2. NOMENCLATURE AND LITERATURE CITATIONS a. AUTHOR CITATIONS Author citations follow the abbreviations in Sayre et al. (l964). 6. JOURNAL CITATIONS Journal citations follow the abbreviations in the World List 9f_§cientific Periodicals (Brown and Stratton, 1963-1965). The titles of journals not included in the World List have been abbre- viated in a similar manner. 61 62 c. SYNONYMY Wherever possible, I have attempted to provide a full synonymy for each taxon. In the citation of synonyms, I have used cf. and fide in the sense used in Index Muscorum (Wijk et al., 1959-1969), i.e., the former as "confer, compare, a reference to an implicit statement by an author concerning a question of taxonomic synonymy," and the latter as "according to, a reference to an explicit statement by an author concerning a question of taxonomic synonymy." d. TYPIFICATIONS I have attempted to indicate the status of typification of all taxa. The vast majority of taxa from the Valdivian and Magellanian regions as well as the Falkland Islands were originally described by Hooker, Taylor, or Stephani. These individuals worked in a period prior to the establishment of the current type method, and the terms holotype and isotype should not be applied to the original material on which their taxa were based. I have proposed a lectotype only after Ihave examined a sufficient number of specimens of the original material to state that a single specimen best represents the describ- ing author's concept of the species. Otherwise, the expression original material is used. I have accredited lectotypification to an author if a lectotype is implied by the citation, i.e., in several instances the term type is followed by citation of an i sotype. With regard to lectotypification, I have used the term cf. to refer to an M statement made by an author and the term fide is used in reference to an explicit statement made by an author. If the author 63 used the term type and no clear identification of a particular col- lection is furnished, I have used the expression original material. If an author has merely said type when describing a species after 1952,1 I have used the term holotype if the location and identity of the type is clearly given. If many localities are listed in the original protologue, as often is the case with species described by Stephani. there is an obvious requirement for lectotypification. In instances where only asingle locality is given, there is still no assurance that a single collection or element is involved. Since several collections or elements may be involved, either in one herbarium or distributed in two or more herbaria, I have referred to the specimen(s) as "original material." This is especially pertinent with collections of Skotts- berg (and/or Halle) and Hooker, since I have frequently found more definitive and better representative specimens in an herbarium other than that of the describing author, and I have preferred to designate these as the lectotypes° e. HERBARIUM CITATIONS These follow the abbreviations in Lanjouw and Stafleu (1964). 3. ECOLOGY I have provided ecological notes for each species I have collected in the Falklands. The terminology and concepts used in the M 'The term holotype, lectotype, syntype and 060139439. etc., were adopted at the Seventh International Botanical Congress in Stockholm, 1950 (see Lanjouw 1950 and Troupin, 1949) and are used in the Inter- nat1onal Code for the first time in 1952 (see Lanjouw, 1952). ‘ . 0".- v a. .. . . -14 a. a 1 u ._ e “ . ~.... ‘ . o. u a. . ‘ . ‘. i ‘ \o 'v 0". u x,“ 1‘ . ,‘ . c 1‘ ' \ c 'o . 0 U .. N ’. o “. c -. a ‘1 ‘v Q . 64 ecological discussions have been defined in the ecology section (see section III, B). 4. DISTRIBUTION The distribution. The distribution of each species has been summarized, and the terminology and concepts used have been defined in the phytogeography section (see sections IV A, B). The expression Patagonian Channels, however, has not been defined elsewhere, and I have used it to indicate the highly dissected portion of Chile south of Puerto Montt to and including the Brunswick Peninsula (i.e., lati- tudes 41° 28' S. to 53° 30' 5.). It includes the western portion of the mainland, and the channels near Puerto Natales, S. Skyring, S. Otway, etc. 5. LITERATURE RECORDS An attempt has been made to compile the literature records for each species. References in which taxa were originally described are not repeated in the literature record section. The information is pre- sented as it is given in the original source, and no attempt has been made to edit the extractions. For example, there are several in- stances where a collector is reported for a locality he did not visit. The localities are cited in the manner provided in gazetteers of the various regions, and it is only to this extent that I have edited the extractions. The following gazetteers were utilized: Chile (U.S. Office of Geography: Chile...1967), Argentina (American Geographical Society of New York...Argentina...1944), and New Zealand (New Zea- land. Dept. of Lands and Survey, 1968). Abbreviations of localities follow the American Geographical Society indices to maps of Hispanic America (see American Geographical Society of New York...Argentina. .. T944) and are as follows: Arch. - Archipelago A. - Arroyo B. - Bahia Br. - Brazo Cta. - Caleta C. - Cerro Cord. - Cordillera Ens. - Ensenada Esto. - Estuario P - Fiordo I. - Isla L. - Lago La. - Laguna 6. SPECIMENS SEEN M M. - Monte Mt. - Mount, Mountain Pen. - Peninsula Pta. - Punta Port. - Portezuelo Pto. - Puerto Q. - Quebrada Ran. - Rancho R. - Rio S. - Seno Sa. - Serra, Sierra Vo. - Ventisquero V. - Volcan I have grouped all Falkland collections together, follOwed by additional collections seen. The specimens I have personally col- lected are indicated by number only, without citation of collector. 66 Acomplete set of my collections has been deposited in the Crypto- gamic Herbarium, Michigan State University (MSC). All other spe- cimens are cited with the collector's name provided. B. Key to the Classes and Orders of ngaticae and AnthocerotaeI l. GametOphytic cells (at least superficial cells) each nor- mally with l (-2-several) chloroplasts; cells lacking oil bodies; gametophyte prostrate, thallose, never with leaves, never with air chambers, with stomata on ventral surface; sporOphyte linear, 2 valved, with a columella; archegonia embedded and antheridia endogenous. . . . .Class ANTHOCEROTAE, Order ANTHOCEROTALES l. GametOphytic chlorOphyllose cells with numerous chloro- plasts; chlorOphyllose cells usually all (or many) with oil bodies; gametophyte thallose or leafy, without sto- mata; sporOphyte a non-linear capsule, usually 4 valved, without a columella, without stomata; sex organs exo- genous, not embedded ........... -. 2. . .Class HEPATICAE 2. Rhizoids typically dimorphic, some tuberculate, others smooth; plants clearly thalloid, without leaf-likelobes, the thallus firm, fleshy, Opaque, with ventral scales, with air chambers of air canals opening dorsally by pores; cells typically 1Key adapted from Schuster (1963a). a... . 67 dimorphic, a small minority of generally smaller cells each with a single, large oil body, but no chloroplasts, the large majority of cells with chlorOplasts only; capsule wall always unistratose; seta short, not extruding the sporophyte to any extent. Archegonia on Specialized thallus branches (carpocephala, archegoniophores) ..... Order MARCHANTIALES 2. Rhizoids, if present, all smooth, isolated thicken- ings excepted; plants leafy or thallose, if thallose : delicate and translucent, without air chambers or pores; cells not sharply dimorphic, oil bodies usu- ally present in all chlorophyllose cells of gameto- Phyte; capsule wall 2-10 stratose; seta usually long and extruding the sporophyte ............ 3 3. Plants clearly and uniformly leafy; archegonia usually terminal on shoots resulting in a cessation of plant growth; capsule wall 2-10 stratose, the .cells variously thickened or lacking band-line thickenings . Order JUNGERMANNIALES Plants usually thallose (leafy in Noteroclagg); ”Che" gonia and antheridia scattered on dorsal thallus sur- face (occasionally on short thallus branches), or in dorsal 9170095. not causing cessation in growth of plant; capsule wall 2-5 stratose ........ Order METZGERIALES 68 C. Order Jungermanniales: Key t9 the Genera of the Falkland Islandsl 1. Leaves of main stems with orientation (and usually in- sertion) incubous, at least at dorsal end of insertion, the leaves usually incubously shingled (but ventral portions sometimes modified to form a lobule or water sac); rhizoids always restricted in origin, never scattered on stem, often lacking or rare .......... 2 1. Leaves of main stems varying from transverse in inser- tion to succubously inserted and oriented, to almost longitudinally (horizontally) oriented; never with ventral margin of leaf bearing, or transformed into, a lobule or sac; rhizoids often copious, Often scat- tered on ventral face of stem ............... l4 2. Leaves divided into a larger dorsal lobe and a solitary, smaller ventral lobe, which is usually sac like or pouch like; branching exclusively lateral, the branches terminal or infra-axillary, neither vegetative nor sexual branches ever ven— tral or axillary; underleaves 0-2 lobed or absent. . . .3 1 Leaves various, but never with ventral base or lobe modified to form a flap or inflated sac; branching various; underleaves always very large, \varying from 2-4 lobed ................. 8 _ 1 This eneric key is ada ' 'd bl dification pted, with conST era, e mo . , from Schuster (1963a) with portions from Schuster (l964b, 1965a, 1966). 69 3. Underleaves lacking; rhizoids (when present) in fascicles from lobuli, absent from stem. Branches uniformly infra-axillary; plants light to yellow green; leaves with a wide, J- or U-shaped insertion; perianth flat- tened, mouth wide, truncate ................. 539213 3. Underleaves present, conSpicuous, some or all of them with rhizoids at their bases ................ 4 4. Lobuli, at least in large part, galeate, almost free from the dorsal lobe; plants often with reddish or brownish, or dark green coloration ...... 5 4. Lobuli not saccate or galeate, united for most of their length with dorsal lobe along an elon- gate keel (the water-sac formed partly by lobule, partly by the opposed portion of the lobe); plants green to yellow green. Plants often small and delicate; perianth usually pentagonal, with a beak .......................... 6 . Underleaves undivided (in Falkland species); plants Without perianths, but with a clavate, variously para- Phyllose coelocaule. Leaves entire or ciliate . . . .Gackstroemia Underleaves bifid; plants with perianths, but without a coelocaule. Leaf margins entire ..... 4 ...... M 70 6. Hyaline papilla of lobule distal to apical tooth. Leaves convex, often strongly so; lobuli always : inflated; plants firm, often dull and opaque . .Cheilolejeunea 6. Hyaline papilla of lobule proximal to apical tooth.......... ................ 7 7. Lobe apices acute to accuminate, narrowly ovate to lanceolate to elliptical, 1.5 x 3 x as long as wide. Leaves ocellate; underleaves obdeltoid, with divergent blunt to rounded lobes ............... Harpalejeunea l. Lobe apices blunt to rounded, usually less than 1.5 X as long as wide. Leaves with or without ocelli. Plants delicate, usually with leaves flat or weakly convex; leaf trigones absent or small ............ Lejeunea 8. Plants only with lateral, terminal branching, without ventral branches and never with ventral flagella or stolons; leaves divided for 0.6 or more their length; leaf and underleaf margins and segments often ciliate or laciniate. Leaves nearly transversely inserted; cells rigid, with large, course, sometimes confluent trigones; plants with- out perianths but with a fleshy coelocaule densely covered with scales and paraphyllia; plants with subfloral innovations ................ Lepicolea ,‘s . V . 71 8. Plants with branching not exclusively lateral and terminal; sometimes vegetative and/or sexual branches partly or wholly ventral in origin and abbreviated; plants often with ventral stolons or flagella (at least from older parts of plant); leaves rarely very deeply lobed, never with longly ciliate margins ..... 9 9. Plants iSOphyllous (both on main stems and branches), the underleaves similar in size, and usually in form, to lateral leaves; branches usually few and irregular, subfloral innovations excepted, all or in large part ventral and axillary, slender flagella usually lack- ing; sexual branches never abbreviated and ventral. Lateral leaves 2-4-fid; leaf insertion only weakly incubous, the orientation subtransverse .......... 10 9. Plants normally somewhat to strongly aniSOphyllous, the underleaves differing in size (and usually in form) from lateral leaves, at least on the branches, underleaves often (if lobed) with fewer lobes than lateral leaves, or else more shallowly lobed; branches (usually) in large part lateral, usually terminal, the plants often regularly l-2-pinnate1y branched; stolons, flagella or rhizomes bearing vestigial leaves usually present; sexual branches always short, postical .................. 