Cryptogamie, Bryologie, 2012, 33 (3): 199-255 © 2012 Adac. Tous droits réservés Checklist of the bryophytes of Madagascar Lovanomenjanahary MARLINE a,b,c**, Roger Lala ANDRIAMIARISOA d, Jacques BARDAT e, Min CHUAH-PETIOT f, Terry A.J. HEDDERSON b, Catherine REEB e, Dominique STRASBERG a, Nicholas WILDING b, Claudine AH-PENG a,b aUniversité de La Réunion, UMR PVBMT, 15 avenue René Cassin, BP 7151, 97 715 Saint Denis Cedex 9, France bUniversity of Cape Town, Botany Department, Private Bag X3, 7701 Rondebosch, South Africa cUniversité d’Antananarivo, Faculté des Sciences, Département de Biologie et Ecologie Végétales, BP 566, Antananarivo, Madagascar dMissouri Botanical Garden Madagascar, PO Box 3391 Antananarivo 101 Madagascar eMuséum National d’Histoire Naturelle, UMR CNRS-7205, Département de Systématique et Evolution, 57 rue Cuvier, 75005 Paris France fUniversity of Sain Malaysia, School of Biological Sciences, 11800 USM Pulau Penang, Malaysia (Received 26 August 2011, accepted 11 January 2012) Abstract – This updated checklist of the bryophytes of Madagascar was compiled from the literature, herbarium and recent collections. A total of 1144 species and infraspecific taxa are recorded for Madagascar composed of 751 mosses, 390 liverworts and 3 hornworts. 28.67% of the species reported for the Island are endemics. Twenty-four taxa are newly recorded for Madagascar and two of them, Fissidens cyathaeicola Brugg.Nann. and Taxithelium nepalense (Schwägr.) Broth. are reported for the first time for the Sub-Saharan Africa region. Hornworts / liverworts / mosses / Madagascar INTRODUCTION Madagascar is located in the Indian Ocean, between 11˚57’ and 25˚39’ latitude, 400 km from the eastern coast of Africa. It is the world’s fourth largest island, with a total area of 590 000 km2, the distance between Cap d’Ambre * Author for correspondence and reprints: marlinelova@gmail.com 200 L. Marline et al. (North) and Cap Sainte Marie (South) is nearly 1570 km, and the distance from West to East is around 560 km (Ingram & Dawson, 2005). The island consists of a central plateau of volcanic origin which culminates to the north in peaks over 2800 meters. To the East, the plateau falls directly and abruptly onto a narrow coastal plain, while in the West it descends more slowly to a broader plain, along the Mozambique Channel. The island is considered a biodiversity hot spot: 85% of the flora (Gautier & Goodman, 2003) and 90% of the fauna are endemics (Goodman, 2008). The tropical forests of Madagascar are among the highest priority areas in the world for biodiversity conservation (Myers et al., 2000). Eighty percent of Madagascar’s original forests have disappeared in the 1500-2000 years since the arrival of the first humans (Harper et al., 2007). At present, only 12% of the surface of the island is covered by natural forest. These remant forests are highly endangered by threat of deforestation. Cultivation involving slash and burn agriculture (locally called tavy) is the main cause of deforestation. Tavy consists of clearing and burning native forest. The burnt area is used for various plantations until the soils are impoverished, and then abandoned (Ingram & Dawson, 2005). The increasing population and the absence of resources and techniques to improve agricultural productivity and pasture lands threaten to destroy what remains of this unique biology (Sussman et al., 1994). The classification by Du Puy and Morat (1996) divides the primary vegetation of Madagascar into two formations: the evergreen and the deciduous formations. The evergreen formations (East and Center) are: Coastal forest (East); Evergreen humid forest, low elevation (0-800 m); Evergreen humid forest, mid elevation (800-1800 m); Evergreen humid forest, lower montagne (1800-2000 m); Mountain, Schrubland (Philipia) (> 1800 m); Evergreen, sclerophyllous (Uapaca) woodland (800-2000 m) and Marshland (0-800 m) The deciduous formations (West and South) are: Coastal forest (West); Deciduous, seasonally dry, western forest (0-800 m); Deciduous, dry, southern forest and schrubland (0-300 m). The distribution of different vegetation formations in Madagascar is shown in Fig. 1. The study on the flora of Madagascar has, for a long time, focused on vascular plants, with few studies being undertaken on bryophytes. Indeed, it was only during the 19th century that collectors first began to collect bryophytes in the Island (Dorr, 1997). There are more than 40 botanists that collected bryophytes on the island during the 19th century. The majority of which were French botanists and explorers. Collections through this period, by Cardot, Tixier, Cramers, Chénagon, and Perrier de la Bathie are mainly located at Paris herbarium (PC). Biographical data of bryophyte collectors are reported in Dorr (1997). The oldest collections are located at PC. From 1960, date of the independence of Madagascar, duplicates of the collections were hosted at the National Herbarium of Madagascar (TAN), of which more than a thousand samples still need to be identified. Nowadays, collections of Madagascan bryophytes are hosted in different herbaria around the world. This article aims to provide a comprehensive list of bryophytes from Madagascar based on literature records and additional collections, as well as provide information on the type specimens for endemic species. We hope that this preliminary work will be helpful for other bryologists and will stimulate and facilitate future work on the bryoflora of Madagascar. Checklist of the bryophytes of Madagascar 201 Fig. 1. Vegetation map of Madagascar (Du Puy & Morat, 1996) (reproduction with the permission of IRD). METHODS This checklist was compiled from the literature, data from various herbaria, and recent collections by the authors. This work was greatly facilitated by the ground work of O’Shea (2006) and Wigginton (2009) for their respective checklists of mosses, liverworts and hornworts for sub-Saharan Africa. 202 L. Marline et al. Presentation of the list – Synopsis of families and genera Families are ranked alphabetically within each order and sub order. The numbers in brackets against each genus indicate the number of taxa within the genus. Synopses of genera and family are based on the work of CrandallStotler et al. (2009) for liverworts, Renzaglia et al. (2009) for hornworts and Goffinet et al. (2009) for mosses. – Format of the list The taxa are arranged alphabetically within the three divisions of bryophytes: Marchantiophyta (liverworts), Anthocerotophyta (hornworts) and Bryophyta (mosses) and within each family. Scientific names appear in bold italic type followed by the authors who discribed the taxa. Endemic taxa are followed by “End”. New records from Madagascar are preceded by *. Herbarium abbreviations follow Thiers (2010). Uncompletely identified species that have been reported by O’Shea (2006) and Wigginton (2009) are taken into account in this checklist. The references following each taxon are the record of the presence of the taxon in Madagascar, it refers mainly to the work of O’Shea (2006) and Wigginton (2009). For new records from herbarium specimens, the references following the taxa are: locality, habitat, altitude, geographic coordinates, collection date, collector(s), collector’s number, and herbarium. For new records that were recently collected during the BRYOLAT expedition in November 2009 and other fieldtrips, as well as for type specimen information, references following the species names are: locality, microhabitat, geographic coordinates, date, collector, collection number, and determinator. When “unknown” is specified after type, this means that no information was given by the specimens, except “Madagascar” as main locality. “N.S.” is used when the holotype or herbarium are “not specified” and “#” is used when the information is considered doubtful. RESULTS Synopsis of the genera and families of Liverworts, Hornworts and Mosses DIVISION MARCHANTIOPHYTA (LIVERWORTS) CLASS MARCHANTIOPSIDA Subclass Marchantiidae ORDER MARCHANTIALES Family Aytoniaceae Asterella (2) Plagiochasma (1) Family Cyathodiaceae Cyathodium (1) Family Dumortieraceae Dumortiera (1) Family Marchantiaceae Marchantia (2) Family Ricciaceae Riccia (5) Family Targioniaceae Targionia (2) Checklist of the bryophytes of Madagascar 203 CLASS JUNGERMANNIOPSIDA Subclass Pelliidae ORDER FOSSOMBRONIALES Family Fossombroniaceae Fossombronia (1) ORDER PALLAVICINIALES Family Pallaviciniaceae Pallavicinia (1) Symphyogyna (2) Symphyogynopsis (1) Subclass Metzgeriidae ORDER PLEUROZIALES Family Pleuroziaceae Pleurozia (1) ORDER METZGERIALES Family Aneuraceae Aneura (1) Riccardia (9) Family Metzgeriaceae Metzgeria (6) Subclass Jungermanniidae ORDER PORELLALES Family Frullaniaceae Frullania (38) Family Lejeuneaceae Acanthocoleus (2) Acrolejeunea (3) Archilejeunea (4) Brachiolejeunea (1) Bryopteris (1) Caudalejeunea (4) Ceratolejeunea (6) Cheilolejeunea (14) Cololejeunea (54) Colura (9) Diplasiolejeunea (26) Drepanolejeunea (9) Frullanoides (1) Haplolejeunea (1) Lejeunea (22) Lepidolejeunea (1) Leptolejeunea (2) Lopholejeunea (12) Marchesinia (2) Mastigolejeunea (2) Metalejeunea (1) Microlejeunea (5) Odontolejeunea (1) Otolejeunea (3) Prionolejeunea (1) Ptychanthus (1) Pycnolejeunea (1) Schiffneriolejeunea (6) Symbiezidium (2) Taxilejeunea (3) Thysananthus (1) Xylolejeunea (1) Family Porellaceae Porella (5) Family Radulaceae Radula (11) ORDER JUNGERMANNIALES Family Adelanthaceae Adelanthus (2) Family Arnelliaceae Gongylanthus (1) Family Balantiopsidaceae Isotachis (1) Family Calypogeiaceae Calypogeia (7) Mnioloma (1) Family Cephaloziaceae Cephalozia (1) Iwatsukia (1) Family Cephaloziellaceae Amphicephalozia (1) Cephaloziella (2) Cylindrocolea (2) Kymatocalyx (3) Family Herbertaceae Herbertus (2) Familly Jamesoniellaceae Cryptochila (1) Cuspidatula (1) Denotarisia (1) Syzygiella (3) Family Jungermanniaceae Gottschelia (1) Notoscyphus (1) Solenostoma (4) Family Lepidoziaceae Amazoopsis (1) Bazzania (11) Kurzia (2) Lepidozia (2) Telaranea (3) Family Lophocoleaceae Chiloscyphus (8) Clasmatocolea (1) Conoscyphus (1) Heteroscyphus (2) Leptoscyphus (2) 204 L. 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Family Mastigophoraceae Mastigophora (1) Family Plagiochilaceae Plagiochila (21) Family Scapaniaceae Anastrophyllum (1) Andrewsianthus (1) Plicanthus (2) Family Schistochilaceae Gottschea (1) Schistochila (2) DIVISION ANTHOCEROTOPHYTA (HORNWORTS) CLASS ANTHOCEROTOPSIDA ORDER ANTHOCEROTALES Family Anthocerotaceae Anthoceros (1) Dendroceros (1) ORDER NOTOTHYLADALES Family Notothyladaceae Phaeoceros (1) DIVISION BRYOPHYTA (MOSS) Superclass I CLASS SPHAGNOPSIDA ORDER SPHAGNALES Family Sphagnaceae Sphagnum (16) Superclass II CLASS ANDREAEOPSIDA ORDER ANDREAEALES Family Andreaeaceae Andreaea (2) Superclass III CLASS POLYTRICHOPSIDA ORDER POLYTRICHALES Family Polytrichaceae Atrichum (1) Pogonatum (4) Polytrichastrum (1) Polytrichum (5) CLASS BRYOPSIDA SUBCLASS FUNARIIDAE ORDER GIGASPERMALES Family Gigaspermaceae Gigaspermum (1) Oedipodiella (1) ORDER FUNARIALES Family Funariaceae Entosthodon (4) Funaria (5) Physcomitrium (2) SUBCLASS DICRANIDAE ORDER GRIMMIALES Family Grimmiaceae Grimmia (3) Bucklandiella (1) Family Ptychomitriaceae Ptychomitrium (2) ORDER ARCHIDIALES Family Archidiaceae Archidium (2) ORDER DICRANALES Family Bruchiaceae Trematodon (7) Family Calymperaceae Calymperes (10) Leucophanes (3) Mitthyridium (2) Octoblepharum (2) Syrrhopodon (23) Family Ditrichaceae Ceratodon (1) Distichium (1) Ditrichum (4) Garckea (1) Pleuridium (1) Family Dicranaceae Anisothecium (1) Aongstroemia (2) Dicranella (6) Dicranoloma (3) Dicranum (1) Holomitrium (11) Leptotrichella (2) Leucoloma (55) Microcampylopus (1) Pseudephemerum (1) Family Erpodiaceae Erpodium (2) Family Eustichiaceae Eustichia (1) Checklist of the bryophytes of Madagascar Family Fissidentaceae Fissidens (40) Family Leucobryaceae Atractylocarpus (1) Campylopus (23) Leucobryum (19) Ochrobryum (1) Family Rhachitheciaceae Rhachithecium (1) Family Rhabdoweisiaceae Oreoweisia (1) ORDER POTTIALES Family Pottiaceae Anoectangium (3) Barbula (6) Bryoerythrophyllum (1) Chionoloma (1) Didymodon (1) Gymnostomiella (1) Gymnostomum (1) Hymenostylium (1) Hyophila (7) Leptodontium (3) Splachnobryum (2) Streptopogon (2) Tortella (1) Tortula (4) Trichostomum (6) Weisiopsis (1) Weissia (1) Family Serpotortellaceae Serpotortella (2) SUBCLASS BRYIDAE Super order Bryanae ORDER SPLACHNALES Family Meesiaceae Leptobryum (1) Family Splachnaceae Brachymitrion (1) Tayloria (4) ORDER BRYALES Family Bryaceae Anomobryum (2) Brachymenium (12) Bryum (26) Mniobryum (1) Rhodobryum (3) Family Mniaceae Epipterygium (1) Mielichhoferia (2) Plagiomnium (1) Pohlia (4) 205 Pseudopohlia (1) Schizymenium (2) ORDER BARTRAMIALES Family Bartramiaceae Bartramia (2) Breutelia (8) Philonotis (7) ORDER ORTHOTRICHALES Family Orthotrichaceae Cardotiella (4) Groutiella (1) Macrocoma (1) Macromitrium (24) Schlotheimia (31) Ulota (1) Zygodon (4) ORDER HEDWIGIALES Family Hedwigiaceae Braunia (1) Bryowijkia (1) Hedwigia (1) Family Rhacocarpaceae Rhacocarpus (1) ORDER RHIZOGONIALES Family Rhizogoniaceae Pyrrhobryum (1) Seper order Hypnanae ORDER HYPNODENDRALES Family Racopilaceae Powellia (2) Racopilum (10) ORDER HOOKERIALES Family Daltoniaceae Adelothecium (1) Calyptrochaeta (1) Daltonia (5) Distichophyllum (2) Family Hypopterygiaceae Hypopterygium (1) Lopidium (1) Family Leucomiaceae Leucomium (1) Family Pilotrichaceae Callicostella (13) Cyclodictyon (3) Hookeriopsis (1) Lepidopilidium (5) Lepidopilum (3) ORDER HYPNALES Family Amblystegiaceae Amblystegium (2) Hygrohypnum (1) 206 L. 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Family Anomodontaceae Herpetineuron (1) Family Brachytheciaceae Aerolindigia (1) Brachythecium (2) Eurhynchium (2) Helicodontium (1) Palamocladium (1) Rhynchostegiella (1) Rhynchostegium (5) Schimperella (1) Squamidium (1) Family Catagoniaceae Catagonium (1) Family Cryphaeaceae Cryphaea (2) Schoenobryum (1) Family Entodontaceae Campylodontium (1) Entodon (5) Erythrodontium (1) Mesonodon (1) Family Fabroniaceae Fabronia (8) Rhizofabronia (1) Family Hypnaceae Ectropothecium (12) Gammiella (1) Glossadelphus (3) Hypnum (2) Mittenothamnium (7) Phyllodon (2) Rhacopilopsis (2) Taxiphyllum (1) Vesicularia (9) Family Leptodontaceae Leptodon (1) Family Leskeaceae Pseudoleskea (1) Family Leucodontaceae Leucodon (2) Pterogoniadelphus (1) Pterogonium (2) Family Meteoriaceae Aerobryopsis (1) Barbella (1) Floribundaria (4) Papillaria (5) Trachypodopsis (1) Trachypus (2) Family Neckeraceae Homaliodendron (2) Neckera (2) Neckeropsis (4) Orthostichella (5) Pilotrichella (4) Pinnatella (1) Porotrichum (5) Thamnobryum (1) Family Phyllogoniaceae Phyllogonium (1) Family Prionodontaceae Prionodon (1) Family Pterigynandraceae Trachyphyllum (2) Family Pterobryaceae Calyptothecium (2) Hildebrandtiella (3) Jaegerina (2) Orthostichidium (1) Orthostichopsis (5) Pterobryopsis (1) Renauldia (3) Family Pylaisiadelphaceae Isopterygium (14) Taxithelium (7) Wijkia (4) Family Regmatodontaceae Regmatodon (1) Family Rigodiaceae Rigodium (1) Family Rutenbergiaceae Rutenbergia (5) Family Sematophyllaceae Acanthorrhynchium (1) Acroporium (5) Clastobryophilum (1) Macrohymenium (4) Meiothecium (3) Radulina (1) Rhaphidorrhynchium (3) Sematophyllum (11) Trichosteleum (10) Warburgiella (1) Family Stereophyllaceae Entodontopsis (2) Stereophyllum (1) Family Symphyodontaceae Symphyodon (1) Family Thuidiaceae Pelekium (7) Rauiella (1) Thuidium (3) Checklist of the bryophytes of Madagascar 207 Liverworts reported from Madagascar Adelanthaceae Adelanthus decipiens (Hook.) Mitt. (Pócs & Geissler, 2002) Adelanthus lindenbergianus (Lehm.) Mitt. (Grolle, 1995) Aneuraceae *Aneura pseudopinguis (Herzog) Pócs – Ambavaniasy, réserve de Vohimana, bois mort oblique en décomposition, alt. 849 m, S 18˚55’43.6’’ E 48˚29’56.6’’, 15/04/2010, Reeb, Andriamananatena & Bidault CRAE 106, TAN, det. Reeb & Andriamananatena; Ambavaniasy, réserve de Vohimana, forêt ombrophile, tronc vertical d’arbre vivant, alt. 830 m, S 18˚55’09.33’’ E 48˚30’49.0’’, 16/04/2010, Reeb, Andriamananatena & Bidault CRAE120 TAN, det. Reeb & Andriamananatena; Anjangoveratra, forêt du Makirovana, forêt ombrophile dense orientale, au dessus d’arbre mort horizontal, alt. 432 m, S 14˚10’13.9’’ E 49˚57’12.9’’, 04/05/2010, Reeb CR220, TAN, det. Reeb. Riccardia amazonica (Spruce) Schiffn. ex Gradst. (Pócs & Geissler, 2002) Riccardia erosa (Steph.) E.W. Jones (Pócs & Geissler, 2002) Riccardia longispica (Steph.) Pearson (Grolle, 1995; Pócs & Geissler, 2002) Riccardia multifida (L.) Gray (Grolle, 1995) Riccardia nudiflora (Steph.) Grolle (Grolle, 1995) Riccardia saccatiflora (Steph.) S.W. Arnell (Grolle, 1995) Riccardia sp. (Aneura caespitans Steph.) (Grolle, 1995) Riccardia sp. (Aneura corbieri Steph.) (Grolle, 1995) Riccardia sp. (Aneura dismieri Steph.) (Grolle, 1995) Arnelliaceae Gongylanthus ericetorum (Raddi) Nees (Grolle, 1995) Aytoniaceae Asterella coronata (Steph.) H.A. Mill. (Grolle, 1995) Asterella wilmsii (Steph.) S.W. Arnell (Grolle, 1995) Plagiochasma microcephalum (Steph.) Steph. (Grolle, 1995) Balantiopsidaceae Isotachis aubertii (Schwaegr.) Mitt. (Grolle, 1995; Pócs & Geissler, 2002) Calypogeiaceae Calypogeia annabonensis Steph. (Pócs & Geissler, 2002) Calypogeia arguta Nees & Mont. (Grolle, 1995) Calypogeia bidentula (F. Weber) Nees (Grolle, 1995) *Calypogeia fissa (L.) Raddi – Ankaratra, sur sol, alt. 2000 m, 27/03/1972, Cremers 1947, TAN. Calypogeia longifolia Steph. (Grolle, 1995; Pócs, 2005) Calypogeia mascarenensis Bischl. (Grolle, 1995) Calypogeia microstipula (Steph.) Steph. (Grolle, 1995) Mnioloma fuscum (Lehm.) R.M. Schust. (Grolle, 1995; Pócs & Geissler, 2002) 208 L. Marline et al. Cephaloziaceae Cephalozia connivens (Dicks.) Lindb. subsp. fissa Vá≈a (Grolle, 1995; Pócs, 1995; Pócs & Geissler, 2002) Iwatsukia jishibae (Steph.) N. Kitag. (Pócs, 1995; Pócs & Geissler, 2002) Cephaloziellaceae Amphicephalozia geisslerae Pócs & Vá≈a (Pócs & Vá≈a, 2001; Pócs & Geissler, 2002) End, Type – Antsiranana, Manongarivo Special Reserve, NW Madagascar, alt. 1300-1350 m, 10/03/1999, Geissler 19708/3, holotype G, paratype EGR, G, PRC. Cephaloziella kiaeri (Austin) S.W. Arnell (Grolle, 1995; Pócs & Geissler, 2002) Cephaloziella sp. (Grolle, 1995) Cylindrocolea atroviridis (Sim) Vá≈a (Grolle, 1995) Cylindrocolea madagascariensis (Steph.) R.M. Schust. (Grolle, 1995) End, Type – Imerina, Hildebrandt, herb (N.S.). Kymatocalyx dominicensis (Spruce) Vá≈a (Gradstein & Vá≈a, 1999) Kymatocalyx madagascariensis (Steph.) Gradst. & Vá≈a (Grolle, 1995; Pócs, 1995; Gradstein & Vá≈a, 1999; Pócs & Geissler, 2002) Kymatocalyx rhizomatica (Herzog) Gradst. & Vá≈a (Pócs & Geissler, 2002) Cyathodiaceae Cyathodium africanum Mitt. (Pócs & Geissler, 2002) Dumortieraceae *Dumortiera hirsuta (Sw.) Nees – Anjozorobe, forêt ombrophile d’altitude, sur des terres humides, 02/1959, Bosser 12648, TAN; Marojejy National Park, Talus terreux ombragé, alt. 690 m, S 14˚26’07.8” E 49˚45’49.9”, 27/11/2009, Ah-Peng 349, det. Ah-Peng; Ambavaniasy, réserve de Vohimana, forêt ombrophile d’altitude, au pied d’une cascade, alt. 849 m, S 18˚55’43.6’’ E 48˚29’ 56.6’’15/04/2010, Reeb, Andriamananatena & Bidault CRAE93, TAN, det. Reeb; Anjangoveratra, forêt du Makirovana, sur falaise suintante, alt. 508 m, S 14˚10’17.4’’ E 49˚57’07.5’’, 07/05/2010, Reeb CR302, TAN, det. Reeb. Fossombroniaceae Fossombronia sp. (Pócs & Geissler, 2002) Frullaniaceae Frullania anderssonii Aongstr. (Grolle, 1995) Frullania angulata Mitt. var. angulata (Grolle, 1995; Pócs & Geissler, 2002) Frullania angulata var. laciniata Demaret et Vanden Berghen (Grolle, 1995) Frullania apicalis Mitt. (Grolle, 1995; Pócs & Geissler, 2002) Frullania apiculata (Reinw. et al.) Nees (Grolle, 1995; Pócs, 1995; Pócs & Geissler, 2002) Frullania arecae (Spreng.) Gottsche (Grolle, 1995) Frullania borbonica Lindenb. (Grolle, 1995) Frullania bullata Steph. (Grolle, 1995) Frullania caffraria Steph. (Grolle, 1995) Frullania capensis Gottsche (Grolle, 1995) Frullania depressa Mitt. (Grolle, 1995) Frullania diptera (Lehm.) Drège (Grolle, 1995) Checklist of the bryophytes of Madagascar 209 *Frullania ecklonii (Spreng.) Gottsche et al. – Ambatovory, Tananarive, vestige de forêt, sur un tronc d’arbre, 04/1957, Bosser 11021, TAN. Frullania ericoides (Nees) Mont. (Grolle, 1995) Frullania gabonensis Vanden Berghen (Grolle, 1995; Pócs, 1995) Frullania grossiclava Steph. (Grolle, 1995; Pócs, 1995; Grolle, 2001; Pócs & Geissler, 2002) Frullania humbertii Vanden Berghen (Grolle, 1995) Frullania imerinensis Steph. (Grolle, 1995; Pócs & Geissler, 2002) Frullania lindenbergii Lehm. (Grolle, 1995; Pócs, 1995; Grolle, 2001; Pócs & Geissler, 2002) Frullania longistipula Steph. (Grolle, 1995) End, Type – Imerina, Ankadivavala, Camboué 162, holotype G. Frullania loricata Pearson (Grolle, 1995; Pócs & Geissler, 2002) Frullania loricata var. laxa Pearson (Grolle, 1995) End, Type – unknown. Frullania nodulosa (Reinw. et