11 Le... 72 10. Male bracteoles with antheridia; leaf margins entire in Falkland species (occasionally dentate only at base of ventral margin); perianth distinct; plants usually without conspicuous pigmentation, at most palebrown . . . . . . . . . . . . . . . . . . Triandrpphyllum 10. Male bracteoles without antheridia; leaf margins usually a dentate; a firm fleshy coelocaule pres- ent, with vestigial perianth at its tip; plants usually with reddish and/or chestnut brown to fuscous pigmentation ................ Isotachi s 11. Lateral branches irregular, plastic, i.e., Frullania-, Microlepidozia- (and in M. mollis Acromastigum-type), intercalary branching both ventral and lateral. Under- leaves frequently with 1 segment shorter than others . . Microlepidozia ll. Lateral branches all of Frullania type ........... 12 12. Stem without a hyaloderm layer (cortical cells at thick walled, not conSpicuously larger than medullary cells); leaf cells often with thick walls and :guttulate lumina; leaves usually con- spicuously asymmetric (dorsal margin longer and/or arched) ....................... Lepidozia 12. Stem with a conspicuous hyaloderm layer, formed of large cells, with walls little or not thickened, D r a u. a. a... .. .. , i. . -4 I v». , n 1 '- - .—. v- ‘- n ,‘ r g n- P. \. .u a u. I n ., .5. a » c 1 . I . . .a .4 ' I .b . . b o . ' I . .I t . . . ‘- ~ I . m a a . c" i . 73 contrasted to smaller medullary cells often with thick walls; leaf cells without trigones, usually large, hyaline, thin or slightly thick-walled; leaves usually symmetrical .............. l3 l3. Leaves with lamina 5-12 cells high, distinct; leaf lobes triangular, never filiform. Lamina cells in longitud~ inal rows ....................... Neol epi dozi a l3. Leaves with lamina 0-4 cells high; leaf lobes uniseriate throughout, filiform and terete, at most 2 cells wide in basal cell tier ................... Telaranea 14. Stem with underleaves conspicuous throughout (not merely with a minute underleaf here and there), even on sterile stems (except Lophozia, which is gemmiparous and has quadrifid leaves with cilia on the ventral margins) ................ 15 14. Stem with underleaves either lacking on sterile stems, or minute, or mere small, scattered cilia or laciniae, or groups of slime papillae, never conspicuous (exc. Lophozia). Rhizoids almost always scattered over ventral stem surface if present at all .................... 33 15. Leaves sharply complicate bilobed, with a i smaller (rarely subequal) dorsal lobe lying over a larger ventral lobe . . 4 ..................... 16 .4 74 15. Leaves not complicate bilobed if lobed at all, the lobes 17. 17. lying in nearly the same plane, or leaf merely concave or t naviculariform, never folded ............. 17 16. Apex of dorsal lobe oriented towards apex of ventral lobe, keel between lobes usually with a broad wing, leaf lobes undivided, never bifid; cells of leaf lobes irregularly arranged, not in tiers; sporo- phyte in a terminal coelocaule lying in the axis of the stem .................... Schistochila l6. Apex of dorsal lobe at angle to axis of ventral lobe, the lobes i divergent, keel between lobes wingless, leaf lobes bifid; cells of leaf lobe oriented in i distinct tiers; sporOphyte in a marsupium ..................... Balantiopsis Mature leaves 3-4 (5-6) lobed, often very deeply so, the lobes usually ending in cilia or acuminate, often the lobe margins with cilia or teeth; stem without a distinct, pellucid hyaloderm. Underleaves very large, ca. 0.5-0.9 the area of leaves, similarly lobed and/or ciliate; rhizoids usually sharply restricted to under- leaf bases ..... p ................... 18 Mature leaves unlobed to 2(3-4) lobed (occasionally with accessory small lobes or teeth, or occasional leaves deeply trifid), if leaves lobed, margins entire or dentate, but not with long cilia; stem sometimes with a hyaloderm ..................... l9 1 V , \ 75 18. Lobes of mature leaves (usually of vegetative shoot sectors, at least in and below gynoecia) with op- posed, sharp teeth or cilia, usually of both disk margins and of lobes, rarely of one only; peri- chaetial bracts freely .Spinose-dentate to copiously ciliate; perianth trigonous, wide at open mouth, the mouth .+. ci 1i ate .................. Temnoma 18. Lobes of mature leaves quadrifid, without cilia or teeth (or, rarely, with an isolated tooth on one or both margins of the disk); perichaetial bracts 4-fid, without trace of teeth or cilia; perianth terete below, basically trigonous only in distal 0.2-O.25, closely contracted to the narrow mouth, the mouth denticulate-lobulate to short ciliate. Leaves and underleaves equally (3) 4-fid, never bisbifid, unistratose throughout, with segment apices setaceous; plants distinctly aniSOphyllous, the underleaves at most 0.5 X the area of lateral leaves, stem with undifferentiated cortex, corti- cal cells not distinctly smaller than medullary; pl ants brown ................... Archeochaete 19. Plants with perfect isophyl 1y; leaves undivided or retuse, 2-3 stratose in median, basal portion; leaf and under-. leaf margins and apices with slime papillae. Branching exclusively intercalary; perianth trigonous, fusiform 2). 76 to elongate, contracted toward the mouth, the mouth with small lobes and slime papillae .......... Hygrobiellgasis . Plants quite anisophyllous, or if nearly iSOphyllous, as in Pachyglossa, then with stiffly patent leaves without slime papillae, never julaceous; leaves undivided to bifid, unistratose, or if pluristratose at the base, as in Clasmatocolea Leorgiensis, then strongly aniso- phyllous; leaves and underleaves without slime papillae. Terminal branching present or absent ........... 20 20. 20. Rhizoids frequent to common, scattered over ven- tral stem surface; underleaves usually ovate to triangular, unlobed or obscurely bilobed, or large, bifid and ciliate; cells each with dense, con5picuous papillae (exc. Cephaloziella dusenii where small or absent ................. 21 Rhizoids always restricted in origin, only at underleaf (sometimes also at leaf) bases, often largely confined to reduced leaves and underleaves of stolons, flagella, or rhizomes, never scattered; cells smooth or with scattered, small fine papillae (if, rarely, with coarser papillae, see Clasmato- colea) ........................ 24 Underleaves large, ciliate and/or bifid; without stolons or flagella; vigorous plants, usually i brownish, to - . a ‘ r a .- " .«Ao ~ - .. .- i. ' . a .. h o 1, .- - i No .-. .- v ., . - ~- .1 'o . o ..' 7“ v . o ‘v ~. .~ . u. u..’ ‘ .. ‘ a . H, I. w ‘. o ’2 21. 23. 23. 77 3-5 mm wide. Cells with small or moderately coarse trigones; cuticle with high, coarse, hemispherical papillae; perianth apex twisted ............ Roivainenia Underleaves small or minute, eciliate, usually unlobed; often with flagella or stolons; small plants (shoots to 0.7-2 mm wide) ..................... 22 22. Leaves quite unequally bilobed, the dorsal lobe smaller, succubously inserted throughout, large (leaf width much greater than stem width); peri- anth lacking, plants with marsupia ......... Acrobolbus 22. Leaves subequally or equally bilobed, with at least dorsal half transverselyinserted (occa- sionally dorsal half subsuccubous in Cephalolobpg scabrellus), small (little broader than stem), distant; perianth present. Plants with Cephaloz- iella-like facies ................ . . . 23 Branching exclusively lateral intercalary; leaf cuticle with conspicuous, very coarse, hemispherical, hyaline papillae ...................... .Qphalolobps Branching usually ventral intercalary, occasionally of Frullania type; leaf cuticle (at least in _C. dusenii) smooth to at most with low, rounded papillae . . . . C_ephaloziella 24. Stem very soft, pellucid, with a cortex of large hyaline cells surrounding a a small celled medulla 25. 25. 78 (which is usually visible through the cortex by transmitted light); cells not collenchymatous, hyaline, large; ventral terminal branching present (in Paracromastigm) ................. 25 24. Stem without a transparent, colorless hyaloderm; the cortical cells never conspicuously larger than medullary cells, the medullary cells not visible through the cortex; cells various, often with dis- tinct trigones; ventral terminal branching absent. . . 27 Leaves cephalozioid (leaves at least feebly dorsally inclined), transverse to distinctly succubously in- serted, transverse-to basically succubously oriented. Branching mostly intercalary, both lateral and ventral, Frullania- type branches rare or sporadic, Acromastigum- type branching in some taxa, Microlepidozia-type branching absent ................. Pseudocephalozi a Leaves lepidozioid (leaflobes turned ventrally), transversely to feebly incubously inserted, basic- ally incubously or transverselyoriented . . . ...... 26 26. Branching predominantly or nearly exclusively lateral- and ventral—intercalary, with rare or sporadic Frullania- and Microlepidozia-type branches, never with Acromastjgum-type branching; antheri di a1 stalk l-seri ate ........... Hyalolepidozia 27. 27. 79 26. Branching predominantly or nearly exclusively term- inal, of Acromastigum- and Frullania-types, never with Microlepidozia-type branching, with rare ventral-intercalary branches, never with lateral- intercalary branching; antheridial stalk 2-seriate ....................... Paracromastigum Plants producing marsupia, perianths absent; gynocia (and androecia) always on abbreviated ventral branches. Leaves ovate, undivided or bidentate (occasionally short bifid in S. vasculosum), cuticle very densely papillose; underleaves 1/2-3/4 bifid (in Falkland Species), free from lateral leaves, much smaller than leaves; plants dull, Opaque, gray or light-green plants developing (often) some brownish pigmentation. . . . Saccogynidium Plants producing perianths, perigynia lacking; gynoecia on apices of unmodified stems or short lateral branches, rarely on short ventral branches ........ 28 28. Cell walls with enlarged, knot-like secondary swellings giving the leaf and underleaf margins a crenulated appearance; stems with single layered cortex of larger cells than medullary. Leaves transversely (or occasionally subsuccubously) inserted ...................... Ejgafettoa 'r .l‘ u 29. 29. 31. 80 28. Cell walls without secondary thickenings, the leaf and underleaf margins plane; stems never with cortex of larger cells than medulla .......... 29 Plants rigid, erect, almost iSOphyllous, with Spreading, pluristratose leaves and underl eaves; both leaves and underleaves unlobed or at most retuse at apex, both nearly transversely inserted; plants stoloniferous . . Pachyglossa Plants lax, or at least not rigid, usually prostrate, _+_ anisophyllous, the leaves and underleaves unistratose; stolons usually lacking .................. 3O 30. Perianth strongly laterally compressed, mouth truncate, wide, 2-lipped. Plants often i in- tensel y brownish ................. L_eptoscyphus 30. Perianth trigonous to trigonous inflated ....... 31 Perianth a: inflated, restricted to strongly abbrev- iated lateral-intercalary branches; leaves unlobed or to 3 spinose-dentate. . . . . . . . .......... Chiloscyphus Perianth normally on more or less long branches, (at least in large part; branches rarely abbreviated, but never consistently so); leaves undivided or bifid. . . 32 32. Leaves convex and with apices decurved or de- flexed ...... .................l;gphocolea , o o-‘ ' .n c l- .o u - ,, .. o a. , ID , o- o a -. i ‘o p. ' .1. - V . a o , H. _ . s . o,” I . a w... a o,‘ ..| H "a. I ‘o e- i . ~.. 0'. . . n 1.. ‘1. -'s . c 0‘ l .- o 1“ u .. I 9. , . s 33. 33. 35. 35. 81 32. Leaves deeply adaxially concave (at least slightly concave in C. vermicularis). Leaves frequently suborbincular to reniform in shape ....... Clasmatocolea Leaves a. sharply complicate-bilobed, transversely to succubously inserted and oriented. Leaf margins (and/or apices) often :t denticulate to ciliate .......... 34 Leaves lobed or not, but if lobed, never with lobes sharply bent over each other- Perianth (if present), not sharply dorsiventrally compressed ........... 35 34. Leaf lobes : short bifid; rhizoids (Often present only on small plants) in fascicles from the stem at the ventral leaf bases; terminal branching frequent, or Frullania type ......... Blepharidgphyllum 34. Leaf lobes undivided, never bifid; rhizoids scattered over ventral stem surface; branching only intercalary, lateral ............ .D_ipl ophyllum Stem soft textured, consisting of a conspicuous, hyaline cortex of enlarged cells (surrounding a medulla of much smaller, t thick-walled cells usually distinct by transmitted light, or medul- lary cells also soft but large). Leaves bilobed; cells thin-walled, t hyaline . . . . . . . . . . . . . . . 35 Stem anatomy very various, but never with a hyaline cortex of large cells surrounding amedulla of smaller, 3: thick-walled cells. . 37 37. 37. 82 36. Leaf insertion transverse at least dorsally, extended to stem midline; pigmentation, when present, purplish or brownish; leaves : canaliculate, plano-distichous; branching plastic, always with free Frullania-type branches, plus lateral-intercalary and ventral- intercalary branches; gynoecia terminal on leading axes ..................... Metagygrobiel la 36. Leaf insertion oblique throughout, the dorsal end oblique to almost longitudinal; pigmentation never purplish, usually absent; leaves never plano- distichous, flat to concave but not canalicul ate; branching from axils of lateral leaves absent; gynoecia normally on sort, ventral branches which Occasionally are elongated ...... . . ..... Cgphalozia Leaves inserted t transversely (at least in part of the dorsal half of the insertion) and transversely oriented, generally appearing 1 pectinate when not densely appressed-imbricate. . . . . . . . . . . . . . . . 38 Leaves obliquely inserted and :- succubously oriented, at least half of the insertion quite oblique on stem tonearlyhorizontal.................,,41 38. Leaves unlobed or to 1/5 bifid, margins often with l-several rows of hyaline cells forming a border......................l_igrzogobryum 39. 39. 83 38. Leaves usually bilobed O.35-O.5 their length; leaves not with l-several rows of leaf cells forming a hyal ine border .................... 39 Plants minute or small, wiry; the leaves remote, bilobed 0.5-0.75 their length; leaf lobes (2) 3-8 cells broad. Cells very small, equally thick walled, lacking trigones; perianth mouth crenulate .............. C_ephaloz1'ella Plants large; the leaves bilobed ca. 0.35-0.45 their length, their lobes 8 or more cells broad ........ . 40 40. 40. Cells with coarse, conspicuous trigones and thin to sinuous intervening walls; leaves strongly antically secund, the dorsal end of the line of insertion arcuate and : decurrent along dorsal midline of stem ................. AnastrOphyllum Cells non-collenchymatous or with rather small trigones; leaves laterally patent and pectinate- distichous, not antically second; insertion dorsally transverse, not decurrent. Cuticle with large, coarse papillae; plants green, leaves brownish distally; lateral-intercalary branching only; leaf lobes : sul cate; pl ants autOECi OUS o o o o o o o o o 9 o o o o o o g g . .A—CIO‘ ophozia 41. 41. 43. 43. 84 Plants with perianths terminal on leafy, main or lateral stems; androecia intercalary or terminal on leading stems. . . . . ...................... 42 Plants without perianths, with pendent perigynia or without perianth and without perigynium, the erect shoot-calyptra fleshy and rigid. Leaves various, often entire or irregularly dentate ............ 48 42. Branching uniformly arising from the dorsal end of the leaf axil, all intercalary. Flagella common, wiry, positively geotrOpic; leaves bilobed; leaf cells with large trigones .......... . .Andrewsianthus 42. Branching various, if intercalary, arising from the ventral portion of the leaf axil , near the ventral portion of leaf base, or ventrally ...... 43 Perianth laterally sharply compressed, wide and trun- cate at mouth. Vegetative branches lateral intercalary and/or terminal and of Frullania-type, never ventral intercal ary ...................... flagiochi la Perianths terete below, pluriplicate and contracted to the mouth. . . . ..................... 44 44. Leaves unlobed, usually rounded at apex ........ 45 44. Leaves consistently 2-4 lobed. Underleaves lacin- iate to ciliate or reduced .............. 45 . - in _ ’ n -. F. ‘ . O s ‘ I n u ,r . 1 D.- . . v o l , ' .OM. ' t n d'. ‘ t I ‘§ I 45. 45. 47. 47. 85 Ventral leaf margin frequently slightly to strongly re— flexed, leaves short bifid; underleaves present, of only a few cells only toward the stem apices (in Falkland species) ........................ Anastrepta Ventral leaf margin plane, not reflexed, leaves 2-4 lobed, often deeply so; underleaves, when present, may be large and conspicuous (ciliate in the Falkland species , _L_. hatcheri) ................... Lophozia 46. Involucral bracts unlobed and edentate; bracteole lacking; plants soft textured, usually prostrate growing, small in size .............. Jungermannia 46. Involucral bracts t lobed and/or dentate, : smaller than leaves; usually with bracteole present; plants ’1 firm, usually erect growing, often robust, highly pigmented. Cells often rigid and thick walled ............. . . . 47 Terminal branching completely absent; flagellae regu- larly produced. Bracteoles nearly as long as bracts; perianth mouth entire to weakly crenulate; plants usually brown to red brown .............. Cryptochila Terminal branching present, if only isolated; flagel- “*— lae only rarely and Sporadically produced, mostly absent ........................ 9311195001911 a . . v . . ‘1 P ‘ . a 'I a. . ‘7 o ' ‘ 'r- u‘ ‘0. I I ‘u._ r '9. 51 .Q 'Q.‘ .u ., . .l' ‘ a. .' _ u I“ .- I. u o. o . .'a 49. 49. 51. 48. 48. 86 Plants without a marsupium, the erect calyptra be- coming thick, green, massive, enclosing the develop- ing sporophyte; leaves undivided; plants erect . . . . 49 Plants with a marsupium; leaves undivided or 2-4 lobed, plants erector depressed ........... 50 Stem with a collapsing hyaloderm (thus scabrous with age); dorsal leaf-margin 4-5 cells thick (in Falkland species); plants green, without secondary pigmentation. Subhyalodermal cells in 2-4 layers of thick walled cells; stems 20-45 cells in diameter . . . . ...... Wettsteinia Stem without a collapsing hyaloderm, smooth, never roughened; leaves unistratose throughout; plants green to red brown ................... Adelanthus 50. 50. Leaves unlobed, leaf margins entire; marsupium slenderly cylindrical, deeply penetrating sub- stratum. Plants closely prostrate ...... . . .L_e_thocole_a_ Leaves 2-4 lobed (or if undivided, then with leaf margins dentate to lobate), leaf margins entire-dentate-lobate; marsupium : conoidal to wide cylindrical in shape, not deeply penetrat- ing substratum ................... . 51 Leafy shoots .+. ascending, not adhering to substratum by rhizoids. Gynoeci a and androecia variable in position, acrotonic or basitonic; plants a clear, 87 translucentrto opaque green, often whitish when dead; capsule tips acute . ................. Marsupidium 51. Leafy shoots -: creeping, often entire axis including shoot tips adhering to the substratum by rhizoids ..... 52 52. Leaves bilobed, often asymmetrically so; cuticle coarsely papillose; rhizoids few, arising from ventral leaf bases, never in dense, decurrent bundles on stem; plants without gemmae; antheridia solitary. Capsule tips acute ............ Agrobol bus 52. Leaves 3-4 lobed; cuticle smooth; rhizoids dense, decurrent along stem in fascicles; plants gemmi- parous; antheridia 1-10 per bract ........ Austrolophozia 88 HERBERTACEAE K. Miill. (Freib.) _ex Fulf. 8: Hatch. Bryologist 61 (4): 284. 1959. K. M1111. (Freib.) i_r_1_Rabenh. Kryptogamenfl. 6 (l): 557. 1954, _no_m. pg. Triandrophyllum Fulf.&Hatch. Bryologist 64 (4): 349. 1962. Triandrophyllum Fulf. LHatch. Bryologist 61 (4): 277. 1959, nom. illeg., sin. gen. typ. Triandrophyllum Grolle, Bryologist 64 (1): 25. 1961, m. pu_d_. Triandroihyllum subtrifidum (Hook. f. 8 Tayl.) Fulf. 8 Hatch. Lwlgermannia subtrifida Hook. f. 81 Tayl. London Jour. Bot. 3: 579. 1844. Isotachis subtrifida (Hook. f. 8 Tayl.) Mitt. mHook. f. Bot. Ant. Voy. 2 (2): 149. 1854. Mastigophora subtrifidp (Hook. f. a Tayl.) Hodg. Revue Bryol. Lichén. 18 (1-2): 25. 1949. hiandrophyllum subtrifidum (Hook. f. & Tayl.) Fulf. a Hatch. Bryologist 64 (4): 350. 1962. Lectotype (_cj. Fulford 1963b): Tasmania, H9333 (BM - pppligl). Mripensis Spruce, Trans. Proc. Bot. Soc. Edinb. 15: 339. 1885, m. _de_e_ Fulford & Hatcher (1959). Original material: Ecuador, "Andes Quintenses," M. Pichincha, _S_prygg §_.!_1_. (FH. G. NY) - cited in Fulford (1963b). o .1.‘ . . v ‘- 89 Isotachis angeps Mass. Nuovo G. Bot. Ital. 17: 219. pl. 15. _f_. _9_. 1885, _S_m. _fid_e Fulford 3 Hatcher (1959). Original material: Chile, Prov. Magallanes, I. Basket, Spegazzini s.n. (G) - cited in Fulford (1963b). Isotachis nordenskjoeldii Steph. Sp. Hep. 3: 659. 1909, pm. _f_i_c_l_e_ Fulford 6 Hatcher (1959). Original material: Chile, Prov. Magallanes, L. Dickson, Nordenskjiild 905 p.p. (FH) - cited in Fulford (1963b). Hastquphora beckettiana Steph. Sp. Hep. 4: 35. 1909, gm. pf. Hodg- son (1949). Original material: New Zealand, South Island, Wai- mate, 1901 Beckett 254 (G) - cited in Grolle (l964c). moscyphus schismoides Steph. K. Svenska VetenskAkad Handl. 46 (9): 38.313. _f_.g. 1911, 331p. ji_dg Grolle (l964c). Leptoscyphus schismoides (Steph.) Kiihnem. Revta Cent. Estud. Doct. Cienc. Nat., B. Aires l: 177. 1937. Mylia schismoides (Steph.) Kiihnem. Lilloa 19:341. 1949. Lectotype (gt. Grolle 1964c): Falkland 15., Mt. Adam, 700m., lelg 81 Skottsberg _s_._r_1_. (UPS - 119111191). _IS_0’C_aChiS halleana Steph. K. Svenska VetenskAkad. Handl. 46 (9): 69. L21. _a_,p_. 1911, syn. fide Fulford & Hatcher (1959). Original material: Chile, Prov. Magallanes, F. Peel; Argentina, Tierra del Fuego, R. Azopardo and Ushuaia, R. Olivia, Halle 81 Skottsberg (a) - cited in Fulford (1963b). Mlanciloba Steph. K. Svenska VetenskAkad Handl. 46 (9): 70. _C_. 261,31. 1911, _syn. flgg Fulford & Hatcher (1959). Original material: Falkland 15., Mt. Usborne, Skottsberg §_._p. (G) - cited in Fulford (1963b). 90 Isotachis mutabilis Herz. Archos Esc. Farm. COrdoba 7: 26. 1938, pyp. fl_d_e_ Fulford & Hatcher (1959). Original material: Chile, Prov. Cautin, Pucon, Hosseus 3.3. (G) - cited in Fulford (1963b). Taxonomical remarks: Triandrophyllum subtrifidum is closely related to T. trifidum. The former has underleaf margins and ventral leaf margins entire or only occasionally with a single tooth, and with leaf cells thin walled to very slightly thickened. Triandrgahyl- _lu_m_trifidum possesses underleaf margins and ventral leaf margins with several sharp teeth, especially toward the bases and leaf cells uniformly thick walled. One collection, Engel 2869, has several under leaf and ventral leaf margins dentate and leaf cells slightly thickened. Hhile the plant is somewhat intermediate between _T_. subtrifidum and _T_. trifidum, I believe it is but a variant of l. subtrifidum. Hodgson (1967, p. 197) states in her discussion of l. trifidum in New Zealand, Examination of many specimens may reveal more and more of this species with the toothed lower margins. Excepting these toothed margins I can find no other character dis- tinguishing l. trifidum from _T_. subtrifidum, nor do Ful- ford and Hatcher (1961, p. 350) mention any other in their key to the four species of the genus. With the marginal vagaries of Isotachis species in mind, one cannot help but wonder about this, but it is doubtless the only prac- ticable way to treat with this complex. Further study of the complex is necessary and may Show _T_. trifidum to be conspecific with I. subtrifidum. Ecology: In the Falklands this species is of sporadic occur- rence on soil or rock under rock ledges of sheltered high altitude cliffs. It occurs rarely in stands of Hebe in dwarf shrub heaths. 91 Phytogeography: This species has an amphipacific temperate distribution with occurrences in New Zealand and Tasmania and north in the Andes to Guatemala. Literature Records (FALKLANDS): Anonxmous - (Fulford & Hat- cher1959, 1962, Kiihnemann 1937 as Isotachis lanciloba 8 LgptosEyphus schismoides, 1949 as l. lanciloba & Mylia schismoides); Skottsberg - Fulford 1963b, Stephani 1922 as L. lanciloba), Mt. Usborne (Bonner 1966 as l. lanciloba), Mt. Adam (Skottsberg 1913 & Stephani 1922 as Leiosgyphus schismoides); Skottsberg g Halle - Mt. Adam (Grolle 1962). Additional Literature Records: Tierra del Fuego: Anonxmous (Fulford a Hatcher 1959, 1962, Grolle 1969, Kiihnemann 1937 8 1949 as Isotachis anceps 3 _I_. halleana); Dusén- Fuegia, I. Desolacion (Steph- ani 1909 as _I_. ancgps); Halle - Fuegia (Bonner 1966 as Isotachis hal- l_e_a_np, Fulford l963b, Stephani 1922 as _I_. halleana); Santesson - Ushuaia (Arnell 1955 & Miiller 1955 as l. nordenskjoeldii); Spegazzini .. I. Basket (Bonner 1966 & Massalongo 1927 as _I_. anceps , I. Basket-Cabo Desolacion (Fulford l963b). . PATAGONIA: Anonymous - (Fulford 81 Hatcher 1959, 1962), Strait of Magellan (Kiihnemann 1937 & 1949 as _I_. anceps); Bugg- R. Aisen (Stephani 1900 as l. anceps); Hosseus - Pucon (Bonner 1966 as _I_. mutabilis, V. Villarrica (Fulford l963b); Nordenskjold .. (Fulford 1963b), L. Dickson (Bonner 1966 as _I_. nordenskjoeldii; M- C. Tesoro (Herzog 1954 as _I_. m ; Skottsberg - (Fulford l963b); Spegazzini - Strait of Magellan (Stephani 1909 as L. m), BOLIVIA: 9.119.959. - Unduavi (Fulford 1963b); [I_e_r_zg_g_ - Tablas - 3400 m, (Stephani 1916 as _I_. rim). PERU: 1m - Cuzco (Fulford l963b) 92 ECUADOR: Jameson - Quito (Fulford l963b); Spruce — M. Pichincha (Bonner 1966 as Isotachis ripensis, Fulford 1963b). COLOMBIA: MEE' Huila-Cauca: Péramo de Las Papas, Cauca: Pa’ramo de Las Papas (Fulford 1963b). COSTA RICA: Standley — San 0056: Las Nubes - 1500-1900 m. (Fulford l963b, Herzog 1938). GUATEMALA: Anonymous - (Grolle 1969); Steyermark - San Marcos: V. Tacana (Fulford l963b). TRISTAN DA CUNHA: Christophersen A Mejland - Cave Gulch (Arnell 1958 as Isotachis halleana). NEW ZEALAND: Anonymous - (Bonner 1966 as Isotachis subtrifida. NORTH ISLAND (NEW ZEALAND): Hooker - Bay of Islands (Hooker 1867 & Mitten 1885 as Isotachis). TASMANIA: Anony- m- Mt. Wellington (Rodway 1916 as Isotachis); Hooker (G. L. 8 N. 1847 as Jungermannia); Lawrence (Mitten 1860 as Isotachis); Weymouth (Pearson 1924 as Isotachis). FALKLAND SPECIMENS SEEN: EAST FALKLANDS: MT. USBORNE REGION, ridge between Mt. Usbornes l 81 2, 685 m. (2515. 