al.) Gottsche et al. var. nodulosa (Grolle, 1995; Pócs, 1995) Frullania obscurifolia Mitt. (Grolle, 1995; Pócs, 1995) Frullania purpurea Steph. (Grolle, 1995; Pócs, 1995; Pócs & Geissler, 2002) Frullania repandistipula Sande Lac. (Grolle, 1995) Frullania rigida Steph. (Grolle, 1995) End, Type – unknown. Frullania schimperi var. laciniata Vanden Berghen (Grolle, 1995) End, Type – Onraedt 74M8078c, holotype BR. Frullania schimperi Nees var. schimperi (Grolle, 1995; Pócs & Geissler, 2002) Frullania serrata var. pertenuis (Nees) Gottsche et al. (Grolle, 1995; Pócs, 1995) Frullania serrata Gottsche var. serrata (Grolle, 1995) Frullania socotrana Mitt. (Grolle, 1995) Frullania spongiosa Steph. (Grolle, 1995; Pócs, 1995) Frullania trinervis (Lehm.) Drège (Grolle, 1995) Frullania usambarana Schiffn. ex Steph. (Grolle, 1995; Pócs, 1995) Frullania usambarana var. reducta Vanden Berghen (Grolle, 1995) Frullania vanden-berghenii Pócs (Grolle, 1995; Pócs, 1995) Frullania variegata Steph. (Grolle, 1995; Pócs, 1995; Pócs & Geissler, 2002) Herbertaceae Herbertus dicranus (Taylor ex Gottsche et al.) Trevis. (Grolle, 1995; Pócs, 1995; Grolle, 2001; Hodgetts, 2008) Herbertus juniperoideus (Sw.) Grolle (Grolle, 1995; Grolle, 2001; Pócs & Geissler, 2002; Hodgetts, 2008) Jamesoniellaceae Cryptochila grandiflora (Lindenb. et Gottsche) Grolle (Grolle, 1995) Cuspidatula contracta (Reinw. et al.) Steph. (Pócs, 1995) Denotarisia linguifolia (De Not.) Grolle (Pócs & Geissler, 2002) Syzygiella purpurascens (Steph) Feldberg et al. (Grolle, 1995; Pócs, 1995; Pócs & Geissler, 2002) Syzygiella concreta (Gottsche) Spruce (Grolle, 1995; Pócs & Geissler, 2002) Syzygiella geminifolia (Mitt.) Steph. (Grolle, 1995; Pócs, 1995; Pócs & Geissler, 2002) 210 L. Marline et al. Jungermanniaceae Gottschelia schizopleura (Spruce) Grolle (Grolle, 1995; Pócs, 1995; Pócs & Geissler, 2002) Notoscyphus lutescens (Lehm. et Lindenb.) Mitt. (Grolle, 1995; Pócs & Geissler, 2002) Solenostoma borgenii (Gottsche) Steph. (Grolle, 1995) Solenostoma onraedtii (Và≈a) Và≈a et al. (Grolle, 1995) Solenostoma renauldii (Và≈a) Và≈a et al. (Grolle, 1995) Solenostoma sphaerocarpum (Hook.) Steph. (Grolle, 1995) Lejeuneaceae Acanthocoleus aberrans (Lindenb. et Gottsche) Kruijt (Kruijt, 1988; Grolle, 1995) Acanthocoleus madagascariensis (Steph.) Kruijt (Grolle, 1995; Pócs, 1995; Pócs & Geissler, 2002) Acrolejeunea aulacophora (Mont.) Steph. (Grolle, 1995) Acrolejeunea emergens (Mitt.) Steph. var. emergens (Grolle, 1995; Pócs & Geissler, 2002) Acrolejeunea pycnoclada (Taylor) Schiffn. (Grolle, 1995) Archilejeunea alata Steph. (Grolle, 1995) Archilejeunea brevilobula Steph. (Grolle, 1995) End, Type – Oriental coast, Sakara, hototype G. Archilejeunea helenae Pócs et Gyarmati (Pócs & Sass-Gyarmati, 2006) End, Type – Mananara-Nord Biosphere Reserve, lowland rainforest on the East slopes of Mahavoho Hill, epiphyllous on relatively rigid leathery leaf, alt. 250-350 m, S 16˚27’ E 49˚46.9-47.5’, 14-16/08/1998, Pócs & Szabó 9878/ FM, holotype EGR. Archilejeunea nobilis (Gottsche) Steph. (Grolle, 1995) End, Type – unknown. Brachiolejeunea phyllorhiza (Nees) Kruijt et Gradst. (Grolle, 1995) Bryopteris gaudichaudii Gottsche (Grolle, 1995; Pócs, 1995) Caudalejeunea grolleana Gradst. (Grolle, 1995) End, Type – Nosy Be, Lokobe natural reserve, on bark in primary forest, alt. 100 m, 15/07/1971, Onraedt M5119, holotype BR, isotype JE. Caudalejeunea hanningtonii (Mitt.) Schiffn. (Grolle, 1995) Caudalejeunea lewallei Vanden Berghen (Grolle, 1995) Caudalejeunea recurvistipula (Gottsche) Schiffn (Grolle, 1995) Ceratolejeunea andringitrae Pócs (Pócs, 2011a) End, Type – Ihorombe Region in former Fianarantsoa Province, Andringitra National Park, Montane rainforest on the West side of Korokoro River, around Camp II, epiphyll, alt. 750-1000 m, S 22˚13’ E 47˚1-2’, 20-23/09/1994, Pócs & A. Szabó 9472/ AF, holotype EGR, isotypes MO, TAN. Ceratolejeunea belangeriana (Gottsche) Steph. (Bonner, 1953; Grolle, 1995; Pócs & Geissler, 2002; Pócs, 2011a) Ceratolejeunea diversicornua Steph. (Pócs, 2011a) Ceratolejeunea cornuta (Lindenb.) Schiffn. (Grolle, 1995; Pócs, 2011a) Ceratolejeunea papuliflora Steph. (Bonner, 1953; Grolle, 1995; Pócs, 2011a) Ceratolejeunea saroltae Pócs (Pócs, 2011a) End, Type – Toamasina Province, Maromizaha forest, Mossy montane rainforest with bamboo (Nastus sp.), undergrowth on the summit ridge of Mt. Maromizaha, south of the Andasibe National Park, on twigs, alt. 1080-1214 m, Sarolta Pócs 9890/BJ, 26/09/1998, holotype EGR, paratype EGR. Checklist of the bryophytes of Madagascar 211 Cheilolejeunea cordigera (Steph.) Grolle (Grolle, 1995; Pócs & Geissler, 2002) Cheilolejeunea cordistipula (Steph.) Grolle ex E.W. Jones (Grolle, 1995; Malombe, 2007) Cheilolejeunea decursiva (Sande Lac.) R.M. Schust. (Grolle, 1995; Pócs & Geissler, 2002; Malombe, 2007) Cheilolejeunea intertexta (Lindenb.) Steph. (Grolle, 1995; Pócs & Geissler, 2002; Malombe, 2007) Cheilolejeunea krakakammae (Lindenb.) R.M. Schust. (Grolle, 1995; Pócs & Geissler, 2002; Malombe 2007) Cheilolejeunea montagnei (Gottsche) R.M. Schust. (Grolle, 1995; Pócs & Geissler, 2002; Malombe, 2007) Cheilolejeunea pluriplicata (Pearson) R.M. Schust. (Grolle, 1995; Malombe, 2007) Cheilolejeunea pocsii E.W. Jones (Grolle, 1995; Malombe, 2007) Cheilolejeunea serpentina (Mitt.) Mizut. (Grolle, 1979; Grolle, 1995; Pócs, 1995; Pócs & Geissler, 2002; Malombe, 2007) Cheilolejeunea surrepens (Mitt.) E.W. Jones (Grolle, 1995; Pócs & Geissler, 2002) Cheilolejeunea trapezia (Nees) Kachroo et R.M. Schust. (Malombe, 2007) Cheilolejeunea trifaria (Reinw. et al.) Mizut. (Grolle, 1995; Pócs & Geissler, 2002) Cheilolejeunea usambarana (Steph.) Grolle (Malombe, 2007) Cheilolejeunea xanthocarpa (Lehm. & Lindenb.) Malombe (Pócs, 1995; Malombe, 2007) Cololejeunea adnata Tixier (Tixier, 1985; Grolle, 1995) End, Type – Tamatave, Foulpointe, forêt d’Analalava, alt. 100 m, 05/12/1975, Tixier 7637, holotype PC. Cololejeunea andapania Tixier (Tixier, 1985; Grolle, 1995) End, Type – DiégoSuarez, massif de l’Anjanaharibe, Ouest d’Andapa, Haute Andramota, bassin de la rivière Lokoho, forêt ombrophile sur gneiss et granite, alt. 700 m, 10/12/1950, Humbert, Capuron & Cour 1/E, holotype PC. Cololejeunea androphylla var. madecassa Tixier (Grolle, 1995) End, Type – Antananarive, Mandialaza, alt. 1200 m, Tixier 7670, holotype PC. Cololejeunea ankaiana Tixier (Tixier, 1985; Grolle, 1995) End, Type – Route de Tamatave, Moramanga - Anosibeanala, alt. 1000 m, 18/4/1978, Tixier s.n., holotype PC. Cololejeunea appressa (A. Evans) Benedix (Grolle, 1995; Pócs & Geissler, 2002) Cololejeunea auriculata (E.W. Jones) R.M. Schust. (Tixier, 1985; Grolle, 1995; Pócs & Geissler, 2002) Cololejeunea bidentula (Steph.) E.W. Jones (Grolle, 1995) Cololejeunea bolombensis (Steph.) Vanden Berghen (Tixier, 1985; Grolle, 1995; Pócs & Geissler, 2002) Cololejeunea borbonica Tixier (Grolle, 1995) Cololejeunea camusii Tixier (Tixier, 1985; Grolle, 1995) End, Type – Antananarivo, Ambatoloana, Andasibe, alt. 1200 m, 10/11/1976, Tixier 9593, holotype PC. Cololejeunea capuronii Tixier (Grolle, 1995) End, Type – Massif de l’Anjanaharibe, à l’Ouest d’Andapa, Andramonta (pente et sommet nord), alt. 500-700 m, Humbert, Capuron & Cours s.n., holotype PC. Cololejeunea cardiocarpa (Mont.) A. Evans (Tixier, 1985; Grolle, 1995) Cololejeunea cristata (Steph.) R.M. Schust. (Tixier, 1985; Grolle, 1995) 212 L. Marline et al. Cololejeunea cuneata (Lehm. et Lindenb.) Herzog (Grolle, 1995; Pócs, 1995) Cololejeunea decemplicata (Steph.) Tixier (Grolle, 1995) Cololejeunea deroinii Tixier (Tixier, 1993; Grolle, 1995) End, Type – DiégoSuarez (Antsiranana), Massif du Marojejy, rive gauche de la rivière Manantenina, alt. 350 m, Deroin & Badré 5, holotype PC. Cololejeunea diplasiolejeuneoides Tixier (Tixier, 1985; Grolle, 1995) End, Type – Tamatave, Périnet, forêt d’Antaniditra, alt. 950 m, 6/7/1977, Tixier 10637, holotype PC. Cololejeunea disciflora Tixier (Grolle, 1995) End, Type – Antananarivo, Carion, Angavobe, alt. 1600 m, Tixier 10834, holotype PC. Cololejeunea distalopapillata (E.W. Jones) R.M. Schust. (Tixier, 1985; Grolle, 1995; Pócs & Geissler, 2002) Cololejeunea dolichodonta Tixier (Tixier, 1985; Grolle, 1995) End, Type – Fianarantsoa, Ambatofinandrahana, alt. 1650 m, 21/3/1978, Tixier 11394, holotype PC. Cololejeunea duvignaudii E.W. Jones (Tixier, 1985; Grolle, 1995; Pócs & Geissler, 2002) Cololejeunea duvignaudii var. papillata Tixier (Grolle, 1995) Cololejeunea elegans Steph. (Tixier, 1985; Grolle, 1995; Pócs & Geissler, 2002) Cololejeunea falcata var. madecassa Tixier (Grolle, 1995) End, Type – Tamatave, Périnet, alt. 917 m, Tixier 10160, holotype PC. Cololejeunea floccosa (Lehm. et Lindenb.) Schiffn. (Grolle, 1995) Cololejeunea floccosa var. trivittata Tixier (Grolle, 1995) Cololejeunea furcilobulata (Berrie et E.W. Jones) R.M. Schust. (Grolle, 1995) Cololejeunea hildebrandii (Austin) Steph. (Tixier, 1985; Grolle, 1995) Cololejeunea leloutrei (E.W. Jones) R.M. Schust. (Grolle, 1995; Pócs & Geissler, 2002) Cololejeunea leloutrei var. microlobulata Tixier (Tixier, 1985; Grolle, 1995) End, Type – Tananarive, Ambatolampy, station de Manjakatompo, alt. 1700 m, 27/10/1976, Tixier 9385, holotype PC. Cololejeunea leloutrei var. ugulurica Pócs ex Tixier (Grolle, 1995) Cololejeunea lemuriana Tixier (Tixier, 1985; Grolle, 1995) Cololejeunea marginata (Lehm. et Lindenb.) Schiffn. (Grolle, 1995; Pócs & Geissler, 2002) Cololejeunea microscopica (Taylor) Schiffn. var. exigua (A. Evans) Bernecker et Pocs (Grolle, 1995; Pócs & Bernecker, 2009) Cololejeunea minutissima (Sm.) Schiffn. (Grolle, 1995) Cololejeunea moramangae Tixier (Grolle, 1995; Pócs & Bernecker, 2009) Cololejeunea mocambiquensis S.W.Arnell (Grolle, 1995) Cololejeunea obliqua (Nees et Mont.) Schiffn. (Grolle, 1995; Pócs & Geissler, 2002) Cololejeunea obtusifolia (E.W. Jones) Tixier (Grolle, 1995; Pócs & Geissler, 2002) Cololejeunea peraffinis var. serrulata Benedix (Pócs & Geissler, 2002) Cololejeunea plagiochiliana Tixier (Grolle, 1995) End, Type – Périnet, Benoist s.n., holotype PC. Cololejeunea platyneura (Spruce) A. Evans (Grolle, 1995; Pócs & Geissler, 2002) Cololejeunea praeruptorum Tixier (Tixier, 1985; Grolle, 1995) End, Type – Mandialaza, forêt de l’Angavo, 19/12/1975, Tixier 7670, holotype PC. Checklist of the bryophytes of Madagascar 213 Cololejeunea pteroporum Tixier (Tixier, 1985; Grolle, 1995) End, Type – DiégoSuarez, station de roussettes, alt. 900 m, 06/11/1975, Tixier 7463, holotype