2531 _B_ 81 2152) WEST FALKLANDS: MT. ADAM, 700 m., 13 December 1907 flp_l_l_e_ and Skottsberg 14L syntype of Leioscyphus schismoides (S-PA); ridge south of north- ern Take, 610 m. (3026, 3036 A, 3040, _8_v_ 3042 A). WESTPOINT ISLAND, near The Waterfall, 30-90 m. (2869-c.9). Triandrophyllum Species Excluded from the Falklands Triandrophyllum durum (Steph.) Fulf. & Hatch. The Falkland report originates from Grolle (1962) who stated Leioscyphug schis- moides, which is based upon a Falkland Island collection, was a sync. nm of _T_. durum. Triandrophyllum durum (Steph.) Fulf. & Hatch. is a 93 synonym of I. trifidum fide Fulford and Hatcher (1962). Grolle (l964c) later placed _L_. schismoides in the synonymy of _T_. subtrifidum. BLEPHAROSTOMACEAE Engel,l K. MIIll. (Freib) 1A Rabenh. Kryptogamenfl. 6(1): 590. 1954, ppm. ppg. Caulis irregulariter ramosus, ramificatione variabili , termin- ali ramificatione plerumque Frullaniae modo, minus frequens Micropolepido- ziae vel Acromastigi modo, intercalaribus ramis ventralibus vel later- alibus. Caulis cellulae omnes subaequimagnae. Plantae isophyllae ad moderate anisophyllae; folia transversa ad succubas, profunde quadrif- ida, rarius bilobata; amphigastria caulina similis foliis in forma. Rhizoidea solum ex amphigastriorum basibus rarius e foliorum basibus. Plantae dioicae vel monoicae. Gynoecia terminalia in caulinis majoribus. Perianthium unistratosum, teretium basin versus, 3-4-5 (-6) plicatum apicem versus; calpytra crassa. Capsula valvulis 2-5 stratosis. Sporae 8-16 u, aequales vel duplo elaterum latitudine; elateres 1-2 spiri . Typus: Blepharostoma Dum. Recueil Obs. Jungerm. 18. 1835. ( 1The diagnosis is based on the information in Schuster 1965)), C). u 94 ARCHEOCHAETE Schust. J. Hattori Bot. Lab. 26: 262. 1963. Archeochaete kuehnemannii Schust. Archeochaete kuehnemannii Schust. J. Hattori Bot. Lab. 26: 262. 1963. Holotype: Argentina, Terr. Tierra del Fuego, ca. 16-17 km. W. of Ushuaia, on road to Lapataia, C. Bandera, February 1961 Schustgg g Gamundi d_e_ Amos 58852 (RMS - hog 3:11,). Taxonomical remarks: Schuster (1965b) in his description of A_rchaeochaete kuehnemannii states the leaf lobe apices are uniseriate for a length of 4-6 (7) cells. The Falkland plants, however, have lobe apices uniseriate for a length of (1) 2-4 cells. Schuster (lg. 515.. p. 816), in distinguishing Archaeochaete states, . The more attenuate leaf apices, with the uniserate tipsalonger than in any species of Lophochaete, are (also) d1st1nct1ve. In Lpphochaete the tips are formed by, at NOSt. 2-3 superimposed cells. While the genera Qphochaete and Archaeochaete are distinct and well dellDEA. the leaf lobe apices should not be used as a character to differentiate them. Qphochaete has (a) leaves and underleaves sub- E9““ in Size and form and bisbifid and (b) perianths 4-5-6 plicate d'SFAW- Ancheochaete has (a) underleaves at most 1/2 the leaf $128 and leaves, when quadrifid, with sinuses subequal and (b) peri- anths 3 plicate distally. The only other member of the genus, A. temnomoides Schust., TS endemic to Tristan da Cunha. 95 Ecology-Phytogeoggphy: This rare species was found in a feldmark pool and on a cushion plant on a sheltered high altitude cliff. The type was collected in a Sppgngppnl bog in Tierra del Fuego, and I have found it in the same habitat in the Brunswick Peninsula. The ecology of the Falkland plants thus deviates con- siderably from that of the only known previous collections. FALKLAND SPECIMENS SEEN: EAST FALKLANDS: MT... OSBORNE REGION, summit of Mt. Usborne 1, c. 700 m. (_2__4_9_6_ _0_). WEST FALKLANDS: MT. ADAPT, ridge south of northern lake, 610 m. (3046). TEMNOMA Mitt. i_r_rHook. f. Handb. N. Z. Flora 753. 1867. _T_§_i_n_n_p_rpa_ Mitt. Phil. Trans. R. Soc. 168 (extra vol.): 32, 33. 18.79, _e_nr;. pyppggl Ignition, 1884). _H 1I regard the use of the genus "Teinnoma" as an typographic error since (a) Mitten used the spelling ____lemnoma on herbarium speci- mens; Dr. G. L. Smith [I am grateful to Dr. G. L. Smith, associate curator Of the New York Botanical Garden, for providing this infor- mation] states (in litt.), I have lOOkE—Et Mitten's handwritten labels on his specimens of Temnoma (in The New York Botanical Garden), and I can see how one might read “Teinnoma” if he didn't know the name. I'm inclined to think it's merely an error; (A) Temnoma is Greek derived from the word "Temno" meaning "cut off," 9'19 "Iganppp" has no Greek derivation, being merely a nonsense word wh1ch Mitten would have had little cause to use; further, Mitten stated in his original description for Temnoma: "from the truncate 96 Temnoma maadriparti tum (Hook.) Mitt. Jmermannia quadripartita Hook. Musci Exot. 2: p_1_. _l_l__7_. j. _-g. 1820. Temnoma quadripartitum (Hook.) Mitt. J. Linn. Soc. 15: 68. 1876. Blepharostoma guadripartitum (Hook.) Trev. Memorie Ist. Lomb. Sci. Lett. 111. 4: 417. 1877. Original material: Argentina, Terr. Tierra del Fuego, I. de los Estados, Mghgjgg (K.V) - cited in Schuster (1966c). Jungermannia poflphylla Angstr. Ofvers. K. VetenskAkad. FOrh. 29 (4): 11. 1872, §y_n. fide Pearson (1887); non Jungermannia pedo- phylla Thunb. Prodr. Fl. Cap. 2: 174. 1823 (= Symphyogyna pod - phylla (Thunb.) Nees 81 Mont. _i_n_G. L. & N. Syn. Hep. 481. 1846). Original material: Chile, Prov. Magallanes, Pto. del Hambre, Andersson §_.A, (S-PA) - cited in Schuster (1966c). _I_epidozia randii S. Arnell, Svensk. Bot. Tidskr. 47 (3): 417. j. 6. 1953, gyp. fide Schuster (1966c). Original material: Marion IS., @3276 (S-PA) - cited in Schuster (1966c). Taxonomical remarks: The Falkland material is referable to var. _qppdripartitum and not var. pseudopungens or var. randii. Ecology: This taxon is rather rare in the Falklands, where only a few gatherings were made in Cortaderia (wet depressions) and dwarf shrub heaths (under rock overhangs of outcrops). k. mouth of the perianth"; (c) as Temnoma is a Mitten genus, there would have been little reason for this author to alter the Spelling 12 Years later, ((1) in my opinion the fact that "Teinnoma" is twice mentioned in 1879 merely indicated the editor chose to use one spell- lng consistently, and (e) Mitten (1884') is merely a list of Kerguelen hepaticae based upon his 1879 paper, thus the presence of "Teinnoma" is a repetition of the earlier error and has little significance. o. .u-o u- , '64 1 . o 0 I . . a . a o I .n .a.. U. r _‘t I I. 1. ,., - D s .- ‘V I: . .I H. n I'l" . . u... . F n . ‘ .- l ‘I . n . 'o n 97 Phytogeography: This taxon has an amphipacific temperate distribution with northward extensions in the American sector (West Patagonia north to 39° 16' S., Andean Patagonia at P. N. Nahuel Huapi), and in the New Zealand sector into the mountains of North Island. It is also present in the subantarctic on Kerguelen and Marion Islands. The report from Tristan da Cunha in Arnell (1958) is based on a specimen of Archeochaete temnomoids (see Schuster 1966c). The report from Inaccessible Islands in Arnell (1958) is questionable (see Map 1). Literature Records TIERRA DEL FUEGO: Anonymous-(Kfi‘hnemann 1949), Fuegia (Schuster 1959), I. de los Estados (Schuster 1966c as var. randii); Davis - I. Hermite (Taylor 81 Hooker 1847b as Jungep- mannia); Dusén - Pto. Angosto (Schuster1966c as var. randii); Gas- pgflB. Angelito, Valle delle Fate 8: S. Agostini (Gola 1923 as BJppharostoma); Halle - Fuegia (Schuster 1966c as var. M); 5.9.119. ISkottsberg - Cta. Gomez (Schuster 1966c as var. 331911); My; - I. Clarence (actually I. Capitan Arecena) (Bescherelle & Massalongo 1.889 as gghgermannia, Schuster 1966c as var. _igypi_c3), I. Hermite (Fulford l963b, Schuster 1966c as var. gym); 9,» £3; Hatcher - Fuegia (Evans 1898 as Blepharostoma); M_e_n_z_i_gs_ - I. de los Estados (G. L. & N. 1845, Taylor 6 Hooker 1847b as Jungermannia, Fulford 1953b, SChUSteC 19556); Santesson - S. de Sorondo (Arnell 1955 as Blepharostoma, Hodgson 8 Allison 1962, Schuster 1966c as var. typica); Schuster - C. Garibaldi (Schuster 1966c as var. typica); Skottsbegg 8 Halle-Cta. Gomez (Stephani 1911 as Blepharostoma); Spegazzini - M. Conegliano & Pto. c... \ n.1, 98 Cook (Massalongo 1885 as Jungermannia, 1927 as Blepharostoma. PATA- GONIA: Anonypous - Strait of Magellan, West Channel (Schuster 1966c as var. typica), Strait of Magellan (Schiffner 1895 as Blepharostoma); Andersson - Pto. del Hambre (Angstrom 1872 as Jungermannia quadri- partitum, Fulford 1963b, Schuster 1966c as var. typica & var. randii); Cunningha - B. Halt (Fulford 1963b, Schuster 1966c as var. typica A randii); Donat - M. Tronador (Grolle l964a, Herzog 1957 as Blephar- ostoma, Schuster 1966c as var. typica); Ousén - Pto. Bueno (Schuster 1966c as var. randii, Stephani 1900 as Blepharostoma), R. Aisen (Ful- ford 1963b, Schuster 1966c as var. typica); Hariot - Patagonia (Schuster1966c as var. randii); Hosseus - Pucon (Grolle l964a, Her- 2091938 as Blgpharostoma); Oberdorfer - C. Puntiagudo (Grolle l964a); Schwabe - P. Magdalena (Grolle l964a, Herzog 1954 as Blepharostoma); Apttsberg 8: Halle - F. de Los Ventisqueros (Stephani 1911 as Blephar- ostoma). TRISTAN DA CUNHA: Christophersen & Mejland - (Arnell 1958 as Blepharostoma). INACCESSIBLE: Christophersen & Mejland (Arnell 1958 as Blepharostoma). MARION: Rand (Schuster 1966c as var. randii). KERGUELEN: Anonmous - Schiffner 1895); flppAg: (Mitten 1879); Moseley (Mitten 1876, 1879, 1884, Schuster 1966c as var. pypi__a_). ST. PAUL: WM - (Schuster 1966c as var. m, Stephani 1909). AUCKLAND: £09333: (Hodgson 1962). NEW ZEALAND. SOUTH ISLAND: Berggren - Bealey (Arthurs Pass Natl. Park) (Schuster 1966c as var. _typj_c_a_); Holdsworth - Arthur's Pass (Hodgson 8 Allison 1962, Schuster 1966c as var. M); M - S. of Haast Pass (Schuster 1966c as var. §ypi_cp 8 var. pseudopungens), Milford Sound ~ on track to Bowen Falls, Humboldt Mts., track from Lake Howden to Lake Mackenzie (Schuster 1966c as var. o O o u. ‘ I ‘ -. |"'I . III-O . ., n o .,,. In _ v o - to is. i . a c 9 . V I .‘ .. ‘ ,. ">1 1 O '0. II 5 ., . \f '\ n H \ u 99 gypipg). NORTH ISLAND (New Zealand); Oflgg - Ruahine Mts. (Hodgson IAllison 1962, Schuster 1966c as var. m); Hodgson - Ruahine Hts. (Schuster 1966c as var. Mp); Schuster - Mt. Hector (Schuster 1966c as var. gym 8 var. M), Mt. Egmont (Schuster 1966c as war. M). FALKLAND SPECIMENS SEEN: EAST FALKLANDS: MT. USBORNE REGION, The Gap, 275-290 m. (gym-c. 0+ + young 9 8 gig/'3). WEST FALKLANDS: FOX BAY REGION, summit of East Head, 180 m. (3428-c. per.); summit of Fox Bay Mt., 307 m. (3446-c. 0+). ADDITIONAL SPECIMEN SEEN: CHILE. AISEN: R. Aisen, January 1897 Dusen as Trichocolea verticillata (NY). ISOTACH I DACEAE Hatch. Nova Hedwigia 2: 579. 1960 as Isotachaceae; corrected by Grolle (l964c). Isotachis Mitt. igHook. f. Bot. Ant. Voy. 2. (2): 148. 1854. lOO Isotachis humectata (Hook. f. 8 Tayl.) Steph. Jungermannia humectata Hook. f. 8 Tayl. London Jour. Bot. 3: 462. 1844. Lophocolea humectata (Hook. f. 8 Tayl.) Steph. Bull. Herb. Boissier II. 6: 656. 1906 (Sp. Hep. 3: 72). Isotachis humec- flflook. f. 8 Tayl.) Steph. Sp. Hep. 3: 654. 1909. Lecto- type (_f_i_d£ Stephani 1906): Falkland Is., Hooker s.n. (BMI). Jungermannia madida Hook. f. 8 Tayl. London Jour. Bot. 3: 465. 1844, _n_o_n_Jungermannia madida Nees 1p Mart. Fl. Bras. 1: 362. 1833 (=2 Porella, fide Swails 1970, p. 249). Isotachis madida (Hook. f.8Tay1.) Mitt. 31 Hook. f. Bot. Ant. Voy. 2 (2): 149. p_1_. £8. _f_._2_. 1854. Original material: Chile, Prov. Magallanes, Cago de Hornos , prk_er;§.p. (NY, BM) - cited in Hatcher (l960). Isotachis fusca Steph. K. Svenska VetenskAkad. Handl. 46 (9): 68. f. 25 d, e. 1911, _S_y_n_. figs; Hatcher (1960). Original material: Chile, Prov. Chiloe, V. Corcovado, 31 July 1908 Hp_ll_e_ and im- MBA (NYI); Prov. Magallanes, F. Peel, F. de Los Ventisqueros MASkottsberg; Argentina, Terr. Tierra del Fuego, Ushuaia, R. Olivia, Halle 8 Skottsberg (non vidi). [sptachis pallens Steph. K. Svenska VetenskAkad. Handl. 46 (9): 70. f. 25 h-l. 1911, _S_yp. _f_j_dp Hatcher (1960). Original material: Chile, Prov. Chiloe, I. Chiloe, R. Pudeto, Skottsberg (G) - cited in Hatcher (1960). Wstriolata Steph. K. Svenska VetenskAkad. Handl. 46 (9): 71. f. 27 c2 d. 1911, gyp. _fidg Hatcher (1960). Original material: Chile, Prov. Magallanes, S. Otway, R. Grande, Skottsberg 1}} (MA) - cited in Hatcher (l960). . .4:- . I as ' - I I a I 101 Isotachis aeggifoliata Steph. Sp. Hep. 6: 349. 1922, gyp. fide Hatcher (1960). Original material: Bolivia, without specific location, Herzog 2848 (G) - cited in Hatcher (l960). Isotachis flavicans Steph. Sp. Hep. 6: 351. 1922, pyp. fide Hatcher (1960). Original material: Chile, without specific location, Skottsberg 739 (G) - cited in Hatcher (1960). Taxonomical remarks: Fulford (l963b, p. 71), in the generic key, spearates Triandrgahyllum from Isotachis in a couplet based upon the character "line of insertion hook-form or recurved at the dorsal end . . .," with Triandrophyllum possessing the character and App- _t_a_ch_i_s_lacking it. I have found A. humectata to have a strongly re- curved ("hook-form") line of insertion at the dorsal end, and Hatcher (l96l,p), 18, f. 583) illustrates this in a male plant of _I_. _hppigg- pajp. Vegetative characters are of little value in distinguishing the two genera and androecial and gynoecial characters must be relied upon. Hatcher (1960) records only ventral-intercalary branches for the genus Isotachis. Isotachis humectata, however, may produce ventral-intercalary branches as well as Frullania-type branches, the latter being quite commonl y developed. Frullania-type branching within Isotachis has been documented by Schuster (1963a, 1964d) and Crandall (1969). There has been considerable confusion regarding the type 10- cality and systematic position of Isotachis humectajgg. The type locality for Jungermannia humectata is the Falkland Islands, and the 102 plant was originally described by Hooker and Taylor (1844, p. 462). Stephani (1906) transferred the species to Lophocolea and commented, "Das originalexamplar dieser Pflanze in Kew Herbarium . . ." which effectively lectotypifies the plant to a specimen in the Kew Herbar- ium. Three years later Stephani transferred the species to Isotachis, listing it from both Campbell and Falkland Islands. Hatcher (1961) was in error in his treatment of the taxon. He states (p. 31) for A. humectata "Campbell's Island: without loc., Hooker, a portion of the original material of A. humectata (FH)," and treated the plant as a synonym of _I_. intortifolia without mentioning the Falkland Island locality, the Lophocolea transfer or the Kew Herbarium typi- fication. Ecology: This species occurs sporadically in streams in Cortaderia heaths above 215 m. where it often forms very extensive mats. The species seems to be restricted to Cortaderia heaths in the Falklands. Phytogeography: This Andean South American species occurs in the Falklands, Tierra del Fuego, Patagonian Channels, Valdivian region (West Patagonia north to 36° 43' S. and in Andean Patagonia in P. N. Nahuel Huapi), central Chile (Valparaiso), in the Andes north to Colombia and Tristan da Cunha (see Map 2). Literature Records (FALKLANDS): Anonmous - (Kiihnemann 1937 61949, Bonner 1966); Hooker (G. L. 8 N. 1847, Taylor 8 Hooker 18476, Stephani 1906, 1909). Additional Literature Records: AW (Kfihnemann 1937 8 1949 as _I_. fusca 8 _I_. madida), M. Foster (Hatcher 1960 as _I_. 111351133); 103 Eggp- Pto. Angosto (Fulford 1963b 8 Matcher 1960 as A. madida); Hooker - I. Hermite and/or Cabo de Hornos (Fulford l963b as A. madida, G. L. 8 N. 1847 as _I_. _1_n_agi_g_a_, Hatcher 1960 as _I_. _m_1p_c_i_j_d_a_, Taylor 8 Hooker 1847b as _I_. madida; Hyades - I. Hoste (Bescherelle 8 Massa- longo 1886 as A. madida); Skottsberg - Pto. Cook (Stephani 1905a as A. madida); Skottsberg 8 _l-Iill_e - L. Fagnano (Stephani 1911 as _I_. m_a_- 11'_d_a; Sggazzini - Pto. San Juan del Salvamiento (Massalongo 1885 as _I_. M8 1927). I. de los Estados (Fulford 1963b 8 Hatcher 1960 as _I_. m). PATAGONIA: Anonmus - (Kiihnemann 1937 8 1949 as _I_. M81. striolata); Dusén - Strait of Magellan (Fulford 1963b 8 Hatcher 1960 as A. M), Pto. Bueno (Fulford 1963b, Hatcher 1960 IStephani 1900 as _I_. madida), I. Guaitecas (Fulford 1963b 8 Hatcher 1960 8 Stephani 1900 as _I_. madida), Pto. Octay, Corral, Concepcion, Talcahuano (Fulford 1963b 8 Hatcher 1960 as A. madida); Hosseus - Corral (Herzog 1938 as A. madida), Valdivia (Fulford l963b 8 Hatcher 1960 as I_. madida); Junie - Pilmaiquen (Fulford 1963b 8 Hatcher 1960 as I. madida); Schwabe - I. Magdalena - Pto. Puyuhuapi (Herzog 1954 as I. madida); Skottsberg - I. Riesco (Fulford 1963b 8 Hatcher 1960 as 1. 93111.92): I. Chiloe (Stephani 1922 as _I_. pallens), Patagonia (Stephani 1922 as 1. Egg); Skottsberg 8 £111.19. - S. Otway - R. Grande (Stephani 1911 as A. madida), R. Pudeto (Stephani 1911 as _I_. 9991.91); W - Corral (Hatcher 1960 as I. madida); W .. L. Todos los Santos (Fulford 1963b 8 Hatcher 1960 as _I_. E9193). BOLIVIA: H_e_rng (Fulford 1963b 8 Hatcher 1960 as _I_. M). COLOMBIA: Bischler - Antioquia (Fulford 1963b 8 Hatcher 1960 as 1.1131133); Ae_i_r_- - Bogota (Fulford 1963b 8 Hatcher 1960 as _I_. _m_a_d_‘_l_c_l_a_). 104 TRISTAN DA CUNHA: Christgphersen 8 Mea‘jland (Fulford 1963b 8 Hatcher 1960 as A. madida), 550-800 m., 3rd Gulch-550-600 m. 8 above Burnt- wood 700 m. (Arnell 1958 as _I_. 1_n_a_d_ig_q 8 A. striolata); A853 - Soggy Plain, 740 m. (Arnell 1958 as A. M). FALKLAND SPECIMENS SEEN: EAST FALKLANDS: 59913; §_._n_., syntypes of Jungermannia humectata (FH, MICH). MT. USBORNE REGION, southeast slope of Mt. Usborne 2, c. 455 m. (2611, 2619 _B_, 2620 8 26_2_l_). DARWIN SETTLEMENT, near mouth of Ceritos arroyo (__2_6__7_6_ 8 M). WEST FALKLANDS: MT. ADAM, basin east of summit, 580-595 m. (_2_9_9_3_). ADDITIONAL SPECIMENS SEEN: CHILE. MAGALLANES: I. Desola- cion, Pto. Angosto, 16 April 1896 my); (NY). LEPICOLEACEAE Schust. Nova Hedwigia 5: 27. 31 January 1963. Schust. _epg Fulf. Hem. N.Y. Bot. Gdn. ll (1): 30. 15 March 1963. Lepicoleaceae Schust. Revue Bryol. Lichén. 26 (3-4): 126. 1957, 1193. 113551. Lepicolea Dum. Recueil Obs. Jungerm. 20. 1835. i . 105 Key to the Falkland Island Species of Lepicolea Leaf segments ending in a uniseriate tip of a few to several elongate cells, the second, third and fourth cell from tip elongate ...... . ....... A. ochroleuca . Leaf segments ending in a uniserate tip of a flag-celled row of short (subquadrate to short-rectangular) cells which are often caducous; the third and fourth cell from the tip short. Vittae commonly present . . . . L. rigida Lepicolea ochroleuca (Spreng.) Spruce Jungermannia ochroleuca Spreng. Syst. Veget. 4 (2): 325. 1827. Sendtnera ochroleuca (Spreng.) Nees _i_r_1_ G. L. 8N. 240. 1845. @eroma ochroleuca (Spreng.) Mitt. illHook. f. Handb. N. Z. Flora. 754. 1867. Herbertia ochroleuca Trev. Memorie Ist. Lomb. Sci. Lett. III. 4: 397. 1877. Lepicolea ochroleuca (Spreng.) Spruce, Trans. Proc. Bot. Soc. Edinb. 15: 345. 1885. Original material: South Africa, Cape of Good Hope, ELSE—"$1” (NYI. STR - cited in Scott. 1960). Jungermannia hirsuta Nees _eA Hook. f. 8 Tayl. London Jour. Bot. 3: 289 (_ip errore pro 389), 475. 1844, m. M. §_e_hdtnera ochroleuca _B_ mexicana Gott. Kongel. Danske Vi- densk. Selsk. Naturvidensk. Math. Afh. II. 6:140. 1863, §_y__. _f_i_d_e_ Scott (1960). 106 Taxonomical remarks: This is the first authentic report of Lapicolea ochroleuca from the Falkland Islands, as all previous re- ports (G. L. 8 N. 1845 as Sendtnera, Hooker 8 Taylor 1844 and Taylor IHooker 1847b as Jungermannia hirsuta) were based upon a Hooker collection which is actually A. rigjgg. _E_cpl_ogy: This species is rare in the Falklands, where it occurred on wet rock walls at the summit of Mt. Kent, 455 111. Only afew "clones" were observed. The second and only other locality for the species was deep in a cool, cave-like hollow of a rock out- crop in a dwarf shrub heath. Here, as in the above local, the plants (again only a few) grew on an extensive, flattish area of the outcrop. The species is common in the wet Magellanian moorland, where it may form extensive mounds over bark. It also is quite common on M- fpgypbark in the Valdivian region. The species appears to require ahigh rainfall and cool temperatures, the absence of the former (as well as the bark habitat) presumably accounting for the rarity of the taxon in the Falklands). The above two sites apparently possessed the precise conditions for growth of the species; both were cool, and received moisture through rock seepage, the former was cool owing to altitude, the latter, 180 m. was cool owing to a particular niche. The Port Stanley region has cooler temperatures and a higher rainfall than other Falkland regions, a point which almost undoubtedly accounts for the presence of _L_. ochroleuca in this part of the Falklands. Phytogeography: This amphiatlantic species occurs in South Africa, Tristan da Cunha the Falklands, Tierra del Fuego, Patagonian 107 Channels, Valdivian region (West Patagonia north to 39° 16' 5., Andean Patagonia at P. N. Nahuel Huapi), Juan Fernandez and north in the Andes to Mexico. The report from Brazil in Stephani (1909) and India in Hooker (1867) are regarded as erroneous. The New Zealand reports are either A. attenuata or _L_. scolopendra, and that from Tasmania is likely A. scolopendra (fide citations in Scott 1960) (see Map 3). Literature Records (note: locality information is omitted for extra-southern South American collections): TIERRA DEL FUEGO: W- (Arnell 1953b, Hooker 1867 as Sendtnera, Ku'hnemann 1937, 1949, Mitten 1855 as Sendtnera, Stephani 1909); Albatross - Pto. Churruca (Evans 1892); M - Pto. Angosto (Fulford l963b, Scott 1960, Stephani 1901a), R. Azopardo Stephani 1901a); GaSperi - M. Sarmiento, I. Laberinto (Gala 1923); A. _B_. Hatcher - Ens. Villarino (Evans 1898); _H_o_gl_<_e_r_ - I. Hermite and/or Cabo de Hornos (Fulford l963b, G. L. 8 N. 1845 as Sendtnera, Hooker 8 Taylor 1844 8 Taylor 8Hooker 1847b as _J_. AM, Scott 1960); Naumann - B. Tuesday (Schiffner 1890 as Sendtnera); Pennington - Ens. Villarino 8 Ushuaia (Dusén 1905); Racovitza - I. Clarence (actually I. Capitan Arencena) (Stephani 1901); Savatier - Pto. Churruca (Bescherelle 8 Massalongo 1889 as Leperoma); Skottsberg 8 Halle - R. Fontaine, Cta. Gomez (Stephani 1911), Pto. Cook (Stephani 1905a); Spegazzini - I. Basket, M. Sarmiento, Pen. Brecknock 8 B. Slogget (Massalongo 1885 as M). PATAGONIA: Anonymous (Kiihnemann 1937, 1949 Stephani 1909), Valdivia, I. Chiloe (Gay 1852); Albatross - Pto. Mayne (Evans 1892); £91199. - I. Chiloe (Fulford l963b, Scott 1960); Cunningham - Pto. Gallant, 108 a. Sholl, a. Halt (Fulford 1963b, Scott 1960); pgng - I. Newton (Stephani 1900), R. Aisen, I. Guaitecas, L. Llanquihue, L. Nahuel Huapi (Puerto Blest) (Fulford 1963b, Scott 1960, Stephani 1900); m- Canal Smyth (Scott 1960); @111 - Valdivia (Fulford l963b, Scott 1960); Hollermayer - Panguipulli (Reimers 1926); flossemg - Corral, Pucon, Petrohue (Herzog 1938); Lechler - Rada York (Fulford 1963b, Scott 1960); Santesson - R. Aisen, Pto. Montt, Valdivia, R. Enco, Enco, Rinihue, C. Tralcan (Arnell 1955), Rinihue, Enco (Miller 1955); Savatier - Pto. Bueno, Pto. Eden, I. Madre de Dios (Besche- relle 8 Massalongo 1889 as Leperoma); Schwabe - Calbuco, R. Puelo (Herzog 1939); Pto. Puyuhuapi 8 Termas de Puyehue (Herzog 1954), L. Pellaifa, Huitrapulli-Aleucapi, Chaihuin-R. Colun, C. Lungoico (Her- zog 1960); Skottsberg 8 M - Pto. Cutter, 1. Atalaya, I. Pacheco, F. Peel, Cta. Rayo, Cta. Connor, 8. Chacabuco, I. de San Pedro, Ancud (Stephani 1911); Sleumer - Pen. Quetrihue (=Pen. Beatriz), (Milller1955);IAax__t_eL - Corral (Fulford l963b); Thomasson - L. Villarrica (Thomasson 1963). JUAN FERNANDEZ. MAS A TIERRA: mgflyggl (Crandall 1969); Kunckel (Arnell 1957); Skottsberg (Arnell 1957, Fulford 1963b, Scott 1960). MAS AFUERA: Skottsberg (Arnell 1957, Fulford 1963b, Herzog 1942, Scott 1960, Stephani 1911)... BOLIVIA: My (Spruce 1890). HONDURAS: Standley (Fulford 1963b). GUATEMALA: Steyermark(Fulford1963b). MEXICO: Anonymous (Fulford l963b, G. L. 8 N. 1845 8 Hooker 1867 8 Mitten 1855 as Sendtnera, Scott 1960). BRAZIL: Anonymous (Stephani 1909). TRISTAN DA CUNHA: Christophersen _8_Mejland (Arnell 1958); Wace (Arnell 1958). I 1-.cI- an «I Inc. '-v. v 109 INACCESSIBLE: Christophersen §_Me,jland (ArnelT 1958). SOUTH AFRICA: Anonmous - (Arnell 1963, Hooker 1867 & Mitten 1855 as Sendtnera, Sim 1926, Stephani 1909); Eaton (Mitten 1877 as Legeroma); Eaton, Ecklon, Marloth, Milne, Rehmann (Scott 1960). CAMPBELL: Anonymous - (Mitten 1855 as Sendtnera); Hooker; (Hooker 1867 as Sendtnera, Taylor & Hooker 1847a as g. m). AUCKLAND: m (o. L. 8. N. 1845 as .5393.- ne_ra). NEW ZEALAND. SOUTH ISLAND: Buchanan, Hector, Lyall (Hooker 1867 as Sendtnera); L_y_a_11 (Mitten 1855 as Sendtnera). TASMANIA: MM (Rodway 1916). JAVA: Anonmous - (G. L. & N. 1845 & Mitten 1855 as Sendtnera). INDIA: Anonymous - (Hooker 1867 as Sendtnera) . FALKLAND SPECIMENS SEEN: EAST FALKLANDS: STANLEY REGION, Goat Ridge, c. 180 m. (_3_1_9_5_); summit of Mt. Kent, 455 m. (373g & 223.2,). ADDITIONAL SPECIMENS SEEN: CHILE. MAGALLANES: Cabo de Hornos, sin. coll. Um) as Jungermannia hirsuta (MICH); I. Deso- lacion, Pto. Angosto, 26 March 1896 99323216 (NY); Canal Smyth, Jan- uary1924 Gusinde 3697 (Cryptogamae exsiccatae editae a Museo Hist. Natur. Vindobonensi, det. Herzog) (NY); B. Sholl, March 1868 SHELTER" 113111332 (NY); Pto. Mayne, 5 February 1888 _Al_b_a_g_o_s_§__.p_. (BM, NY); 8. Halt, Cunningham §_.p_. (NY). CHILOE: I. Guaitecas, May 1897 M $.11. (BM). VALDIVIA: Corral, Thaxter 116, 119 - c. coel. + cap., BE. l_4_§_ (MICH); Arique, Lechler 639 _13 (NY). * ._ _. .. .,t. .. ..... .. . . .. . .. u v .a ..u .. u ... .... n .- vol .- no.» u .n -.u N . e .. u . ..- . . _ n. x; :. ..... . . - w. v... _ Z . 110 Lepicolea rigida (De Not.) Scott Sendtnera rigida De Not. Memorie Accad. Sci. Torino II. 16: 229. pl. XV,_1_-_9_1855. Herbertia rigida (De Not.) Trev. Memorie Ist. Lomb. Sci. Lett. III. 4: 397. 1877. Lgeroma rigida (De Not.) Mass. Nuovo G. Bot. Ital. 17: 252. 1885. Lepicolea MWe Not.) Scott, Nova Hedwigia 2 (1-3): 148. 1960. Original material: Chile, "Valparaiso," P_ug_c_i_o_ (NYI). Lepicolea seriata Herz. Hedwigia 66: 91. f. 8. 1926, EL. fide Scott (1960). Original material: Chile, Prov. Aisen, Pta. Leopardos, Reichert a Hicken s.n. (JE) - cited in Scott (1960). Ecology: See comments for the ecology of L. ochroleuca, which also apply here. Hooker visited only East Falkland and it is likely that his collection was made in a niche similar to those of L. ochroleuca. Phytogeograghy: This species occurs in the Falklands, Tierra del Fuego, Patagonian Channels and Valdivian region north to 46° 32’ 8.. 