PC. Cololejeunea raduliloba Steph. (Tixier, 1985; Grolle, 1995) Cololejeunea saltuum Tixier (Tixier, 1985; Grolle, 1995) End, Type – Fianarantsoa, Ranomafana, forêt du pk 24, alt. 1200 m, 30/12/1976, Tixier 10038, holotype PC. Cololejeunea sambiroana Tixier (Tixier, 1985; Grolle, 1995) End, Type – Sambirano, forêt d’Anjozobe, alt. 300m, 22/08/1977, Tixier 10744, holotype PC. Cololejeunea selangorensis Tixier (Tixier, 1985; Grolle, 1995) Cololejeunea subinflata Tixier (Tixier, 1985; Grolle, 1995) End, Type – Antananarivo, Manjakandriana, lac Mantasoa, 19/04/1979, Tixier 8158, holotype PC. Cololejeunea succinea Tixier (Tixier, 1985; Tixier, 1979a) End, Type – DiégoSuarez, Montagne d’Ambre, station des Roussettes, forêt de montagne, 03/11/1975, Tixier 7395, holotype PC. Cololejeunea tamatavensis Tixier (Tixier, 1985; Grolle, 1995) End, Type – Foulpointe, forêt d’Analalava, alt. 100 m, 05/12/1975, Tixier 7230, holotype PC. Cololejeunea tanzaniae Pócs (Grolle, 1995; Pócs & Geissler, 2002) Cololejeunea vidaliana Tixier (Grolle, 1995) Cololejeunea zenkeri (Steph.) E.W. Jones (Pócs & Geissler, 2002) Colura benoistii Jovet-Ast (Grolle, 1995) Colura bicornis Jovet-Ast (Grolle, 1995; Pócs, 1995) End, Type – Route de Moramanga à Anosibe, pk 29, forêt marécageuse, sur feuille de palmier, 15/09/1953, Millot & Bosser 6319, herb (N.S.) Colura digitalis (Mitt.) Steph. (Pócs, 1995; Pócs & Geissler, 2002) Colura heimii Jovet-Ast (Grolle, 1995) Colura humbertii Jovet-Ast (Grolle, 1995) Colura inornata Jovet-Ast (Grolle, 1995; Pócs, 1995) End, Type – Route de Moramanga à Anosibe, pk 29, forêt marécageuse, sur feuille de palmier, 15/09/1953, Millot & Bosser 6323, herb (N.S.) Colura obesa Jovet-Ast (Grolle, 1995) Colura obvoluta Jovet-Ast (Jovet-Ast, 1983; Grolle, 1995) Colura tenuicornis (A. Evans) Steph. (Grolle, 1995) Diplasiolejeunea andringitrae Pócs et Schafer-Verwimp (Pócs & SchaeferVerwimp, 2006) End, Type – Monts d’Andringitra, Nature Reserve, montane rainforest, West side of Korokoro River, around Camp II, alt. 750-1000 m, S 22˚13’ E 47˚1-2’, 20-23/9/1994, Pócs et al., 9472/AM, holotype EGR. Diplasiolejeunea aulae E.W. Jones (Grolle, 1995) Diplasiolejeunea auriculata Tixier (Grolle, 1995) End, Type – Fianarantsoa, forêt de Tsarafidy, Tixier 8908, holotype PC. Diplasiolejeunea cavifolia Steph. (Grolle, 1995; Pócs & Geissler, 2002) Diplasiolejeunea cobrensis Gottsche ex Steph. (Grolle, 1995; Pócs & Geissler, 2002) Diplasiolejeunea cobrensis Pócs subsp. antsirananae (Pócs, 2001; Pócs & Geissler, 2002) End, Type – Province d’Antsiranana, Réserve Spéciale de 214 L. Marline et al. Manongarivo, on the rain forest covered north ridge leading to the 1869 m high summit of Manongarivo massif, epiphylous, alt. 850-950 m, N 13˚59’ E 48˚26’, 3/1999, Geissler 19549/8, holotype G. Diplasiolejeunea comorensis Tixier (Grolle, 1995) Diplasiolejeunea cornuta Steph. (Grolle, 1995; Pócs, 1995) Diplasiolejeunea ensifera Tixier (Grolle, 1995) End, Type – Antananarivo, Andasibe, alt. 1400 m, Tixier 9626, holotype PC. Diplasiolejeunea hamata Tixier (Grolle, 1995) End, Type – Fianarantsoa, Vohiparara, Tixier 11373, holotype PC. Diplasiolejeunea insignis Tixier (Grolle, 1995) End, Type – Tananarive, Carion, Angavokely, alt. 1800 m, 27/01/1977, Tixier 10258, holotype PC. Diplasiolejeunea jonesii Tixier (Grolle, 1995) End, Type – Tamatave, Antaniditra, Vohiparara, alt. 1100 m, 30/12/1976, Tixier 10157, holotype PC. Diplasiolejeunea kraussiana (Lindenb.) Steph. (Grolle, 1995) Diplasiolejeunea lemuriana Tixier (Grolle, 1995) End, Type – Fianarantsoa, Ranomafana, alt. 1200 m, 30/12/1976, Tixier 10171, holotype PC. Diplasiolejeunea magnistipulata Tixier (Tixier, 1985; Grolle, 1995) Diplasiolejeunea ornata Pócs et Schäfer-Verwimp (Pócs & Schaefer-Verwimp, 2006) End, Type – Province deToamasina, Mananara - Nord Biosphere Reserve, Lowland rain forest on the NW slope of Behafotra Hill, alt. 250300 m, S 16˚27.2’ E 49˚47.7’, 16/08/1998, Pócs & Szabó 9877/CH, holotype EGR. Diplasiolejeunea palustrium Tixier (Grolle, 1995) End, Type – Tamatave, Périnet, Antaniditra, alt. 1000 m, 06/07/1977, Tixier 10636, holotype PC. Diplasiolejeunea phyllarthonii Tixier (Grolle, 1995) End, Type – Tamatave, Manohitaly (route du Chrome), alt. 1200 m, 03/03/1978, Tixier 11632, holotype PC. Diplasiolejeunea ramicola Tixier (Grolle, 1995) End, Type – Antananarivo, Ambatoloana, 1400 m, Tixier 9629, holotype PC. Diplasiolejeunea ranomafanae Pócs & Schäfer-Verwimp (Pócs & SchaeferVerwimp, 2006) End, Type – Ranomafana National Park, on the summit ridge of Mont Vatolampy near the telecom aerial, 2.3 km W of Ambatolahy village, alt. 1200 m, S 21˚15.10’ E 47˚24.44’, 30/07/2004, Pócs 04128/AK, holotype EGR. Diplasiolejeunea runssorensis Steph. (Grolle, 1995) Diplasiolejeunea subcornuta Tixier (Grolle, 1995) Diplasiolejeunea symoensii Vanden Berghen (Grolle, 1995) Diplasiolejeunea utriculata Steph. (Grolle, 1995) Diplasiolejeunea villaumei Steph. (Grolle, 1995) Diplasiolejeunea zakiae Tixier (Tixier, 1977; Pócs & Geissler, 2002) Drepanolejeunea cambouena Steph. (Grolle, 1995; Pócs, 2011b) Drepanolejeunea cultrella (Mitt.) Steph. (Pócs & Geissler, 2002) Drepanolejeunea geisslerae Pócs (Pócs, 2001; Pócs & Geissler, 2002) End, Type – Antsiranana, Réserve Spéciale de Manongarivo, NW Madagascar, abundant on palm stem, alt. 950-1050 m, N 13˚59’ E 48˚26’, 03/1999, Geissler 19559, holotype G. Drepanolejeunea madagascariensis (Steph.) Grolle (Grolle, 2001; Pócs & Geissler, 2002) Checklist of the bryophytes of Madagascar 215 Drepanolejeunea mascarena (S.W. Arnell) R.L. Zhu et Grolle (Zhu & Grolle, 2003) Drepanolejeunea pentadactyla (Mont.) Steph. (Mizutani, 1975; Pócs, 2011b) Drepanolejeunea physaefolia (Gottsche) Steph. (Grolle, 1995; Pócs, 1995; Pócs & Geissler, 2002) Drepanolejeunea pocsii Grolle (Pócs & Geissler, 2002) Drepanolejeunea trematodes (Nees) Bischl. (Grolle, 1995; Pócs & Geissler, 2002) Frullanoides tristis (Steph.) Van Slageren (Grolle, 1995) Haplolejeunea sticta Grolle (Grolle, 1995; Pócs & Geissler, 2002) Lejeunea alata Gottsche (Grolle, 1995; Pócs & Geissler, 2002) Lejeunea alata var. patriciae Pócs (Pócs, 2001; Pócs & Geissler, 2002) End, Type – Province d’Antsiranana, Réserve Spéciale de Manongarivo, on the rain forest covered north ridge leading to the 1869 m high summit of Manongarivo massif, epiphylous, N 13˚59’ E 48˚26’, 12/03/1999, Geissler 19856/1, holotype G. Lejeunea amaniensis E.W. Jones (Grolle, 1995) Lejeunea anisophylla Mont. (Grolle, 1985; Pócs & Geissler, 2002; Pócs, 2011b) Lejeunea capensis Gottsche (Grolle, 1995) Lejeunea confusa E.W. Jones (Pócs & Geissler, 2002) Lejeunea ecarinata (Steph.) Steph. (Grolle, 1995) Lejeunea eckloniana Lindenb. (Grolle, 1995; Pócs & Geissler, 2002) Lejeunea flava (Sw.) Nees (Grolle, 1995) Lejeunea flavovirens Aongstr. (Grolle, 1995; Pócs & Geissler, 2002) Lejeunea isophylla E.W. Jones (Grolle, 1995) Lejeunea julacea Steph. (Grolle, 1995; Pócs, 2011b) End, Type – Forsyth-Major B 473, holotype G. Lejeunea leucosis Besch. & Spruce (Grolle, 1995; Pócs, 2010a) Lejeunea masoalae Pócs (Pócs, 2010a) End, Type – West side of Masoala Peninsula, above Antongil Bay, montane rainforest on the summit ridge SE of Ambanizana village, alt. 660-720 m, S 15˚37’ E 50˚05’, 09/09/1994, Pócs 9448/AN, holotype EGR, isotype MO, TAN. Lejeunea obtusata Gottsche (Grolle, 1995) Lejeunea papilionacea Steph. (Grolle, 1995; Zhu & Grolle, 2001) Lejeunea phyllobola Nees et Mont. (Pócs, 1995) Lejeunea tabularis (Spreng.) Gottsche et al. (Grolle, 1995; Pócs & Geissler, 2002) Lejeunea tuberculosa Steph. (Grolle, 1995; Pócs & Geissler, 2002) Lejeunea villaumei (Steph.) Grolle (Grolle, 1995; Pócs & Geissler, 2002) Lejeunea vojtkoi Pócs (Pócs, 2010a) End, Type – 10 km North of Fianarantsoa, on decaying wood in montane forest, alt. 1230-1360 m, S 21˚15’ E 47˚14’, 22/09/1994, Vojtkó 9461/AY, holotype EGR. Lejeunea vulgariformis Gottsche (Grolle, 1995; Pócs, 2010a) Lepidolejeunea bidentula (Steph.) R.M. Schust. var. bidentula (Piippo, 1986; Grolle, 1995; Zhu & So, 2001) Leptolejeunea epiphylla (Mitt.) Steph. (Pócs & Geissler, 2002) *Leptolejeunea vitrea (Nees) Schiffn. – Anjozorobe, épiphylle, alt. 300 m, 22/07/ 1977, Tixier 10743, TAN. Lopholejeunea borbonica Steph. (Grolle 1995, Pócs & Geissler, 2002) Lopholejeunea erugata B. Thiers (Grolle, 1995) End, Type – 1896, Forsyth-Major, holotype G. 216 L. Marline et al. Lopholejeunea eulopha (Taylor) Schiffn. (Grolle, 1995; Pócs & Geissler, 2002; Zhu & Gradstein, 2005) Lopholejeunea grandicrista Steph. (Grolle, 1995; Pócs & Geissler, 2002) Lopholejeunea leioptera Gyarmati (Sass-Gyarmati, 2001; Pócs & Geissler, 2002) End, Type – Fianarantsoa, alt. 1590 m, S 22˚11’00” E 46˚53’00”, 19/09/ 1994, Magill & Pócs 12855, holotype MO. Lopholejeunea lepidoscypha Kiaer et Pearson (Grolle, 1995) End, Type – Ankaratra, 1877, Borgen s.n., holotype BM. Lopholejeunea multilacera Steph. (Grolle 1995) Lopholejeunea nigricans (Lindenb.) Schiffn. (Grolle, 2001; Zhu & Gradstein, 2005) Lopholejeunea onraedtii Vanden Berghen (Grolle, 2001) End, Type – Route de Moramanga, Tamatave, 17 km from Moramanga, forêt primaire humide et dégradée, alt. 800 m, 01/1970, Onraedt 70M2030, holotype BR. Lopholejeunea subfusca (Nees) Schiffn. (Grolle, 1995; Pócs & Geissler, 2002; Zhu & Gradstein, 2005) Lopholejeunea tixieriana Vanden Berghen (Grolle, 1995) End, Type – Antanidetra, 07/1977, Tixier 10720, holotype BR. Lopholejeunea utriculata Steph. (Grolle, 1995) End, Type – unknown. Marchesinia excavata (Mitt.) Schiffn. (Pócs & Geissler, 2002) Marchesinia madagassa Steph. (Grolle, 1995) Mastigolejeunea auriculata (Wilson) Schiffn. (Grolle, 1995) Mastigolejeunea rhodesica (Vanden Berghen) E.W. Jones (Grolle, 1995) Metalejeunea cucullata (Reinw. et al.) Grolle (Grolle, 1995) Microlejeunea africana Steph. (Grolle, 1995) Microlejeunea fissistipula Steph. (Grolle, 1995; Pócs & Geissler, 2002) End, Type – unknown. Microlejeunea inflata Steph. (Grolle, 1995; Pócs & Geissler, 2002) Microlejeunea oblongistipula (Gottsche) Pearson (Grolle, 1995) Microlejeunea papulosa (Gottsche) Pearson (Grolle, 1995) End, Type – unknown. Odontolejeunea lunulata (F. Weber) Schiffn. (Grolle, 1995; Pócs, 1995; Pócs & Geissler, 2002) Otolejeunea moniliata Grolle (Grolle, 1985; Grolle, 1995) End, Type – Partie nord du massif du Morojejy, alt. 1450 m, Guillaumet 4133/7, holotype JE, isotypes PC, BR. Otolejeunea rabenorii Tixier (Grolle, 1995) End, Type – Toamasina, route de Lakato, épiphylle en forêt dense à graminées, 02/06/1979, Tixier 12591, holotype PC. Otolejeunea subana Pócs (Pócs, 2004) End, Type – Toamasina Province, Mantadia Forest Reserve, North of Andasibe (Périnet), submontane rainforest, alt. 1030 m, S 18˚52’20” E 48˚28’26”, 02/10/1994, Pócs & Szabó 9485/R, holotype EGR. Prionolejeunea grata (Gottsche) Schiffn. (Grolle, 1995; Pócs & Geissler, 2002) Ptychanthus africanus Steph. (Grolle, 1995; Ahonen et al., 2005) Pycnolejeunea contigua (Nees) Grolle (Grolle, 1995; He, 1999) Schiffneriolejeunea madagascariensis (Steph.) Gradst. (Grolle, 1995) End, Type – unknown. Schiffneriolejeunea pappeana (Nees) Gradst. (Grolle, 1995; Pócs & Geissler, 2002) Checklist of the bryophytes of Madagascar 217 Schiffneriolejeunea pappeana var. bidentata Gradst. et Vanden Berghen (Grolle, 1995) Schiffneriolejeunea pappeana var. integra Gradst. et Vanden Berghen (Grolle, 1995) Schiffneriolejeunea parviloba (Steph.) Gradst. (Pócs & Geissler, 2002) Schiffneriolejeunea polycarpa (Nees) Gradst. (Grolle, 1995; Pócs & Geissler, 2002) Symbiezidium barbiflorum (Lindenb. et Gottsche) A. Evans (Sass-Gyarmati & Pócs, 2002) Symbiezidium madagascariense Steph. (Gradstein & Van Beek, 1985; Grolle, 1995; Pócs, 1995) Taxilejeunea conformis (Mont. et Nees) Steph. (Grolle, 1995; Pócs & Geissler, 2002) Taxilejeunea furcicornuta Grolle (Grolle, 1995) Taxilejeunea sikorae Steph. (Grolle, 1995) End, Type – unknown. Thysananthus spathulistipus (Reinw. et al.) Lindenb. (Grolle, 1995; Pócs & Geissler, 2002) – Marojejy National Park, alt. 650 m, 24/11/2009, Ah-Peng 349, det. Ah-Peng, Bardat. Xylolejeunea grolleana (Pócs) X.-L.He et Grolle (Pócs, 1999; He & Grolle, 2001) Lepidoziaceae Amazoopsis diplopoda (Pócs) J.J. Engel et G.L.S. Merr. (Grolle, 1995; Pócs, 1995) Bazzania curvidens Steph. (Grolle, 1995) End, Type – Alakaty forest, 30/11/1889, Perrot, herb (N.S.). Bazzania deciduifolia Onr. (Onraedt, 1977; Grolle, 1995) End, Type – Province de Fianarantsoa, forêt d’Ambatofitorahana, Onraedt 71M5364, holotype NAM, isotypes BR, JE. Bazzania decrescens (Lehm. et Lindenb.) Trevis. (Grolle, 1995) Bazzania decrescens subsp. pumila (Mitt.) Pócs (Pócs 1995) Bazzania decrescens var. ambahatrae Pócs (Pócs & Geissler, 2002) End, Type – Manongarivo massif, alt. 1300-1400 m, Pócs & Geissler PG 19742, holotype EGR. Bazzania mascarena (Steph.) Herzog (Grolle, 1995) Bazzania nitida (F. Weber) Grolle (Grolle, 1995; Pócs, 1995; Pócs & Geissler, 2002) Bazzania orbanii Pócs (Pócs, 2010b) End, Type – SW Madagascar, Toliara Province, Isalo National Park, “Piscine naturelle”, a deep pool of a brook in a sandstone gorge, shaded by Pandanus pulcher trees, on soil, alt. 800 m, Orbán 9455/J, holotype EGR, isotypes MO, TAN. Bazzania perrotana (Steph.) E.W. Jones (Jones, 1975; Grolle, 1995) Bazzania praerupta (Reinw. et al.) Trevis. (Grolle, 1995; Pócs, 1995) Bazzania reflexa (Gottsche) Steph. (Grolle, 1995) End, Type – unknown. Kurzia capillaries (Sw.) Grolle subsp. capillaris (Pócs, 1995; Pócs & Geissler, 2002) Kurzia capillaris (Renauld ex Steph.) Pócs subsp. stephanii (Grolle, 1995; Pócs, 1995; Pócs & Geissler, 2002) Lepidozia africana Steph. (Grolle, 1995) Lepidozia succida Mitt. (Grolle, 1995; Pócs, 1995) Telaranea bischleriana Pócs (Pócs, 2006) 218 L. Marline et al. Telaranea coactilis (Spruce) J.J. Engel et G.L.S. Merr. (Engel & Merrill, 2004) Telaranea nematodes (Gottsche ex Austin) M. Howe (Grolle, 1995) Lophocoleaceae Chiloscyphus coadunatus (Sw.) J.J. Engel et R.M. Schust. (Grolle, 1995; Pócs & Geissler, 2002) Chiloscyphus concretus (Mont.) J.J. Engel et R.M. Schust. (Grolle, 1995; Pócs & Geissler, 2002) Chiloscyphus difformis (Nees) J.J. Engel et R.M. Schust. (Pócs & Geissler, 2002) Chiloscyphus lucidus (Spreng. ex Lehm.) J.J. Engel et R.M. Schust. (Grolle, 1995; Pócs & Geissler, 2002) Chiloscyphus madagascariensis (Gottsche) J.J. Engel et R.M. Schust. (Grolle, 1995) End, Type – unknown. Chiloscyphus martianus (Nees) J.J. Engel et R.M. Schust. (Grolle, 1995; Pócs & Geissler, 2002) Chiloscyphus muricatus (Lehm.) J.J. Engel et R.M. Schust. (Grolle, 1995; Pócs & Geissler, 2002) Chiloscyphus regularis Steph. (Grolle, 1995) Clasmatocolea vermicularis (Lehm.) Grolle (Engel, 1980) Conoscyphus trapezioides (Sande Lac.) Schiffn. (Grolle, 1995; Pócs, 1995; Pócs & Geissler, 2002) Heteroscyphus dubius (Gottsche) Schiffn. (Grolle, 1995) Heteroscyphus splendens (Lehm. et Lindenb.) Grolle (Pócs, 1995; Pócs & Geissler, 2002) Leptoscyphus expansus (Lehm.) Grolle (Grolle, 1995) Leptoscyphus infuscatus (Mitt.) E.W. Jones (Pócs & Geissler, 2002) Marchantiaceae Marchantia debilis Goebel (Grolle, 1995) Marchantia pappeana Lehm. subsp. pappeana (Grolle, 1995) Mastigophoraceae Mastigophora diclados (Brid. ex F. Weber) Nees (Grolle, 1995; Pócs & Geissler, 2002) Metzgeriaceae Metzgeria consanguinea Schiffn. (So, 2004) Metzgeria furcata (L.) Dumort. (So, 2004) Metzgeria leptoneura Spruce (Grolle, 1995; So, 2004) Metzgeria madagassa Steph. (Grolle, 1995; So, 2004) Metzgeria nudifrons Steph. (So, 2004) Metzgeria quadrifaria Steph. (So, 2004) Pallaviciniaceae *Pallavicinia lyellii (Hook.) – Ambavaniasy, réserve de Vohimana, forêt ombrophile d’altitude, jonction sol-racine au pied d’une micro falaise, alt. 870 m, S 18˚55’19.5’’ E 48˚30’05.7’’, 14/04/2010, Reeb, Andriamananatena & Bidault CRAE 40 & CRAE 48, TAN, det. Reeb & Andriamananatena; Ambavaniasy, réserve de Vohimana, bord de chemin vers relais du naturaliste, terre tassée, alt. 790 m, S 18˚55’14.95’’ E 48˚30’49.58’’, 16/04/2010 Reeb, Andriamananatena & Bidault CRAE 155, TAN, det Reeb & Andriamananatena. Checklist of the bryophytes of Madagascar 219 Symphyogyna brasiliensis Nees et Mont. (Grolle, 1995) *Symphyogyna aff. podophylla (Thunb.) Mont. et Nees. – Ambavaniasy, réserve de Vohimana, grande cascade microfalaise à 3 m du bord, face verticale, alt. 849 m, S 18˚55’43.6’’ E 48˚29’56.6’’, 15/04/2010, Reeb, Andriamananatena & Bidault CRAE 87, TAN, det. Reeb & Andriamananatena. Symphyogynopsis gottscheana (Mont. et Nees) Grolle (Grolle, 1995) Plagiochilaceae Plagiochila angusta Lindenb. (Grolle, 1995; Pócs & Geissler, 2002) Plagiochila barteri Mitt. var. barteri (Grolle, 1995; Pócs & Geissler, 2002) Plagiochila boivinii Steph. (Grolle, 1995) Plagiochila boryana Gottsche ex Steph. (Grolle, 1995) Plagiochila cambuena Steph. (Grolle, 1995) Plagiochila drepanophylla Sande Lac. (Grolle, 1995) Plagiochila fracta Pócs, nom. inval. (Pócs & Geissler, 2002; Heinrichs et al 2005) Plagiochila granditexta Steph. (Grolle, 1995) End, Type – Hildebrandt, herb (N.S.). Plagiochila incerta Gottsche (Grolle, 1995; Pócs & Geissler, 2002) Plagiochila integerrima Steph. (Grolle, 1995) Plagiochila kiaeri Gottsche (Grolle, 1995; Pócs & Geissler, 2002) Plagiochila paucidentata Mont. et Gottsche (Grolle, 1995; Pócs & Geissler, 2002) Plagiochila pectinata Willd. ex Lindenb. (Grolle, 1995; Pócs & Geissler, 2002) Plagiochila punctata (Taylor) Taylor (Grolle, 1995; Pócs & Geissler, 2002; Heinrichs et al., 2005) Plagiochila repanda (Schwaegr.) Lindenb. (Grolle, 1995) Plagiochila repanda var. perrotana (Steph.) Vanden Berghen (Grolle, 1995) Plagiochila rodriguezii Steph. (Grolle, 1995) Plagiochila squamulosa var. crispulo-caudata (Gottsche) Vanden Berghen (Grolle, 1995) Plagiochila squamulosa var. sinuosa (Mitt.) Vanden Berghen (Grolle, 1995) Plagiochila stricta Lindenb. (Lindner et al., 2004) Plagiochila terebrans Nees et Mont. ex Lindenb. (Grolle, 1995; Pócs & Geissler, 2002) Pleuroziaceae Pleurozia gigantea (F. Weber) Lindb. (Pócs & Geissler, 2002) Porellaceae Porella abyssinica var. hoehnelii (Steph.) Pócs (Grolle, 1995) Porella cucullistipula Steph. (Grolle, 1995) Porella madagascariensis (Nees et Mont.) Trevis. (Grolle, 1995) Porella montantii (Steph.) E.W. Jones (Grolle, 1995) Porella triquetra (Steph.) E.W. Jones (Grolle, 1995) Radulaceae Radula ankefinensis Gottsche ex Steph. (Grolle, 1995; Grolle, 2001; Pócs & Geissler, 2002) Radula appressa Mitt. (Grolle, 1995; Pócs, 1995; Pócs & Geissler, 2002) Radula boryana (F. Weber) Mont. (Grolle, 1995; Grolle, 2001; Pócs & Geissler, 2002) Radula comorensis Steph. (Pócs & Geissler, 2002) 220 L. Marline et al. Radula fulvifolia (Hook.f. et Taylor) Gottsche et al. (Grolle, 2001) Radula madagascariensis Gottsche (Yamada, 1975; Grolle, 1995; Pócs, 1995; Grolle, 2001) Radula marojezica E.W. Jones (Grolle, 1995) End, Type – Province Antsiranana, Parc National de Marojejy, Pócs 90114/BN, holotype E, isotype and paratype EGR. Radula quadrata Gottsche (Grolle, 1995) Radula silvestris Gottsche (Grolle, 1995) Radula stenocalyx Mont. (Grolle, 1995; Pócs, 1995; Grolle, 2001; Pócs & Geissler, 2002) Radula voluta Taylor ex Gottsche et al. (Grolle, 1995; Pócs, 1995; Grolle, 2001; Pócs & Geissler, 2002) Ricciaceae Riccia atromarginata var. jovet-astiae Rauh & Buchloh (Grolle, 1995) Riccia cavernosa Hoffm. (Grolle, 1995) Riccia crinita Taylor (Grolle, 1995) Riccia perssonii Sultan Khan (Perold, 1995) *Riccia purpurascens Lehm. – Makirovana, Talus vertical en bord de rizière, Ambodimanga au pied de Makirovana, alt. 41 m, S 14˚11’13.6” E 49˚59’09.2”, 03/05/2010, Reeb CR 203, det. Reeb & Bardat. Scapaniaceae Anastrophyllum piligerum (Nees) Steph. (Grolle, 1995; Pócs, 1995; Pócs & Geissler, 2002) Andrewsianthus aberrans (Nees et Mont.) Grolle (Pócs, 1995) Plicanthus birmensis (Steph.) R.M. Schust. (Grolle, 1995) Plicanthus hirtellus (F. Weber) R.M. Schust. (Grolle, 1995; Pócs & Geissler, 2002) Schistochilaceae Gottschea neesii (Mont.) R.M. Schust. (Grolle, 1995; Pócs, 1995; Pócs & Geissler, 2002) Schistochila alata (Lehm.) Schiffn. (Grolle, 1995; Pócs, 1995; Pócs & Geissler, 2002) Schistochila piligera Steph. (Grolle, 1995; Pócs, 1995; Pócs & Geissler, 2002) End, Type – Camboué, herb (N.S.). Targioniaceae Targionia hypophylla L. (Grolle, 1995) *Targionia dioica Schiffn. – Biosphere Reserve and National Park, MananaraNord, Toamasina, alt. 220 m, 17/08/1998, Pócs & Szabó 9878/CY, EGR. Hornworts reported from Madagascar Anthocerotaceae Anthoceros sp. (Grolle, 1995) *Dendroceros cf. africanus Steph. – Réserve Spéciale Angavokely, alt. 1750 m, 11/2009, Bardat JB MAD 147, det. Bardat. Notothyladaceae Phaeoceros carolinianus (Michx.) Prosk. (Grolle, 1995) Checklist of the bryophytes of Madagascar 221 Mosses reported from Madagascar Amblystegiaceae Amblystegium chalaropelma Müll.Hal. ex Renauld (Crosby et al., 1983) End, Type – Zone du plateau central, Imerina Oriental, Hildebrandt 2076, herb. (N.S.). Amblystegium riparium (Hedw.) Bruch, Schimp. et W.Guembel (Crosby et al., 1983) Hygrohypnum caussequei (Renauld & Cardot) Cardot (Crosby et al., 1983) End, Type – Entre Tananarive et Betafo, 1894, Caussèque, holotype PC. Andreaeaceae Andreaea subulata Harv. (Crosby et al., 1983) Andreaea tsaratananae Thér. (Crosby et al., 1983) Anomodontaceae Herpetineuron toccoae (Sull. et Lesq.) Cardot (Touw, 1993) Archidiaceae *Archidium ohioense Schimp. ex Müll.Hal. – Beza mahafaly, Betioky, Riverine forest adjacent to Sakamena river, open ground on forest floor, alt. 130 m, S 23˚39’ E 44˚38’, 18/05/1987, Phillipson 1818, TAN. *Archidium subulatum Müll.Hal. – Beloha Tsihombe, dalle de gneiss, 10/04/1971, Cornet & Cremers 2014, TAN. Bartramiaceae Bartramia gigantea Bory (Crosby et al., 1983) Bartramia longifolia Hook. (Crosby et al., 1983) Breutelia diffracta Mitt. (Crosby et al., 1983) Breutelia gnaphalea P. Beauv. (Crosby et al., 1983) Breutelia madagassa Thér. var. madagassa (Crosby et al., 1983) End, Type – Mont Tsaratanana, alt. 1200 m, 04/1924, Perrier de la Bathie, holotype PC. Breutelia microdonta (Mitt.) Broth. (Crosby et al., 1983; Virtanen, 1997) Breutelia perrieri Thér. (Crosby et al., 1983) Breutelia sciuroides Thér. (Crosby et al., 1983) Breutelia stuhlmannii Broth. (De Sloover, 1975; Crosby et al., 1983) Breutelia viguieri Cardot (Crosby et al., 1983) End, Type – Ambatolampy, ruisselet près du sommet du Tsiafajavona, alt. 2575 m, 28/11/1912, Viguier 1688ter, holotype PC. Philonotis byssiformis Müll.Hal. (Crosby et al., 1983) End, Type – Zone inférieure des forêts, Nosy Be, Hildebrandt, holotype PC. Philonotis hastata (Duby) Wijk et Margad. (Crosby et al., 1983) Philonotis mauritiana Aongstr. (Crosby et al., 1983) Philonotis mauritiana var. stricta Renauld et Cardot (Crosby et al., 1983) End, Type – Plateau central, environs de Tananarive, Camboué; Entre Tananarive et Betafo, Caussèque. Fianarantsoa, Besson; Ambositra, Soula; Mont Ambohijanahary, PC, holotype (N.S.). Philonotis perpusilla Cardot (Crosby et al., 1983) End, Type – Zone de savane ocidentale, Marovoay, Mathieu, herb (N.S.). Philonotis perrieri Thér. (Crosby et al., 1983) Philonotis sparsifolia (Hampe) A. Jaeger (Crosby et al., 1983) 222 L. Marline et al. Brachytheciaceae Aerolindigia capillacea (Hornsch.) M. Menzel (Crosby et al., 1983) Brachythecium borgenii (Hampe) A. Jaeger (Crosby et al., 1983) Brachythecium decurrens Cardot (Crosby et al., 1983) Eurhynchium spinulinerve Kiaer ex Cardot (Crosby et al., 1983) Eurhynchium striatum (Schreb. ex Hedw.) Schimp. (Crosby et al., 1983) Helicodontium fabroniopsis Müll. Hal. (Crosby et al., 1983) End, Type – Zone supérieure des forêts, troncs d’arbres à Andrangoloaka, Sikora, herb (N.S.). Palamocladium leskeoides (Hook.) E. Britton (Crosby et al., 1983; Hofmann, 1997) Rhynchostegiella microtheca (Renauld et Cardot) Broth. (Crosby et al., 1983) Rhynchostegium angustifolium Renauld & Cardot (Crosby et al., 1983) End, Type – Zone moyenne des forêts, forêt d’Analamazaotra, Camboué & Campenon, herb (N.S.). Rhynchostegium comorae (Müll.Hal.) A. Jaeger (Crosby et al., 1983) Rhynchostegium distans Besch. (Crosby et al., 1983) Rhynchostegium nervosum (Renauld) Cardot (Crosby et al., 1983) End, Type – Zone du Plateau central, monts Ankaratra, Borgen 61, lectotype BM, isolectotype THR. Rhynchostegium pseudodistans Cardot (Crosby et al., 1983) Schimperella rhynchostegioides Thér. (Crosby et al., 1983) End, Type – Mont Tsaratanana, alt. 1200 m, 04/1924, Perrier de la Bathie 176 p.p., lectotype PC. Squamidium brasiliense (Hornsch.) Broth. (Crosby et al., 1983; Allen & Crosby, 1986; Delgadillo, 1995) Bruchiaceae Trematodon curvicollis Cardot (Crosby et al., 1983) End, Type – Entre Savondronina et Ranomafana, Besson, holotype PC. Trematodon decaryi Thér. (Crosby et al., 1983) End, Type – Vallée de l’Anony, sur talus humide, 05/08/1921, Decary, holotype PC. Trematodon lacunosus Renauld & Cardot (Crosby et al., 1983) End, Type – Entre Savondronina et Ranomafana, 1892, Besson, PC. Trematodon longicollis Michx. (Crosby et al., 1983) Trematodon platybasis Müll.Hal. (Crosby et al., 1983) End, Type – Alasora, Ambohipo, Tananarive, alt. 1250 m, Camboué, holotype PC. Trematodon reticulatus Müll.Hal. ex Geh. (Crosby et al., 1983) End, Type – Zone moyenne des forêts, à l’Est d’Ambatondrazaka, Rutemberg, herb (N.S.). Trematodon viguieri Cardot (Crosby et al., 1983) Bryaceae Anomobryum filiforme var. madagassum Renauld et Cardot (Crosby et al., 1983) End, Type – Zone de plateau central, Imerina, Camboué, herb (N.S.). Anomobryum julaceum (P. Gaertn., B. Mey. et Scherb.) Schimp. var. julaceum (Ochi, 1972; Crosby et al., 1983) Brachymenium acuminatum Harv. (Ochi, 1972; Crosby et al., 1983; Delgadillo, 1995) Brachymenium apiculatum P. Beauv. (Crosby et al., 1983) Brachymenium capitulatum (Mitt.) Kindb. (Crosby et al., 1983) Checklist of the bryophytes of Madagascar 223 Brachymenium exile (Dozy et Molk.) Bosch et Sande Lac. (Ochi, 1972; Crosby et al., 1983; Delgadillo, 1995) Brachymenium leptophyllum (Bruch et Schimp. ex Müll.Hal.) Bruch & Schimp. ex A. Jaeger (Crosby et al., 1983) Brachymenium madagassum Hampe (Ochi, 1972; Crosby et al., 1983) End, Type – Plateau central, 1869, Borgen, holotype PC. Brachymenium nepalense Hook. (Ochi, 1972; Crosby et al., 1983; Pócs & Geissler, 2002) Brachymenium nigrescens Besch. (Ochi, 1972; Crosby et al., 1983) Brachymenium notarisii (Mitt.) A.J. Shaw (Crosby et al., 1983) Brachymenium philonotula Broth. (Ochi, 1972; Crosby et al., 1983) Brachymenium pulchrum Hook. (Ochi, 1972; Crosby et al., 1983; Tixier, 1995) Brachymenium rigidum Broth. et Paris (Crosby et al., 1983) Bryum albo-marginatum Cardot (Ochi, 1972; Crosby et al., 1983) End, Type – Mont Tsaratanana, alt. 1200-2400 m, 4/1924, Perrier de la Bathie s. n., holotype PC# Bryum alpinulum var. levieri Renauld et Cardot (Crosby et al., 1983) End, Type – Zone inférieure des forêts, forêt de Fito, Perrot, holotype FI# Bryum alpinum Huds. ex With. var. alpinum (Ochi, 1972; Ochi, 1973; Crosby et al., 1983) Bryum apiculatum Schwaegr. (Ochi, 1972; Crosby et al., 1983) Bryum arachnoideum Müll.Hal. (Ochi, 1972; Crosby et al., 1983) Bryum argenteum Hedw. var. argenteum (Ochi, 1972; Crosby et al., 1983; Pócs & Geissler, 2002) Bryum aubertii (Schwaegr.) Brid. (Ochi, 1972; Crosby et al., 1983) Bryum billardierei Schwaegr. var. billardierei (Ochi, 1972; Mohamed, 1979; Crosby et al., 1983) Bryum caespiticium Hedw. var. caespiticium (Pócs & Geissler, 2002) Bryum capillare Hedw. var. capillare (Ochi, 1972; Crosby et al., 1983) Bryum coronatum Schwaegr. (Ochi, 1972; Crosby et al., 1983; Pócs & Geissler, 2002) Bryum dichotomum Hedw. (Crosby et al., 1983) Bryum erythrocaulon (Schwäegr.) Brid. (Mohamed, 1979; Crosby et al., 1983) Bryum huillense Welw. et Duby (Ochi, 1972; Crosby et al., 1983; Pócs & Geissler, 2002) Bryum imerinanus O’Shea (O’Shea, 2008) Bryum leptospeiron Müll.Hal. (Crosby et al., 1983) Bryum megalacrion Schwaegr. (Ochi, 1972; Crosby et al., 1983) Bryum neelgheriense var. wichurae (Broth.) Mohamed (Pócs & Geissler, 2002) Bryum paradoxum Schwaegr. (Ochi, 1972; Crosby et al., 1983; Delgadillo, 1995) Bryum perrieri Thér. (Crosby et al., 1983) End, Type – Forêt du centre, alt. 2000 m, Perrier de la Bathie 12127p.p, holotype PC. Bryum pseudotriquetrum (Hedw.) P. Gaertn., B. Mey. et Scherb. var. pseudotriquetrum (Crosby et al., 1983) Bryum purpureonigrum Duby (Crosby et al., 1983) End, Type – unknown. Bryum savondroninicus O’Shea (Crosby et al., 1983; O’Shea, 2008) End, Type – Between Savondronina and Ranomafana, Besson, herb (N.S.). Bryum subargenteum Hampe (Ochi, 1972; Crosby et al., 1983) Bryum viguieri Thér. (Ochi, 1972; Crosby et al., 1983) 224 L. Marline et al. Bryum voeltzkowii Broth. (Ochi, 1972; Mohamed, 1979; Crosby et al., 1983) Mniobryum perlaxum Thér. et P. de la Varde (Crosby et al., 1983) End, Type – Totohaza, 05/1931, Grainville, holotype PC. Rhodobryum commersonii (Schwaegr.) Brid. (Crosby et al., 1983; Pócs & Geissler, 2002) Rhodobryum giganteum (Schwaegr.) Paris (Crosby et al., 1983; Ochi, 1972) Rhodobryum ontariense (Kindb.) Paris (Crosby et al., 1983) Calymperaceae Calymperes afzelii Sw. (Crosby et al., 1983; Pócs & Geissler, 2002) Calymperes erosum Müll.Hal. (Crosby et al., 1983) Calymperes hispidum Renauld et Cardot (Crosby et al., 1983; Ellis, 1988) Calymperes loucoubense Besch. (Crosby et al., 1983) End, Type – Zone inférieure des forêts, Nosy Be, Lokobe, 1879, Marie, holotype BM. Calymperes palisotii Schwaegr. (Crosby et al., 1983) Calymperes pallidum Mitt. (Crosby et al., 1983; Delgadillo, 1995) *Calymperes sanctae-mariae var. mayottense Besch. – Mananjary, tronc de Pandanus, 14/07/1972, Cornet & Cremers 2250, TAN. Calymperes taitense (Sull.) Mitt. (Crosby et al., 1983; Orbán, 1995) Calymperes tenerum Müll.Hal. (Crosby et al., 1983) Calymperes venezuelanum (Mitt.) Broth. ex Pittier (Orbán, 2000) Leucophanes hildebrandtii Müll.Hal. (Crosby et al., 1983; Salazar Allen, 1993) Leucophanes renauldii Cardot (Crosby et al., 1983; Salazar Allen, 1993; Pócs & Geissler, 2002) Leucophanes rodriguezii Müll.Hal. (Salazar Allen, 1993) Mitthyridium fasciculatum (Hook. et Grev.) H. Rob. var. fasciculatum (Reese, 1991) Mitthyridium subflavum (Renauld & Cardot) H. Rob. (Müller, 1897; Crosby et al., 1983) Octoblepharum africanum (Broth.) Cardot (Crosby et al., 1983) Octoblepharum albidum Hedw. (Crosby et al., 1983; Pócs & Geissler, 2002) Syrrhopodon africanus (Mitt.) Paris var. africanus (Crosby et al., 1983) Syrrhopodon africanus var. mandrakensis (Tixier) W.D. Reese (Tixier, 1992; Pócs & Geissler, 2002) Syrrhopodon albidus (E.B. Bartram) L.T. Ellis subsp. integrifolius (Pócs & Geissler, 2002) Syrrhopodon apertifolius Besch. (Ellis, 1995) Syrrhopodon armatissimus W.D. Reese (Reese, 1991; Orbán, 1995) Syrrhopodon armatus (Bizot et Onr.) Orbán and W.D. Reese subsp. insularus (Orbán, 1977) Syrrhopodon asper Mitt. (Een, 1993) Syrrhopodon cuneifolius Thér. (Orbán, 1981; Crosby et al., 1983; Pócs & Geissler, 2002) End, Type – Massif de l’Ankaizina, alt. 1200 m, 04/1923, Decary, holotype PC. Syrrhopodon dimorphophyllus L.T. Ellis (Ellis, 2005) Syrrhopodon gardneri (Hook.) Schwaegr. (Orbán, 1981; Crosby et al., 1983; Ellis, 2002; Pócs & Geissler, 2002) Syrrhopodon gaudichaudii Mont. (Orbán, 1981; Crosby et al., 1983; Ellis, 2005) Syrrhopodon graminifolius Renauld et Cardot (Orbán, 1981; Crosby et al., 1983) Checklist of the bryophytes of Madagascar 225 Syrrhopodon hispidocostatus Renauld et Cardot in Renauld (Orbán, 1978; Orbán, 1981; Crosby et al., 1983; Pócs & Geissler, 2002; Ellis, 