73° 52‘ w. (Pta. Leopardos) (see Map 4). Literature Records: TIERRA DEL FUEGO: g. B. Hatcher - Lapataia (Crandall 1969, Fulford 1963b, Scott 1960), Ens. Villarino (Fulford 1963b, Scott 1960); Hooker - Cabo de Hornos, I. Hermite (Ful- ford l963b, Scott 1960); Spegazzini - I. de los Estados (Fulford l963b), Scott 1960). PATAGONIA: Cunningham - Pto. Gallant, Pto. Island (Fulford l963b, Scott 1960); 993! - Strait of Magellan (Fulford l963b, Scott 1960); Dusén - I. Newton (Fulford 1963b. Scott 1960); 111 Gusinde - Canal Smyth (Fulford 1963b); 9_. §_. Hatcher - Patagonia (Scott 1960); Hicken - Pta. Leopardos (Fulford 1963b), Pillwax - Strait of Magellan (Fulford 1963b, Scott 1960), Reichert - Pto. Leopardos (Fulford 1963b). FALKLAND SPECIMENS SEEN: EAST FALKLANDS: 1843 Hooker 3.1. as Jungermannia hirsuta "with a Frullania allied to Jung. lobulata Hook." (FH). ADDITIONAL SPECIMENS SEEN: CHILE. MAGALLANES: Cabo de Hornos, Hooker s_.n. as Sendtnera ochroleuca (NY). ARGENTINA. TIERRA DEL FUEGO: I. de los Estados, 2 February 1882 Spegazzini 31 as Sendt- mochroleuca (NY). CHILE. MAGALLANES: Ensenado Villarino, J. B. Hatcher _3_, 1_3_as L_. ochroleuca (MICH); Cta. Playa Parda, Cunningham Mas 1;. ochroleuca (BM); "Patagonia," J. B. 1191:3113: lg. l_3_. :33, 1; as L_. ochroleuca (NY); Pto. Bueno, May 1896 Dusen 3.1. as I_._. ochro- leuca (NY). AISEN: Pto. Island, Cunningham s.n. (NY). LEPIDOLAENACEAE Nakai i_n_Ogura et al. List Prof. Nakai's Pap. 200. 1943. Gackstroemia Trev. Memorie Ist. Lomb. Sci. Lett. III. 4: 397. 1877. 112 Key to Falkland Island Species of Gackstroemia l. Dorsa1 lobes of main axis armed and auriculate at anti- cal base; cells of dorsal lobes with large knot-like trigones and thickened walls; dorsal lobes of branches with 6-8 cilia; lobules of main axis involuted, widely ovate in shape, often with decurved margins, not inflated .................... g. magellanica 1. Dorsa1 lobes of main axis not armed or auriculate at antical base; cells of dorsal lobes with small tri- gones and thickened walls; dorsal lobes of branches mostly entire; lobules of main axis inflated . . . g. patagonica Gackstroemia magellanica (Lam.) Trev. Jungermannia magellanica Lam. Encycl. Method. Bot. 3: 284. 1789, 321 Jungermannia magellanica Spreng. Annln Wetter. Ges. Naturk. 1 (1): 25. 1809 (= Frullania fide Grolle, 1967). Polyotus magellanicus (Lam.) Gott. _i_nG. L. & N. Syn. Hep. 248. 1845. Gackstroemia magellanica (Lam.) Trev. Memorie Ist. Lomb. Sci. Lett. III. 4: 397. 1877. Lepidolaena magellanica (Lam.) Evans, Contr. U.S. Natn Herb. 1: 140. 1892. Lectotype (_C_f. Grolle 1967): Chile, Prov. Magallanes, Strait of Magellan, Commerson (PC - MM). Mdecipiens Goeb. _e_x_ De Not. Memorie Accad. Sci. Torino II. 15: 223, 1355, 19E!- M” 112.51% Polyotus deciiiens Goeb. Ann. Jard. Bot. Buitzenzorg I. 7: 30. _f_. _2_3_. 1888, gm. fide Grolle (1967). 11.3 Lepidolaena hallei Steph. K. Svenska VetenskAkad Handl. 46 (9): 74. _f. 29911. 1911, syg fide, Grolle (1967), 5911 Lepidolaena halle- aflSteph. Sp. Hep. 6: 370. 1923 ( = _L menziesii fide Grolle, 1967). Lectotype (2:. Grolle 1967): Falkland Is... Mt. Usborne, Meson (UPS) - 292.219.: Lepidolaena skottsbergii Steph. Sp. Hep. 6: 371. 1923, _S_yg _fidg Grolle (1967), non Lepidolaena Skottsbergii Steph. K. Svenska VetenskAkad. Handl. 46 (9): 76. 1911 (= L_. menziesii fide Grolle, 1967). Original material: Falkland 15., 1909 Skotts- £133.11. (G) - cited in Grolle (1967). Ecology: This species is common and exhibits a wide ecolog- ical tolerance in the Falklands as well as southern South America. In the Falklands it is often encountered on soil and old, rotted cushion plants in dwarf shrub heaths. It is also common on soil, rock orAs_t_el_i_g cushions of sheltered high altitude cliffs. I fur- ther encountered it in a moist feldmark (700 m.). Phytogeography: This Species occurs in the Falklands, Tierra del Fuego (widespread). Patagonian Channels, Valdivian region (West Patagonia north to 39° 52' S.) and Juan Fernandez. Australasian records of the species were shown by Grolle (1967) to be plants of 9.. WISE}. (see Map 5). Literature Records (FALKLANDS): Anonymous - (Herzog 1926 as _L_. Skottsbergii, KUhnemann 1937 8: 1949 as Lepidolaena 8. _|:_. hallei); Mg- Mt. Usborne (Grolle 1967); Skottsberg - Mt. Usborne (Grolle 1967); Skottsberg 8: Halle - Mt. Usborne (Stephani 1911 as Lepido- 199931.11edde1 Is. (Grolle1967). ‘7".- I. . I ‘ 1 o. a I I l u 1.. I u i l. I. . II a. h v D u. o. . II 0 7 TO . l r - Q .. t . I 1 u I n . a . v u - «1|. .1 . I . Q I n I IV I o. .o . . v .1. .o. \a I. - u - o u . _..\ u . \ C . 1 ,7 114 Additional Literature Records (note: locality information has been omitted for non-American sector records): TIERRA DEL FUEGO: Anonmou - Strait of Magellan (Stephani 1909 as Lepidolaena), M. Forster (Grolle 1967, Hooker 1867 as Polyotus, Rodway 1916 as Legisl- o_l_ae_n_a_, Schwaegrichen 1814 as Jungermannia), Kiihnemann 1937 8 1949 as gagidolaena); Dusen - R. Azopardo (Grolle 1967, Stephani 1901a as Lepidolaena), Pto. Angosto (Grolle 1967); GaSperi - B. Angelito, S. Agostini, M. Sarmiento, Valle delle Fate, I. Laberinto (Gola 1923 as Lepidolaena); J. E: Hatcher - Fuegia (Evans 1898 as Legidolaena); Hooker - I. Hermite and/or Cabo de Hornos (G. L. 8 N. 1845 as Poly- o_t_u§, Hooker 8 Taylor 1844 8 Taylor 8 Hooker 1847b as Jungermannia Lechler - Rada York (Grolle 1967); Menzies - I. de los Estados (G. L. 811. 1845 as Polyotus, Grolle 1967, Hodgson 1959 as LeLidolaena, Hooker 1818 as Jungermannia, Taylor 8 Hooker 1847b as Jungermannia, Montagne 1845 as Polyotus); Naumann - B. Tuesday (Schiffner 1890 as Polyotus); Racovitza - I. Clarence (actually I. Capitan Arecena) (Stephani 1901b as Lepidolaena); Santesson - Canasaca (Arnell 1955 as gagidolaena); Savatier - I Hermite, I. Hoste, I. Clarence (actu-— ally I. Capitan Arecena), Pto. Churruca (Bescherelle 8 Massalongo 1889 as Polyotus); Skottsberg - Pto. Cook, 8. Tekenika (Stephani 1905a as Lepidolaena); Skottsberg 81m - R. Fontaine (Stephani 1911 as Lepidolaena); Spegazzini - I. de los Estados (Massalongo 1927 as Lepidolaena), I. Hoste, M. Sarmiento, Pto. Vancouver, M. Buenos Aires, B. Blossom, Penguin Rookery (Massalongo 1885 as Polyotus). PATAGONIA: Anonymous (Kil'hnemann 1937 8 1949 as Lepidolaena); Alba- £[9§_§,- Strait of Magellan (Evans 1892 as Lepidolaena); Andersson - 115 Pto. del Hambre (Angstrom 1872 as Jungermannia, Grolle 1967); Comner- .591‘ Strait of Magellan (Gay 1852 as Polyotus, Grolle 1967, G. L. 8 N. 1845 as Polyotus, Hodgson 1959 as Lepidolaena, Hooker 1818 as Jungermannia, Taylor 8 Hooker 1847b as Juggermannia, Montagne 1845 as Polyotus); Dumont d'Urville - Pto. Gallant (Montagne 1845 as Poly- gt_ug), Strait of Magellan (Gay 1852 as Polyotus); DuSén - I. Newton, Pto. Bueno, R. Aisen, Puerto Blest (Stephani 1900 as Lepidolaena), 1. Newton (Grolle 1967); Hollermayer - V. Villarrica (Reimers 1926 as gagidolaena); Jacquinot - Pto. Gallant (Montagne 1845 as Polyotus); m- Corral (Grolle 1967); Kgbitski - Cord. Pelada (Grolle 1967); N_eg_e_r- western Slope of Andes - 39° - 39° 30' S. (Neger 1899 as Qgidolaena); Nobl - B. Borja (Grolle 1967); Santesson - C. Mina Rica (Miiller 1955 as gpidolaena), R. Rubens, Tres Puentes, C. Mina Rica (Arnell 1955), Punta Arenas (Grolle 1967); Savatier - Pto. Bueno. Pto. Eden, I. Madre de Dios, Pto. Barroso (Bescherelle 3' 143558101190 1889 as Polyotus); M - Istmo de quui (Herzog 1954 as £93110; 1.3.9111). C. Tesoro (Herzog 1939, 1954 as Lepidolaena), Pto. Puyuhuapi (Grolle 1957); Skottsberg - Pto. Cuarenta Dias, I. Chiloe (Grolle 1967135&tt5berg J Halle - Pto. Cutter, 1. Atalaya, F. Peel, L- Azara, Cta. Rayo, I. de San Pedro (Stephani 1911 as Lepidolaena), 3. Skyring (Grolle 1967); Spegazzini - R. Santa Cruz (Massalongo 1906 as 101%); Poyo - Huapi, R. Santa Cruz (Massalongo 19271. JUAN FERNANoEz. MAS A TIERRA: Mk3]. - near summit of El Yunque - ca. 900 m. (Arnell 1957 as Lepidolaena). MAS AFUERA: Skottsberg - Camp C0"?”ESpondenC1‘a - ca. 1350 m. (Grolle 1967, Herzog 1942 as Lepidolaena). - .- " . A o .- _—- ‘ ‘ I .. r“ . v . I a - o . y o I h I . . I. 'E .. y; .. '~ .. ‘ . ‘ _. m ' ~.. ~ . \ .-.. ...l ‘ ' a 'o . . o. .u \ .~¢‘_‘ ”u .. ‘I. t... u \ "\ ‘. ‘n c 116 CENTRAL CHILE: £93ng - "Valparaiso" (De Notaris 1855 as Polyotus). CAMPBELL: Hgglger; (G. L. 8 N. 1845 as Polyotus, Hodgson 1959 as Lepidolaena, Hooker 1867 as Polygth, Hooker 8 Taylor 1844 as M99}: 992111)- NEN ZEALAND: Anonymous (Stephani 1909 as Lepidolaena). STENARD ISLAND (NEW ZEALAND): Kirk, Martin (Hodgson 1959 as M- 11933); Ly_a_Il (Martin 1949). SOUTH ISLAND (NEW ZEALAND): Du Rietz, Kirk, Macdonald, Martin, M (Hodgson 1959 as Lepidolaena); LleJ (Hooker 1867 & Mitten 1855 as Polyotus). NORTH ISLAND (NEW ZEALAND): m, Colenso, Hodgson, Mossman (Hodgson 1959 aS Lepidolaena); m (Hooker 1867 8 Mitten 1855 as PM). TASMANIA: Aggy- M- (Hodgson 1959 as Lepidolaena, Hooker 1867 as Polyotus, Rodway 1916 as L_epidolaena, Stephani 1909 as Lepidolaena); Bastow, Rodway (Carrington 8 Pearson 1888 as Polyotus); Hooker (G. L. 8 N. 1845 as m. Hooker 8 Taylor 1844 as Jungermannia); Archer, Gunn, 01d- fieLd (Mitten 1860 as Polyotus); Weymouth - (Pearson 1924 as Lepido- Em). AUSTRALIA: Anonymous (Hooker 1867 as Polyotus, Stephani 1909 as BPidolaeLa); Cunningham (G. L. 8 N. as Pol otu_S_. Hodgson 1959 as L_epidolaena). FALKLAND SPECIMENS SEEN: EAST FALKLANDS: STANLEY REGION: surmn't ridge of north peak of Two Sisters, 245-290 m. (2.7.1.9 5.); summit of Mt. Kent, 455 m. (2151 & gggi B). MT. USBORNE REGION. gap between Mt. Usborne 2 and Table Rock, c. 440 m. (2633 5); south“ east slope of Mt. Usborne 2, c. 455 m. (_2_5___9_fl_ A); ridge between Mt“ Usbornes 1 8 2, 685 m. (£532 A, _2_59_§_ 8. _2_§_5_3__A_); summit of Mt. Us- borne 1, c. 700 m. (mg 3. gig), WEST FALKLANDS: PORT HOWARD, .r s. o , .. . . O p ‘l o ,. . u ‘ II I 117 Freezer Rocks, east slope of Mt. Maria, 320 m. (£1111); pass south- west of Mt. Maria summit, c. 610 m. (_e_ogg g, 3092, 3113); MT. ADAM, ridge south of northern Take, 610 m. (3024 _B-c. perigyn.). WEDDELL ISLAND, rock dome on summit of peak northeast of Mt. Weddell, 335 m. 199.13.96.98 9999.8). ADDITIONAL SPECIMENS SEEN (representative selection): CHILE. MAGALLANES: Cabo de Hornos (M95 (NY); I. Hermite, 111915311th 1. Lennox, 19 March 1900 m (FH, NY); R. Azopardo, 3 March 1896 L881]1_4 (NY); I. Desolacion, B. Tuesday, 2 February 1876 W (FH); B. Ventisqueros, _H_i_1_1_ (FH); B. Isthmus, Savatier (FH); I. New- ton. May 1896 D_us1e_n_ (FH, NY); Pto. Bueno, May 1896 mg (NY- perigyn.); Pto. Grappler, 1893 Douglas (FH); B. Halt Cunningham (NY). AISEN: R. Aisen, January 1897 ggseg (NY). Gackstroemia patagonica (Steph.) Grolle WMagonica Steph. K. Svenska VetenskAkad. Handl. 46 (9): 76. f. 29, e-h. 1911. Gackstroemia patagonica (Steph.) Grolle, J. Hattori Bot. Lab. 30: 14. 1967. Lectotype (9:. Grolle. 1967): Chile, Prov. Magallanes, Canal Gajardo, Ha__ll__e_8 Skottsberg 3.11- (UPS - 993 vidi). M913 This species is very rare in the Falklands and was c011ected but once, at 455 m. in a stream on a slope (southeast) in a W heath. In the Magellanian moorland this species is charac- teriStic of non-forested areas of cushion plants, scattered rocky out- "095 and scattered small pools. 7 . ‘ . I l 5,. . c t —- u .. o. A 118 Phytogeogapgy: This taxon occurs in the Falklands, Tierra del Fuego (R. Azopardo and Pto. Angosto) and southern Patagonian Channels (Canal Gajardo). Literature Regorts: TIERRA DEL FUEGO: Anonym ou_s_ - (Stephani 1923 as Lepidolaena); Dusén - R. Azopardo, Pto. Angosto (Grolle 1967); Skottsberg Jflaflg - R. Azopardo (Grolle 1967). PATAGONIA: Skottsberg 8M - Canal Gajardo (Grolle 1967). FALKLAND SPECIMEN SEEN: EAST FALKLANDS: MT. USBORNE REGION, southeast SlOpe of Mt. Usborne 2, co 455 m. (2_6__1_§_)o ADDITIONAL SPECIMENS SEEN: CHILE. MAGALLANES: FUEGIA, M1133 as Lepidolaena reticulata (NY); R. Azopardo, 600 m., 9 March 1896 Dusen L2; as L. hariotiana (NY); I. Desolacion, Dusen _l_3_7_ as L. hariotiana (FH). Without definite location, Lechler (NY). Lepidolaenaecae Species Excluded from Falklands GLCkstroemia hariotiana (Besch. 8 Mass.) Grolle. This taxon was reported from the Falklands by Stephani (1911 as Lepidolaena). 910119 (1967) did not report the species from the Falklands, and I have searched for the collection in numerous herbaria without success. 1 am at least temporarily excluding the species from the Falkland flora. EPidolaena reticulata (Hook. f. 8 Tayl.) TY‘GV- This 599C165 was "ePOrted for the Falklands by Stephani (1911) and KUhnemann (1937, 1949) Cited the Stephani paper in his catalogues. Grolle (1957: p. 47) .-t‘ 4.. . n .0. '0‘ re- 7 ;.: . ’—b—- so I 'I . r. . . ‘ n ‘ \ 119 states the specimen on which the Falkland report was based is Gack- stroemia magellanica. Lepidolaena reticulata is found on Auckland Island, Campbell Island, New Zealand and Tasmania. LEPIDOZIACEAE Limpr.i_r1Cohn, Kryptogamenfl. Schles. 310. 1875. Hyalolepidozia S. Arnell eyGrolle, Revue Bryol. Lichen. 32: 179. 1963. Hyalolepidozia bicuspidata (Mass.) S . ArneTTe_x GroTTe ggidozia bicuspidata Mass. Nuovo G. Bot. Ital. 17: 239. pl. 22. f. Z_5_. 1885. Hyalolegidozia bicuspidata (Mass.) S. Arnell, Bot. Notiser 115: 213. 1962, nom. illeg. Paracromastigum bicuspi- 93111 (Mass.) Schust. J. Hattori Bot. Lab. 26: 276. 1963. Hyalolepidozia bicuspidata (Mass.) S. Arnell gyGrolle, Revue Bryol. Lichen. 32: 179. 1963. Original material: Argentina, Terr. Tierra del Fuego, I. de los Estados, Pto. Cook, Spegazzini 8.5.9 (G) - cited in Fulford (1968). 120 Taxonomical remarks: After a thorough survey of branch types in the Falkland material of H. bicuspidata, I found both lateral- and ventral-intercalary branches in about equal frequency, and only a single occurrence of a Frullania-type branch. I was unable to find Microlepidozia-type branching. Terminal branching is only rarely de- veloped in the genus; in H. bicuspidata Grolle (l963b) found two Frullania-type branches (Chilean material) and one Microlegidozia- type branch (South African material). The branching pattern in Molepidozia readily distinguishes the genus from the closley re- lated genera Bonneria and Paracromastigum. In Bonneria the branch- ing is plastic with Acromastigum-, Frullania-, Microlepidozia- and ventral-intercalary branches (with lateral-intercalary branches absent). ParacromastimIm has Acromastigum- and Frullania-type branches and rarely ventral intercalary branches (with Microlepi- dozia- _a_nJ lateral-intercalary branches absent). See Schuster (1965a) for a discussion of the evolution and relationships within this com- plex of genera. Fulford (1968, p. 277), in her generic key, utilizes as her primary key character for Hyalolqaidozia, "underleaves and leaves similar in size and form." This statement is presumably based upon examination of the type, although the ventral view of the stem (plate 101, fig. lb) is from non-type material. Massalongo (1885) in his original description of "Lepidozia" bicuspidata states, "Amphigastria fol. 2-4-plo minora . . ." and his figures clearly Show the underleaves as considerably Smaller than the leaves. The 121 underleaves of the Falkland material of H. bicuspidata are ca. 1/3-1/2 the leaf size and are never equal to the size of the leaves. M: This taxon was collected in dwarf shrub heaths where it was found on soil particularly on soil layers over rock. Phytogeography: This amphiatlantic temperate species is known from South Africa (Table Mt.), southern Patagonia (R. Rubens), Pata- gonian Channels, the Valdivian region north to 41° 46' S. (West Pata- gonia), Juan Fernandez and "Frai Jorge" (see Map 6). Literature Records: TIERRA DEL FUEGO: Anonymous - (Kiihnemann 1937:1949 as Lepidozia); Ousén - R. Azopardo (Fulford 1968); m- _z_i_n_i_- Pto. Cook (Fulford 1968, Massalongo 1927 as Lepidozia), Fuegia (Stephani 1909 as Lepidozia), M. Richardson (Fulford 1968). PATAGONIA: MM“ (Grolle 1969); Santesson - R. Rubens (Arne11 1955 as LgLi- 19_z_i_a_); M- Calbuco (Grolle 1963b, Herzog 1939 as Lepidozia , I. Puluqui (Fulford 1968), I. Magdalena, Pto. Puyuhuapi (Herzog 1954 as M). JUAN FERNANDEZ. MAS A TIERRA: Skottsberg - between Piedra Agujeriada 8 Laura (Herzog 1942 as Lepidozia). CENTRAL CHILE: 19111311- "Frai Jorge" (Herzog 1954 as Lepidozia). TRISTAN DA CUNHA: Ebristophersen 113,111.31 - zoo m. a 550-600 m. (Arnell 1958 as 1.921. Mia). SOUTH AFRICA. CAPE PROV.: Anonymous - Table Mt., Disa Gorge (Arnell 1963), Arm-311 - Table Mt. (Grolle 1863b). FALKLAND SPECIMENS SEEN: EAST FALKLANDS: PORT WILLIAM REGION: summit ridge of Mt. Low, c. 245 m. (_3_2_3_3_). STANLEY REGION, Sapper Hill, 135m. (1mg). MT. USBORNE REGION, southeast slope of Mt. Us- borne 2, c. 455 m. (gig-1;). NEST FALKLANDS: WESTPOINT ISLAND, steep Slope 8 cliffs facing The Woolly Gut (2914, 2924, 2931, 8 2912- (2. per.) ..,. ['2‘ ~, . I .5 I _ . v .‘ o '. It» .n u. ‘k T I .1:— 122 Hygrobiel lgasi s (Schust.) Engel, comb. nov. Hygrolembidium subg. Mrobiellopsis Schust. Nova Hedwigia 15: 467. 1968. Type: Hygrolembidi um isophmum Schust. Plants erect, perfectly isophyllous, trigonous, branching only of lateral and ventral intercalary type, stern cortical cells small, thin or slightly thickened, medullary cells larger, lepto- dermous, in many rows; leaves and underleaves closely imbricated, deeply concave, ovate or orbicular, apices plane or retuse, margins and apices with slime papillae, (l-) 2-3 stratose at least locally in median basal portion, cell walls thin or uniformly thickened, trigones absent. Perianths on Short intercalary branches, upper portion pluriplicate, mouth with Slime papillae. Seta of Lepidozia type, epidermal cells in 18-20 rows. Capsule walls 4-6 stratose. Spores red-brown in color, with numerous vermiculate, short to long, closely arranged, irregular, occasionally branched ridges, Elaters bispiral throughout or 3-4 spiraled in median portion. Hygrobiellcmsis isophyllum (Schust.) Engel, comb. nov. Hygrolembidium isophyllum Schust. Nova Hedwigia 15: 467. 21. _5_§_. 1968. Holotype: Argentina. Terr. Tierra del Fuego, C. Gari- baldi, near L. Escondido, Schuster 58319e (RMS 1). 123 Taxonomical remarks: Schuster (1968b) reflected his uncer- tainty regarding the systematic position of Hygrolembidium isophyllum when he stated (p. 467), Allied to Isolembidium and possibly also to Hygrolembidium is another undescribed taxon which serves, in some ways, to connect Hygrolembidium and Isolembidium, yet differs from both taxa. The margins and apices of leaf and underleaf of Hygrobiel- lopsis isophyllum have several (8-20) slime papillae. The papillae are usually mounted directly on the margin but may be on the tip of a 1-several celled tooth. There is no mention of slime papillae or teeth in the original description or indication of their presence in the figures of the taxon in Schuster (1968b). I have, through the kindness of Dr. R. M. Schuster, studied the holotype material of: Mygrolembidium is_ophyllum and confirmed the presence of these struc- tures in the original material. Neither Isolembidium nor Hygrolem- 11'_di_u_n1 possess slime papillae, and the presence of this character in Hygrobiellogsis aids in the differentiation from the two genera. The perianth mouth, bract and bracteole margins and apices are likewise ornamented with slime papillae. The perianth mouth and bracts of Hygrolembidium andinum may have slime papillae, judging from the general appearance of the structures figured, but not described, in Herzog (1954, f. 8, e, h). The leaves are entire and without slime Papillae. I have not seen type material of this taxon. Hygrobiellgasis iSOphyllum was describe-d on the basis of com- pletely sterile material. The perianths of Hygrobiellopsis and Hm. lembidium compare somewhat in general shape. both being trigonous, 124 fusiform to elongate-ovate and contracted toward the mouth. Hm- biellopsis, however, is pluriplicate in the upper 1/2. Both Hygrolembidium and Hygrobiellopsis possess the Lepidozia type seta anatomy (8-16 (-20) rows of large epidermal cells surround- ing an indefinite number of interior rows of cells). While 11ng- lembidium isodictyon has 17 rows of epidermal cells and fl. stereophyl- 19116 rows (fide Herzog, 1951 figs. 7 and 9 respectively). Hygro- biellopsis isophyllum has 18-20 rows. The capsule anatomy differs somewhat in the two genera with bistratose walls in Hygrolembidium (jige description of _l_i_. stereophyllum in Herzog. 1951), and Hygyg- biellopsis with 4—6 layers. Schuster (1968b) stated the leaves of Hygrobiellopsis isophyl- Mwere unistratose, and utilized this character in differentiating it from Isolembidium and Hygrolembidium. I have found the leaves of Hygrobiellopsis to be 2-3 stratose in the median basal portion, with marginal areas unistratose. The leaves, however, may have poly- stratose regions scattered among areas of a single cell layer. The possession of at least locally polystratose leaves aids in the estab- lishing the rather close relationship of this taxon to both Hygrolem- bidium and Isolenbidium. Hygrobiellopsis shares several critical features with Hygro- lembidium and Isolembidium and indeed may be said to "connect" the tNO taxa. _H. isophyllum has the isophylly and polystratose, deeply concave leaves of Isolembidium but differs in (a) stem cells thin walled throughout, without a hyaloderm but with a cortex of small 125 cells, (b) leaf margins with slime papillae, (c) cells thin to uni- formly thickened, with trigones absent, and (d) the exclusively in- tercalary branching. Like Hygrolembidium, Hygrobiellopsis has poly- stratose leaves, exclusively intercalary branching, and a Lepidozia type seta anatomy, but the latter differs in (a) the small celled cortex with an absence of a hyaloderm, (b) the perfect isophylly, (c) the presence of slime papillae on the leaves, (d) the pluripli- cate (upper 1/2) perianth, and (e) the 4-6 stratose capsule walls. I have studied fertile material (9) of this Species from the Brunswick Peninsula (Engel 1982--c. per. + cap.), and from near Puerto Natales (_H_a_t_c_heL H. 1__5-_1_§_-C. per.). On the basis of this material I am providing below, a detailed description of the gynoecium and sporo- phyte. Plants dioicous; androecia not seen. Gynoecia on short inter- calary branches; bracts and bracteoles in 3 series, bracts of inner- most series deeply concave to subnavicularis, ovate, margins and especially apices with Slime papillae, bracts 2 cell layers thick in basal region, marginal 2-6 rows unistratose, cutical very finely papillose; bracteoles concave, not nearly as concave as bracts, broadly ovate, apex broadly rounded, margins and apices with slime papillae; perianth 1/4 - 1/2 exerted beyond bracts, fusiform to wide ovate, contracted toward the mouth, obscurely to distinctly trigonous in lower 1/2, keels, when present, A rounded, not sharp, upper 1/2 Pluriplicate, basal portion of perianth 2 cell layers thick, 3 layers thick only very locally, median portion mostly unistratose, but with ‘u on. 126 scattered regions of 2 cell layers, upper 1/3 unistratose throughout, perianth mouth with several small lobes, with slime papillae, sub- apical perianth cells irregularly rectangular, 52-77 u long, 20-26 u wide. Seta of the Lepidozia type, 10 cells in diameter, epidermal cells in 18-20 very regular longitudinal rows, the cells 52-62 1.1 in diameter, finely papillose, interior cells indefinite in number, (26-) 33-48 1: in diameter, corners slightly thickened. Capsule broadly ovate, 1.05 mm long, 0.86 mm wide, capsule wall 52-59 p thick, of 4-6 layers, the outer layer of large cells and the inner 3-5 rows of smaller cells of approximately equal size, outer layer of cells regularly to irregularly rectangular or subsquarrose in shape, 39-60 u long, 13-26 p wide, 18-26 L: thick, exposed wall smooth, outer layer with radial walls with small to quite large red-brown nodular thicken- ings which have been interconnected by sheetlike thickenings, the thickenings strictly I-Shaped or feebly extending onto outer tangen- tial wall, radial walls occasionally thin walled and without thicken- ings; inner layer of cells irregularly rectangular in shape, 55-79 11 long, 10-22 I: wide, 7-12 p thick, exposed wall smooth, inner wall with usually incomplete semiannular bands or with nodular extensions of various lengths from sparingly developed swellings on the radial walls, rarely with complete semiannular band, swellings often without bands or extensions, radial walls occasionally without thickenings; intermediate cell layers with or without radial thickenings, thicken- ings occasionally extending slightly on to outer tangential walls. 127 Spores red-brown in color, globose or broadly ovate, covered with numerous vermiculate, short to long, closely arranged, irregular, occasionally branched ridges, Spores 17-21 (I. Elaters usually bi- spiral throughout, occasionally 3 or 4 spiraled in median portion and bispiraled only at tips, spirals yellow-brown, elaters gradually tapering at both ends, usually long and Slender, 126-169 u long, 7.8-11.7 0 wide, occasionally shorter and stouter and to 16.9 11 wide. Ecology-Phytogeography: In the Falklands this species seems tote restricted to sheltered high altitude cliffs, where it grows on soil under rock overhangs or on Azorella cushions. In the Brun- swick Peninsula I found the Species only in the relatively dry eastern 911d (8 km. west of Punta Arenas, 305-610 m.), and here it grew on soil among cushion plants. Raymond E. Hatcher made several collections of the taxon between Punta Arenas and Puerto Natales, ca. 100 km. north of the Strait of Magellan, a locality which is on the eastern side of the Andes and thus relatively dry. The only other locality for the Plant is in the mountainous region of Isla Grande de Tierra del Fuego. 0n the mainland this taxon seems to be restricted to deciduous W forests of magellanian South America (see Map 7). FALKLAND SPECIMENS SEEN: EAST FALKLANDS: MT. USBORNE REGION. I‘ldge between Mt. Usbornes 1 8 2, 685 m. (_2_5_2_2_. _2_§_5,5_1- WEST FALK‘ LANDS: PORT HOWARD, pass southwest of Mt. Maria summit, c. 610 m. (112). MT. ADAM, east Side of summit ridge 670-700 m. (35818); summit of southermost peak 685 m. (2991). 128 PLATE 1 Hygrobiellopsis isophyllum (Schust.) Engel. Fig. l. Portion of plant with perianth, X 24. Fig. 2. Capsule with portion of seta, X 24. Figs. 3, 4, 5. Perianth, cross sections at ca. 1/4 distance from apex, middle, and base respectively, X 43. Fig. 6. Perianth mouth showing 5 slime papillae, X 205. Fig. 7. Elater, X 530. Fig. 8. Spore, X 530. Fig. 9. Innermost bract, cross section through base, X 105. Fig. 10. Inner cells of capsule wall, X 530. Fig. 11. Capsule wall, cross section, X 530. Fig. 12. Bract of innermost series, lateral view, X 24. Fig. 13. Bract apex of inner- most series showing a slime papilla, X 530. Fig. 14. Seta, cross section, X 105. Fig. 15. Outer cells of capsule wall, X 530. Fig. 1 from Hatcher H_15-13, Chile, Prov. Magallanes, Pto. Natales. Figs. 2-15 from Engel 1982, Chile, Prov. Magallanes, Brunswick Peninsula, 8 km. W of Punta Arenas. n . s u ..,. .. I n — -. e . 1 .. v 7. .... u‘ n. .' o. a o ‘I I I I - 130 Lepidozia (Dum.) Dum. Recueil Obs. Jungerm. 19. 1835. Pleuroschisma sect. Lepidozia Dum. Syll. Jungerm. 69. 1831. Key to the Falkland Island Species of Lepidozia l. Atleast some leaves and/or underleaves on well developed stems with marginal teeth-laciniae ....... L- chordulifera l. Margins of leaves and underleaves entire, very rarely with an isolated tooth ............. 2 2. Leaf segments in conspicuous pairs, the dorsal and ventral segments often reduced to a tooth of a few cells, leaves often distant, often scale-like; (only above 245 m. in Falklands, common in sheltered high altitude cliffs). . . . . . . . L_. fuegiensis 2. Leaf segments not in conspicuous pairs, dorsal and ventral segments larger, never reduced to a few cells, leaves imbricated, leafy, never scale-like; (rather common below 245 m. in Falklands, absent L. laevifolia in sheltered high altitude cliffs) ...... 131 Lepidozia chordulifera Tayl. Legidozia chordulifera Tayl. London Jour. Bot. 5: 371. 1846. Ju_n_- germannia chordulifera (Tayl.) Hook. f. 8 Tayl. _i_gHook. f. Bot. Ant. Voy. l (2): 442. 1847. Mastigophora chordulifera (Tayl.) Trev. Memorie Ist. Lomb. Sci. Lett. III. 4: 397. 1877. Orig- inal material: Chile, Prov. Aisen 8 Chile, Arch. de los Chonos, 1834 Darwin fl (FH-cited in Fulford 1966, NY!). L_egidozia hastata Steph. Sp. Hep. 3: 605. 1909, syn. fide Fulford (1966). Original material: Chile, Prov. Aisen, R. Aisen, Dusén 833 (G) - Cited in Fulford (1966). Lepidozia cuspidata Steph. K. Svenska VetenskAkad. Handl. 46 (9): 61. f. 23 a-b. 1911, syn. my. Original material: Western Patagonia, Skottsberg _S__.11. (G) - cited in Fulford (1966). L_epidozia effusa Steph. K. Svenska VetenskAkad. Handl. 46 (9): 62. 1.2151111. 1911, in. fig; Fulford 1966. Original material: Chile, Prov. Magallanes, Pto. Ramirez and I. Atalaya, Halle 8 Skottsberg (G) - cited in Fulford (1966). Lepidozia fernandeziensis Steph. K. Svenska VetenskAkad. Hand. 46 (9): 63. 1:. g_4__e. 1911,1 gm. f_i_dg Fulford (1966). Original material: Juan Fernandez, Mas a Tierra, near El Yunque, Skottsberg §_.n_. (G) - cited in Fulford (1966). Lepidozia hariotii Steph. Sp. Hep. 6: 329. 1922, _S_yg. _fj_d_e_ Fulford (1966). Original material: Chile. Prov. Magallanes, I. Hermite, Hariot (G, ex hb. Bescherelle) - cited in Fulford (1966). “_h IFulford (1966) states the description and figures of L. fernan- figfiysis have been interchanged with those of _l_._. M in Stephani 132 Lepidozia microscopica Steph. Sp. Hep. 6: 334. 1922, Eyg. fide Ful- ford (1966). Original material: Chile, Prov. Aisen, Cta. Hale, Dusen _s_._n_. (e) - cited in Fulford (1966). Lepidozia angu_lata Steph. g5. Fulf. Mem. N.Y. Bot. Gdn ll (2): 206. 1966.999- 999;: 919989 Taxonomical remarks: I cannot recognize Lepidozia cuspidata and must treat the species as a synonym of L. chordulifera. This treatment is based upon the study of some 123 collections of the L. ch_ordulifera - "cusgidata" complex from southern Chile, the Falkland Islands and South Georgia. The reasons for this synonymy are out- lined below. Taylor (1960, p. 110) states for Lepidozia cuspidata, The species is known only from the original collection. It is, however, readily distinguished by the broad leaves, very broad underleaves with rounded margins and the Spin- ose margins of both leaf and underleaf-laminas. Fulford (1966, p. 183) states for L. cuspidata in the key to Species, ". . . the lamina of the underleaves with bulging Sides often bearing several teeth; . . . ." I have found the underleaf curvature and armature highly variable, with several combinations of amounts of each. I have examined several specimens in which the underleaf bases vary considerably in curvature on a single stem axis. I have seen several specimens with 2-3 teeth on the dorsal, basal leaf margin and regularly produce underleaf marginal teeth. This condition grades into those specimens with a regular production of a single tooth on the dorsal leaf base and '3 regularly toothed underleaves, which 133 then phases into one in which the leaves and underleaves are occa- sionally toothed, but otherwise entire. Fulford (1966) records for the cell size of the segment bases of Lepidozia chordulifera 10-18 X 18 p, and for L. cuspidata 18-24 X 24 u, and includes the measurements in her key to aid in distinguish- ing the two taxa. I have made cell measurements of the basal segment cells of some 83 collections of the Lepidozia chordulifem-"cuspidata" complex and I find a steady continuum and wide range of cell sizes with no indication that L. chordulifera (3. Eli-1 has smaller cells, or _L_. "cuspidata" larger cells. The basal segment cell size of L. chordulifera (_S_. BIL.) is (lO-) 12-31 (-36) u long, and 10-26 (-33) 1.1 wide. Lc_gl_ogy: This species is very common in the Falklands, where it is able to tolerate, more so than any other Falkland hepatic, quite xeric, exposed conditions. It is very common in crevices and under overhangs of rock outcrops in dwarf Shrub heaths. In this association it is also frequently encountered under Blechnum magel- lgijLLm cover as well as on M cushions. The species is rare in sheltered high altitude cliffs and moist feldmarks. fliytogeogrgghy: This species occurs in South Georgia, the Falklands, Tierra del Fuego, Patagonian Channels, Valdivian region and Juan Fernandez (400-795 m. on Mas a Tierra). The report from Tasmania in Rodway (1916) requires confirmation (see Map 8). Literature Records (FALKLANDS): Anonymous (KiI'hnemann 1937’ 1949); Skottsberg - Port Stanley (Fulford 1966, Skottsberg 1913), Port Harriet (Fulford 1966); Skottsberg 8 Halle - Port Harriet (Stephani 1911). 134 Additional Literature Records: SOUTH GEORGIA: Anonymous - (Stephani 1909); Skottsberg - Royal Bay (Steere 1961, Stephani 1905a). TIERRA DEL FUEGO: Anonymous - Kiihnemann 1937, 1949, Stephani 1909); Chelminski - B. Orange (Fulford 1966 as L. cuspidata), Cunningham - Pto. Churruca (Fulford 1966); Dusen - R. Azopardo, Pto. Angosto (Stephani 1901a); I. Desolacion (Fulford 1966), R. Azopardo (Fulford 1966 as L. cuspidata); Gasperi - B. Angelito (Fulford 1966 as L. chordulifera 8 L. cuspidata, Gola 1923); B. Ainsworth (Fulford 1966, Gola 1923), M. Sarmiento (Fulford 1966 as L. chordulifera 8 L. cuspi- gaLa_, Gola 1923), I. Laberinto (Fulford 1966 as L. chordulifera 8 L. cuspidata, Gola 1923 as L. chordulifera 8 L. hastata), B. Parry (Gola 1923 as L. chordulifera 8 L. hastata);H'assel - Lapataia, Est. Los Cerros, L. Roca, L. M (Fulford 1966), L. Fagnano, L. Roca, Est. La Rubi, Est. Carmen (Fulford 1966 as L. cgpidata); Racovitza - I. Clarence (actually I. Capitan Arecena) (Fulford 1966, Stephani 1901b); Santesson - Pto. Yartou, Ushuaia (Arnell 1955 as L. hastata), 5. A1- vear, Pto. Yartou (Arnell 1955, Taylor 1960), Est. Yendegaia (Arnell 1955 as L. cugpidata); Skottsberg - Ushuaia, I. Navarino, Harberton, B. Tekenika, Pto. Cook (Stephani 1905a), Pto. Cook (Fulford 1966 as as L. chordulifera 8 L. cuspidata); Skottsberg 8 Halle - R. Azopardo (Fulford 1966, Stephani 1911 as L. chordulifera 8 L. 3.9.5.3232), Cta. Gomez, 8. Harris, R. AZOpardo (Stephani 1911); giegazzini - M. Richardson (Massalongo 1885, 1927) I. de los Estados (Fulford 1966). PATAGONIA: Anonymou - (KO‘hnemann 1937 as L. 93311353). Astrolabe - Pto. del Hambre (Fulford 1966); Claude-Joseph-Lanalhue (Fulford 1966); W - Pto. Grappler (Fulford 1966); m - Punta Arenas .u . n' " .- R" . . u- v c .1 0 .- . ‘l ‘ u ‘ ... . s . on e . I o . . a. . '0 I . .‘ '- U 1., a . '- '1 .‘ I '0 i ' 135 (Fulford1966), Arch. de los Chonos (Fulford 1966, G. L. 8 N. 1847, Taylor & Hooker 1847b as J. chordulifera, Taylor 1960); Dusen - Pto. Bueno, S. Molyneux, R. Aisen, I. Guaitecas, La Ensenada (Stephani 1900), Punta Arenas (Stephani 1901a), Pto. Bueno, Cta. Hale (Fulford 1966). R. Aisen, L. Llanquihue (Fulford 1966), Corral (Fulford 1966 as L. cuspidata); Fulford - Fuerte Bulness, between Punta Arenas 8 Hotel Rubens, Pta. Leon (Fulford 1966), between Punta Arenas 8 Hotel Ruben (Fulford 1966 as L. cuspidata; Hahn - Valdivia (Fulford 1966); H'assel - Fuerte Bulnes, R. Blanco (Fulford 1966), Pta. Leon (Fulford 1966 as L. cuspidata); Heggg - Valdivia (Herzog 1923 as L. m- 1i_fe__r;i1_8 L. m); Lechler - Rada York (Fulford 1966); M - Strait of Magellan (Fulford 1966); Popelear Je _T_e__r_l_g_g - Strait of Magellan (Fulford 1966); Santesson - Puerto Montt (Arnell 1955, Taylor 1960), Llanquihue, Alerce (Taylor 1960); Savatier - Pto. Eden (Bescherelle 8 Massalongo 1889); Schwabe - Chalhuin - R. Colun (Her- zog 1960 as L. cuspidata), Pto. Puyuhuapi (Herzog 1939, 1954 as L. fernandeziensis), C. Tesoro, Termas de Puyehue (Herzog 1954), Puca- trihue, Huitrapulli-Aleucapi, Chaihuin - R. Colun, L. Pellaifa (Her- 2091960); Skottsberg - Patagonia (Stephani 1922 as L. cuspidata 8 L. gijSJ, Taylor 1960 as L. cuspidata 8 L. effusa); Skottsberg 8 M13; R. de las Minas, Pto. Cutter, 8. Chacabuco, I. Guaitecas, Pto. Quellon, Pampa del Chalia (Stephani 1911); 23111135- I. Victoria (MOller 1955); Spegazzini Poyo-huapi (Massa- longo 1906, 1927); M - Punta Arenas (Fulford 1966 as L. Lusa- Qita). Corral (Fulford 1966); Thomasson - between L. Villarrica 8 V. Lanin (Thomasson 1963); Warnstorf - Strait of Magellan (Fulford u- u on. -., l I . e o- n v I 0 1| "I... A . '0 . .‘. . u . .- _ I I ‘ u 39... ‘ . 8‘ ... ‘ e .' 'Io. 0v ‘ I. I D u '\ \.. ‘- . i._ , I I I Is. \ ‘ n I“ ' \. n n '- I. i ‘1 \ . ‘0 D I .‘ c a ’0 } u c". ‘ I ’I r T I ,Q o ’l u I 136 1966). JUAN FERNANDEZ. Skottsberg (Stephani 1922 as L. fernandezi- g_n_s_i_§_). MAS A TIERRA: _KJflchJ - Cordon Chifladores (Arnell 1957); Skottsberg (Fulford 1966), Q. Damajuana - 400-500 m., above Plazoleta del Yunque (Arnell 1957), between Piedra Agujeriada 8 Laura-650 m., Pangal-580 m., near El Yunque-795 m., Centinela-BSO m., C. Piramide- 600 m., SalsipuedeS-66O m. (Herzog 1942 as L. fernandeziensis). TAS- MANIA: Anonymou - Mt. Wellington, Mt. Fielt, West Coast (Rodway 1916). ' FALKLAND SPECIMENS SEEN: EAST FALKLANDS: 1843 Hooker 3.11. as Jungermannia laevifolia var. incrassata (FH). PORT WILLIAM REGION, sunmit ridge of Mt. Low, c. 245 m. (3230 8 3238); north side of Gypsy Cove, sea level (3242, 3243, 3252 8 3258); Engineer Point headland, The Narrows, c. 15 m. Imshaug 40618; ridge on Engineer Point peninsula, 18-25 m. (2817). STANLEY REGION, Port Stanley, 8 April 1902 Skotts- nggfl. as Legidozia saddlensis (S-PA); near Sapper Hill, 29 October 1907 Skottsber as L. M (S-PA, UPS); Sapper Hill. 135 m. (2392, 2321.1g911, 2399, 2407-c. per., 2410. 2413-c. sporo., 2414 a 2419); Tumbledown Mt., 155-230m. (2375 A. 2380, 238482391);‘Goa’t‘Ridge, c. 180m. (3118, 3205, 320883216); summit ridge of north peak of Two Sisters, 245-290 111. (2703 A, 2727, 2732 J, 2735 8 2736); sunmit of Mt. Kent, 455 m. (fl). MT. USBORNE REGION, gap between Mt. Usborne 281able Rock, c. 440 m. (M _B_); summit of Mt. Usborne 1, c. 700 m. (_2__4_9_5_). NEST FALKLANDS: PORT HOWARD, Freezer Rocks, east Slope of Mt. Maria, 320 m. (M 8 1131). FOX BAY REGION, summit of East Head, 180m. (1111 8 31g); valley east of Sulivan House, c. 75 m. (_3§_3_l_). MT. ADAM, east side of sumnit ridge, 670-700 111. (3211). HILL COVE REGION, summit of East French Peak, 305 m. (2211); SUM“: 01’ W851: 137 French Peak, 290 m. (2962 8 2963); summit of Mt. Fegen, 335-360 m. (3065, 3073 8 3076); ridge of north slope of Mt. Fegen, c. 275 m. (1151 a 3062). NMHTIONAL SPECIMENS SEEN: SOUTH GEORGIA: Royal Bay, Moltke Harbour, 29 April 1902 Skottsberg _.p_. (S-PA); Cumberland Bay, May 1902 Skottsberg s.n. (S-PA). CHILE. MAGALLANES: Cabo de Hornos, Hooker i-fl» (NY). ARGENTINA. TIERRA DEL FUEGO: I. de los Estados, B. Blossom, March 1882 Spegazzini 4_8_ (NY); I. de los Estados, Pen- guin Rookery, February 1882 Spegazzini 13 (NY). (?) CHILE. MAGAL- LANES - (?) ARGENTINA. TIERRA DEL FUEGO: L. Fagnano, 10 March 1908 Halle_l7_5 as L. pallida (UPS). MAGALLANES: R. Azopardo, 2 March 1908 Halle 8 Skottsberg 175 as L. pallida (S-PA, UPS), March 1896 Dusen 8_3_a_as L. setiformis (S-PA), March 1896 Dusen §_._1_1_. as L. sad- dlensis (S-PA); _S_. Almirantazgo, 2 May 1908 Skottsberg 8 Halle l7 ——_ as L. ngLi_d_a (UPS); 8. Keta, 24 February 1929 Roivainen 2028 as L. W (S-PA); I. Riesco, R. Grande, 16 April 1908 fla_l_l__e_8 M. 933L151 as L. M (UPS); S. Skyring, Pta. Eulogio, 22 April 1908 fla__l_l_e8 Skottsberg 175 as L. pallida (UPS); B. Halt, Cunningham go as L. pallida (G). Lepidozia fuegiensis Steph. ngLdozia fuegiensis Steph. K. Svenska VetenskAkad. Handl. 46 (9); 53. f. 24, f-g. 1911. Original material: Chile, Prov. Magai- lanes, Pto. Cutter, Cta. Gomez, R. Fontaine and R. Azopardo, fla_1__1_e_8 Skottsberg (G) - Fulford (1966). 138 Lepidozia minuta Steph. Sp. Hep. 3: 603. 1909 non Lepidozia minuta Col. Trans. Proc. N. Z. 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