2003; Ellis, 2005) Syrrhopodon involutus Schwaegr. (Orbán, 1981; Crosby et al., 1983) Syrrhopodon mauritianus Müll.Hal. ex Aongstr. (Orbán, 1981; Crosby et al., 1983) Syrrhopodon parasiticus (Sw. ex Brid.) Besch. (Crosby et al., 1983) Syrrhopodon pottioides Orbán (Orbán, 1995) Syrrhopodon prolifer var. acanthoneuros (Müll.Hal.) Müll.Hal. (Orbán, 1981; Crosby et al., 1983; Ellis, 2003) Syrrhopodon prolifer Schwaegr. var. prolifer (Orbán, 1981; Crosby et al., 1983; Delgadillo, 1995) Syrrhopodon pulcher W.D. Reese (Orbán, 1995) Syrrhopodon spiralis Renauld et Cardot (Ellis, 2005) Syrrhopodon tixieri W.D. Reese (Tixier, 1992) End, Type – Ranomafana, Fianarantsoa, route de Mananjary, pk 24, corticole, alt. 900 m, Tixier 10152, holotype PC, paratype MO. Syrrhopodon vardei L.T. Ellis (Ellis, 2005) Catagoniaceae Catagonium nitens (Brid.) Cardot subsp. nitens (Crosby et al., 1983) Cryphaeaceae Cryphaea jamesonii Taylor (Crosby et al., 1983; Rao, 2001) Cryphaea rutenbergii Müll.Hal. (Crosby et al., 1983; Rao, 2001) Schoenobryum concavifolium (Griff.) Manuel (Crosby et al., 1983; O’Shea, 2003) Daltoniaceae Adelothecium bogotense Hampe (Crosby et al., 1983; Delgadillo, 1995) Calyptrochaeta asplenioides (Brid.) Crosby (Crosby et al., 1983) Daltonia angustifolia Dozy et Molk. var. angustifolia (Crosby et al., 1983) Daltonia cardotii Bizot et Onr. (Crosby et al., 1983) Daltonia forsythii Cardot (Crosby et al., 1983) End, Type – Zone moyenne des forêts, forêt d’Ambohimitombo, alt. 1350 m, 1895, Forsyth Major, holotype PC. Daltonia latimarginata Besch. (Crosby et al., 1983) Daltonia latimarginata var. madagassa Renauld (Crosby et al., 1983) *Distichophyllum mascarenicum Besch. – Marojejy National Park, scattered elfin forest, soil bank above the stream, S 14˚26’ E 49˚45’, alt. 1050 m, Magill, Pócs & La Farge 9971, TAN. Distichophyllum rakotomariae Crosby (Crosby et al., 1983; Touw, 1993) End, Type – Montagne d’Ambre, Diégo-Suarez, alt. 900-1100 m, M.R. & C.A Crosby 6883, holotype MO, isotypes PC, TAN. Dicranaceae Anisothecium madagassum Thér. (Crosby et al., 1983) End, Type – Montagne d’Ambre, forêt, alt. 1000-1200 m, 10/1926, Perrier de La Bathie, holotype PC. Aongstroemia filiformis (P. Beauv.) Wijk et Margad. (Crosby et al., 1983; Delgadillo, 1995) Aongstroemia julacea (Hook.) Mitt. (Crosby et al., 1983) Dicranella acroclada Cardot (Crosby et al., 1983) 226 L. Marline et al. Dicranella borbonica var. madagascariensis Thér. (Crosby et al., 1983) End, Type – Montagne d’Ambre, Forêt du versant oriental, 1899, Girod, holotype PC. Dicranella madagassa Renauld (Crosby et al., 1983) Dicranella pervilleana Besch. (Crosby et al., 1983) Dicranella polii Renauld et Cardot (Crosby et al., 1983) Dicranella subsubulata (Hampe ex Müll.Hal.) A. Jaeger (Crosby et al., 1983) Dicranoloma billardierei (Brid. ex Anon.) Paris (Crosby et al., 1983) Dicranoloma billardieri var. patentifolium (Renauld et Paris) Thér. (Crosby et al., 1983) Dicranoloma billardieri var. scopareolum (Müll.Hal.) Thér. (Crosby et al., 1983) Dicranum johnstonii Mitt. (Pócs & Geissler, 2002) Holomitrium borbonicum Besch. (Crosby et al., 1983) Holomitrium borbonicum var. hamatum (Müll.Hal.) Thér. (Crosby et al., 1983) Holomitrium camptocarpum P. de la Varde (Crosby et al., 1983) End, Type – Pentes orientales du massif de Marojejy, à l’Ouest de la rivière Manantenina, affluent de la Lokoho, sylve à lichens sur gneiss et quartzite, alt. 1400 m, 15/12/1948, Humbert, holotype PC. Holomitrium cylindraceum var. cucullatum (Besch.) Wijk et Margad. (Crosby et al., 1983) Holomitrium cylindraceum (P. Beauv.) Wijk & Margad. var. cylindraceum (Crosby et al., 1983) Holomitrium cylindraceum var. obtusifolium (Besch.) Wijk et Margad. (Crosby et al., 1983) Holomitrium gracilisetum Thér. (Crosby et al., 1983) End, Type – Ankaizina, alt. 1200-1500 m, 04/1933, Decary, holotype PC. Holomitrium hillieri Thér. (Crosby et al., 1983) End, Type – Forêt de Manongarivo, entre Nosy Be et Analalava, alt. 600 m, S 13˚55’00” E 48˚24’00”, 1909, Perrier de la Bathie, holotype PC. Holomitrium lepervanchei Thér. (Crosby et al., 1983) Holomitrium rhyparodictyon P. de la Varde (Crosby et al., 1983) End, Type – Pente orientale du massif de Marojejy, sylve à lichens et végétation éricoide des sommets, affluent de la Lokoho, alt. 1500 m, 17/12/1948, Humbert, holotype PC. Holomitrium subglobosum Thér. (Crosby et al., 1983) End, Type – Ifanadina, forêt, herb (N.S.). Leptotrichella minuta (Hampe) Ochyra (Crosby et al., 1983) Leptotrichella pallidiseta (Brid.) Ochyra (Crosby et al., 1983) End, Type – unknown. Leucoloma albocinctum Renauld et Cardot (Crosby et al., 1983; La Farge, 2002c) Leucoloma allorgei Thér. (Crosby et al., 1983; La Farge, 2002c) End, Type – Befotaka, 20/08/1926, Decary, holotype PC. Leucoloma ambreanum Renauld et Cardot (Crosby et al., 1983; La Farge, 2002c) End, Type – Montagne d’Ambre, près de Diégo-Suarez, 1890, Chénagon, holotype PC. Leucoloma bifidum (Brid.) Brid. (Crosby et al., 1983) Leucoloma boivinianum Besch. var. boivinianum (Crosby et al., 1983) Checklist of the bryophytes of Madagascar 227 Leucoloma boivinianum var. brevifolium Renauld (Crosby et al., 1983) End, Type – Betsileo, entre Savondronina et Ranomafana, 1892, Besson, holotype PC. Leucoloma brevioperculatum Dixon (Crosby et al., 1983; La Farge, 2002c) End, Type – Vohémar, 1814, Thompson 30, holotype BM. Leucoloma brotheri Renauld (Crosby et al., 1983; La Farge, 2002c) *Leucoloma caespitulans (Müll.Hal.) A. Jaeger – Montagne des Français, bois mort, alt. 400 m, 25/08/1973, Cremers 2657, TAN. Leucoloma candidum Broth. (Crosby et al., 1983; La Farge, 2002c) End, Type – Sainte Marie, East Madagascar, 1904, Voeltzkow, holotype BR. Leucoloma capillifolium Renauld (Crosby et al., 1983; La Farge, 2002c) End, Type – Sainte Marie, Sahasifotro, troncs d’arbres, 1894, Darbould, lectotype PC. Leucoloma chrysobasilare (Müll.Hal.) A. Jaeger var. chrysobasilare (Crosby et al., 1983) Leucoloma cirrosulum Renauld (Crosby et al., 1983; La Farge, 2002c) End, Type – Forêt d’Analamazaotra, 1891, Camboué & Campenon, holotype PC. Leucoloma crosbyi La Farge-England (La Farge-England, 1992; La Farge, 2002a; La Farge, 2002c) End, Type – Diégo-Suarez, Montagne d’Ambre, Parc National, 10 km along trail between Petit Lac and Grand Lac, S 12˚34’ E 49˚12’, 1972, M.R. & C.A Crosby 7167, 13.xi. holotype MO, isotypes ALTA, TAN. Leucoloma cuneifolium (Hampe ex Müll.Hal. et Geh.) C.H. Wright (Crosby et al., 1983; La Farge, 2002b; La Farge, 2002c) Leucoloma decaryi Thér. (Crosby et al., 1983; La Farge, 2002c) End, Type – Massif d’Ankaizinana, 10/06/1923, Decary, holotype PC. Leucoloma delicatulum Renauld (Crosby et al., 1983; La Farge, 2002c) Leucoloma dichelymoides (Müll.Hal.) A. Jaeger (Crosby et al., 1983; La Farge, 2002a) Leucoloma fontinaloides Dixon (Crosby et al., 1983; La Farge, 2002a; La Farge, 2002c) End, Type – Ambohimitomba forest, Tanala, 05/01/1895, ForsythMajor 473 b, holotype BM. Leucoloma fuscifolium Besch. (Crosby et al., 1983) Leucoloma fuscifolium var. crispatulum Renauld (Crosby et al., 1983) Leucoloma garnieri Paris & Renauld (Crosby et al., 1983; La Farge, 2002c) End, Type – Ad radices arboris “Jisy”, Dictae secus rivulum “Pleurs des Bœufs”, 24/08/1904, Savelli, lectotype PC. Leucoloma garnieri f. aquaticum Thér. (Crosby et al., 1983; La Farge, 2002c) End, Type – Massif de l’Andringitra, 1921, Perrier de la Bathie, holotype PC. Leucoloma grandidieri Renauld et Cardot (Crosby et al., 1983; La Farge-England, 1992; La Farge, 2002a; La Farge, 2002c) End, Type – Forêt d’Analamazoatra, 1891, Camboué & Campenon, lectotype PC. Leucoloma grimmioides P. de la Varde (Crosby et al., 1983; La Farge, 2002b; La Farge, 2002c) Leucoloma humbertii P. de la Varde (Crosby et al., 1983; La Farge, 2002b; La Farge, 2002c) End, Type – Pentes orientales du massif de Marojejy, à l’ouest de la rivière Manantenina, affluent de la Lokoho sylve à lichens sur gneiss et quartzite, alt. 1400 m, 15/12/1948, Humbert, holotype PC. Leucoloma incrassatum Thér. (Crosby et al., 1983; La Farge, 2002c) End, Type – Massif de l’Ikongo, rochers humides en forêt, Decary, holotype PC. 228 L. Marline et al. Leucoloma lepervancheri Besch. (La Farge, 2002a; La Farge, 2002c) Leucoloma longifolium (Brid.) Wijk et Margad. (Crosby et al., 1983; La Farge, 2002c) Leucoloma madagascariense La Farge (La Farge, 2002a; La Farge, 2002c) End, Type – Province Tamatave, Andasibe (Périnet), Reserve Forestière, montane rainforest with shady, wet habitats along stream leading to lac vert, Canopy reaching 25 m, mostly closed, with tall palms and bamboo, alt. 920-980 m, S 18˚54’00” E 48˚25’3”, 16/03/1990, La Farge-England 5788, holotype ALTA, isotypes MO,TAN. Leucoloma marojeziense La Farge (La Farge, 2002a; La Farge, 2002b; La Farge, 2002c) End, Type – Province Diégo-Suarez, Tahirin-javaboahary Marojejy, north of Manantenina and Loko River Valley, slopes of the Marojejy massif above Mandena village, S 14˚26’ E 49˚44’, La FargeEngland 6298, holotype ALTA, isotypes MO, TAN. Leucoloma membranaceum La Farge (La Farge, 2002a) Leucoloma ochrobasilare Ren. (Crosby et al., 1983; La Farge, 2002c) Leucoloma ochrobasilare (Thér.) La Farge subsp. longifolium (Crosby et al., 1983; La Farge, 2002a; La Farge, 2002c) End, Type – Mont Tsaratanana, forêt à sous bois herbacé, alt. 1200-2400 m, 04/1924, Perrier de la Bathie, holotype PC. Leucoloma ochrobasilare Renauld subsp. ochrobasilare (Crosby et al., 1983; La Farge, 2002a) Leucoloma oncophorellum Müll.Hal. (Crosby et al., 1983; La Farge, 2002c) End, Type – lectotype PC. Leucoloma perrieri Thér. (Crosby et al., 1983; La Farge, 2002c) End, Type – Forêt à l’Est de Tananarive, 11/1927, Perrier de la Bathie, holotype PC, isotype FH. Leucoloma persecundum Besch. (Crosby et al., 1983) Leucoloma persecundum var. perrotii Renauld (Crosby et al., 1983; La Farge, 2002c) End, Type – Environs d’Andevoranto, 1895, Perrot, lectotype BM. Leucoloma pumilum C.H. Wright (Crosby et al., 1983) End, Type – Sur le Plateau central, à Alaby, près de Maritandrano, Rutenberg, herb (N.S.). Leucoloma pusillum Cardot (Crosby et al., 1983; La Farge, 2002c) End, Type – Diégo-Suarez, zone inférieure des forêts, sur écorces, Chénagon, lectotype PC. Leucoloma rutenbergii (Geh.) C.H. Wright (Crosby et al., 1983; La Farge, 2002c) End, Type – Ambatondrazaka, Analamazaotra, Rutenberg, lectotype PC. Leucoloma rutenbergii var. abbreviatum Renauld (Crosby et al., 1983) End, Type – Forêt d’Alakaty, Perrot; Forêt d’Analamazaotra, Camboué & Campenon; holotype (N.S.). Leucoloma rutenbergii var. elatum Renauld (Crosby et al., 1983; La Farge, 2002c) Leucoloma rutenbergii var. perrotii Renauld (Crosby et al., 1983; La Farge, 2002c) End, Type – Environs d’Andevoranto, 1895, Perrot, holotype PC, isotypes BR, FH. Leucoloma sanctae-mariae Besch. (Crosby et al., 1983; La Farge, 2002a; La Farge, 2002c) Leucoloma seychellense Besch. (Crosby et al., 1983; La Farge, 2002a; La Farge, 2002c) Leucoloma sinuosulum Müll.Hal. ex Besch. (Crosby et al., 1983) Checklist of the bryophytes of Madagascar 229 Leucoloma squarrosulum (Geh.) Wright (Crosby et al., 1983) End, Type – Forêt de Vondrozona, 11/1877, Rutenberg, holotype PC. Leucoloma subchrysobasilare Renauld (Crosby et al., 1983; La Farge, 2002a; La Farge, 2002c) End, Type – In montibus Ankaratra, 1877, Borgen, lectotype PC. Leucoloma talazaccii Renauld et Cardot (Crosby et al., 1983; La Farge, 2002a; La Farge, 2002c) End, Type – Ambondrombe, 1894, Talazac, holotype PC, isotypes BR, PC, S. Leucoloma thraustum Hampe ex Besch. (Crosby et al., 1983; La Farge, 2002a; La Farge, 2002c) End, Type – Alamazantraskoven, Borchgrevink, lectotype BM. Leucoloma thuretii Besch. (Crosby et al., 1983; La Farge, 2002a; La Farge, 2002c) End, Type – Côte Est, près de la cascade de Simiang, Bernier, lectotype BM. Leucoloma tuberculosum Renauld (Crosby et al., 1983; La Farge, 2002a; La Farge, 2002c) End, Type – Forêt d’Analamazaotra, 1891, Camboué & Campenon, holotype PC. Leucoloma villaumei Thér. (Crosby et al., 1983; La Farge, 2002c) End, Type – Betsileo, Villaume, holotype PC. Microcampylopus laevigatus (Thér.) Giese & J.-P. Frahm (Crosby et al., 1983; Giese & Frahm, 1985) Pseudephemerum nitidum (Hedw.) Reimers (Crosby et al., 1983) Ditrichaceae Ceratodon purpureus (Reichardt) Burley subsp. convolutus (Crosby et al., 1983; Burley & Pritchard, 1990; Een, 1993) Distichium capillaceum (Hedw.) Bruch, Schimp. et W. Guembel var. capillaceum (Crosby et al., 1983) Ditrichum difficile (Duby) M. Fleisch. (Crosby et al., 1983; Delgadillo, 1995) Ditrichum luteum Dixon et Thér. (Crosby et al., 1983) End, Type – Mont Tsaratanana, Perrier de la Bathie, holotype PC. Ditrichum madagassum (Renauld et Cardot) Paris (Crosby et al., 1983) End, Type –Andrangoloaka, 1891, Sikora, holotype PC. Ditrichum pallidum (Hedw.) Hampe (Crosby et al., 1983) Garckea flexuosa (Griff.) Margad. et Nork. (Crosby et al. 1983) Pleuridium nervosum (Hook.) Mitt. (Tixier, 1979b; Crosby et al., 1983) Entodontaceae Campylodontium flavescens var. viride Cardot (Crosby et al., 1983) End, Type – Plateau central, environs de Tananarive, Sibree, holotype BM# Entodon corbieri Renauld et Cardot (Crosby et al., 1983; Tixier, 1995) End, Type – Montagne d’Ambre, près de Diégo-Suarez, Leloutre, holotype CHE# Entodon felicis Renauld et Cardot (Crosby et al., 1983) End, Type – Fianarantsoa (zone du Plateau central), Félix, holotype PC. Entodon felicis var. julaceus Renauld et Cardot (Crosby et al., 1983) End, Type – Plateau central, Ambohimahatsara, près d’Ambositra, Berthieu, herb (N.S.). Entodon madagassus Geh. (Crosby et al., 1983) End, Type – Ambatondrazaka, Borgen 100, holotype PC. 230 L. Marline et al. Entodon rutenbergii Geh. (Crosby et al., 1983) End, Type – Mont Ankaratra, Borgen 101, holotype PC. Erythrodontium lacoutourei Renauld et Cardot (Crosby et al., 1983) End, Type – Fianarantsoa, 1905, Villaume, holotype BM# Mesonodon flavescens (Hook.) W.R. Buck (Crosby et al., 1983) Erpodiaceae Erpodium beccarii Müll.Hal. ex Venturi var. beccarii (Tixier, 1979b; Crosby et al., 1983) Erpodium madagassum Paris et Renauld (Crum, 1972; Crosby et al., 1983) End, Type –Antsirabe, vallée de la rivière Andrantsay, Galinon, PC, holotype (N.S.). Eustichiaceae Eustichia longirostris (Brid.) Brid. (Crosby et al., 1983) Fabroniaceae Fabronia campenonii Renauld et Cardot (Crosby et al., 1983) End, Type – Forêt d’Amperifery, alt. 1300 m, 06/1890, Campenon, holotype PC. Fabronia crassiretis Renauld et Cardot (Crosby et al., 1983) Fabronia fastigiata Renauld et Cardot (Crosby et al., 1983) End, Type – Zone du Plateau central, Fianarantsoa, 1891, Besson, holotype PC. Fabronia fastigiata var. asperula Renauld et Paris (Crosby et al., 1983) End, Type – Zone moyenne des forêts, cercle militaire des Barii, secteur d’Ivondro, 18/11/1900, Croll, holotype PC. Fabronia garnieri (Paris et Renauld) Renauld et Paris (Crosby et al., 1983) End, Type – Betafo, rive de la Mania, 1902, holotype PC. Fabronia lachenaudii Renauld (Crosby et al., 1983) End, Type –District de Fihaonana, 3/1901, Ab indig. Coll 30, holotype PC. Fabronia motelayi Renauld et Cardot (Crosby et al., 1983) End, Type – Tananarive, dans les jardins de la Résidence, sur les troncs, 15/04/1897, Dorr, holotype PC. Fabronia villaumii Renauld et Cardot (Crosby et al., 1983) End, Type – Fianarantsoa, 1905, Villaume 722, holotype PC. Rhizofabronia persoonii var. sphaerocarpa (Dusén) Bizot ex Ochyra (Crosby et al., 1983; Touw, 1993) Fissidentaceae Fissidens angustifolius Sull. (Bruggeman-Nannenga, 2009; Bruggeman-Nannenga & Arts, 2010) Fissidens aristifer Brugg.-Nann. (Bruggeman-Nannenga, 2009) Fissidens asplenioides Hedw. (Crosby et al., 1983; Bruggeman-Nannenga & Arts, 2010) Fissidens bogosicus Müll.Hal. in Venturi (Crosby et al., 1983; BruggemanNannenga & Arts, 2010) Fissidens borgenii Hampe (Bruggeman-Nannenga, 1993) Fissidens crispulus Brid. (Crosby et al., 1983; Bruggeman-Nannenga & Arts, 2010) *Fissidens cryptarum Müll.Hal. – Tampolo Tamatave, littoral forest base of tree, earthen insect nest, ant or termit, S 17˚16’ E 49˚24’, 26/10/1972, M.R. & C.A Crosby 6700, 6702, 6697, TAN. Fissidens cryptoneuron P. de la Varde (Crosby et al., 1983) Checklist of the bryophytes of Madagascar 231 Fissidens curvatus Hornsch. var. curvatus (Crosby et al., 1983; BruggemanNannenga & Arts, 2010) *Fissidens cyathaeicola Brugg.Nann. – Marojejy National Park, On tree fern 220 m up trail from camp Mantella, alt. 464 m, 24/11/2009, Wilding 4, det. Bruggeman-Nannenga; Marojejy National Park, on tree fern 300 m up trail from camp Mantella, alt. 465 m, 24/11/09, Wilding 6, det. BruggemanNannenga. Fissidens darntyi Schimp. (Crosby et al., 1983) Fissidens ellipticus Besch. (Crosby et al., 1983) Fissidens flaccidus Mitt. (Bruggeman-Nannenga, 2009) Fissidens grainvillei Thér. (Crosby et al., 1983) End, Type – Kiangara, sur écorces, en trés petite quantité, 1931, Grainville, holotype PC. Fissidens hookerioides Bizot et Onr. (Crosby et al., 1983) End, Type – Tananarive, domaine des Frères de Soavimbahoaka, sur latérite ombragée, alt. 1400 m, 06/07/1971, Onraedt 71M5137, holotype PC. Fissidens hymenodon Besch. (Crosby et al., 1983) Fissidens inclusus Bizot et Dury ex Pócs (Bruggeman-Nannenga, 2009) Fissidens intramarginatus (Hampe) A. Jaeger (Bizot, 1974; Crosby et al., 1983; Bruggeman-Nannenga & Arts, 2010) Fissidens leucocinctus Hampe (Crosby et al., 1983; Bruggeman-Nannenga & Arts, 2010) Fissidens madecassus Müll.Hal. (Crosby et al., 1983; Bruggeman-Nannenga, 2009 ; Bruggeman-Nannenga & Arts, 2010) Fissidens mariei (Besch.) Broth. (Crosby et al., 1983) End, Type – Nosy Be, forêt de Lokobe, 04/1879, Marie, holotype PC. Fissidens motelayi Renauld et Cardot (Crosby et al., 1983) End, Type – Andriba, 1897, Dorr, holotype PC. Fissidens ovatus Brid. (Crosby et al., 1983; Bruggeman-Nannenga & Arts, 2010) Fissidens pellucidus Hornsch. var. pellucidus (Crosby et al., 1983; Delgadillo, 1995; Bruggeman-Nannenga, 2010) Fissidens planifrons Besch. (Crosby et al., 1983; Bruggeman-Nannenga & Arts, 2010) Fissidens platyneuros Renauld et Cardot (Crosby et al., 1983) End, Type – Andevoranto, 1895, Perrot, holotype PC. Fissidens plumosus Hornsch. (Bruggeman-Nannenga & Arts, 2010) Fissidens pocsii Bizot et Dury ex Pócs (Bruggeman-Nannenga, 2009; BruggemanNannenga & Arts, 2010) Fissidens punctulatus Sande Lac. (Bruggeman-Nannenga & Arts, 2010) Fissidens ramulosus Mitt. (Crosby et al., 1983; Bruggeman-Nannenga & Arts, 2010) Fissidens reflexus Hampe (Crosby et al., 1983; Bruggeman-Nannenga, 2009) Fissidens rosulatus P. de la Varde (Crosby et al., 1983) End, Type – Tananarive, sur talus, 20/01/1907, Rotereau, holotype PC. Fissidens rufescens Hornsch. (Bruggeman-Nannenga, 2009) Fissidens sciophyllus f. sciophyllus Mitt. (Crosby et al. 1983; BruggemanNannenga, 1993; Bruggeman-Nannenga & Arts, 2010) Fissidens scleromitrius (Besch.) Broth. (Bescherelle, 1891; Crosby et al., 1983) Fissidens serratus Müll.Hal. var. serratus (Crosby et al., 1983; Delgadillo, 1995; Bruggeman-Nannenga, 2010) 232 L. Marline et al. Fissidens submarginatus Bruch ex C. Krauss (Crosby et al., 1983; BruggemanNannenga & Arts, 2010) Fissidens subplanifrons Bizot et Onraedt ex Brugg.-Nann (Bruggeman-Nannenga, 2009) Fissidens subundatus Dury (Crosby et al., 1983) Fissidens zollingeri Mont. (Crosby et al., 1983; Bruggeman-Nannenga & Arts, 2010) Funariaceae Entosthodon borbonicus Besch. (Crosby et al., 1983) Entosthodon marginatulus Müll. Hal. ex Geh. (Crosby et al., 1983) End, Type – Zone moyenne des forêts, forêt à l’Est d’Ambatondrazaka, Rutenberg, herb (N.S.). *Entosthodon mauritianus Schimp. ex Besch. – Befotaka forêt, S 18˚56’47” E 47˚06’03”, 16/08/1926, Decary 77, holotype MO. Entosthodon renauldii Thér. (Crosby et al., 1983) End, Type – Zone du Plateau central, pays Betsileo, 2/1905, Villaume, holotype PC, isotype PC, BM. Funaria decaryi Thér. (Crosby et al., 1983) End, Type – Ambovombe, FortDauphin, Decary, holotype S. Funaria delicatula Thér. (Crosby et al., 1983) End, Type – Environ de Tananarive, Perrier de la Bathie, holotype PC. Funaria hygrometrica var. calvescens (Schwaegr.) Kindb. (Crosby et al., 1983; Pócs & Geissler, 2002) Funaria imerinensis Cardot (Crosby et al., 1983) End, Type – Zone du Plateau central, Imerina, dans un gazon de Webera decurrens Ren. & Card, Villaume, holotype PC. Funaria subleptopoda Hampe (Crosby et al., 1983) End, Type – Imerina, 2/9/1847, Borgen, holotype PC. Physcomitrium coarctatum Cardot (Crosby et al., 1983) Physcomitrium dilatatum Renauld et Cardot (Crosby et al., 1983) End, Type – Environ de Tananarive, in terra, Camboué, holotype PC. Gigaspermaceae Gigaspermum repens (Hook.) Lindb. (Tixier, 1979b; Crosby et al., 1983) *Oedipodiella australis (Wager & Dixon) Dixon – Morondava, forêt, alt. 20 m, 11/04/1979, Tixier 12114, TAN. Grimmiaceae Grimmia elatior Bruch ex Bals.-Criv. et De Not. (Crosby et al., 1983) Grimmia elongata Kaulf. (Crosby et al., 1983) Grimmia longirostris Hook. (Muñoz, 1998) Bucklandiella membranaceum (Mitt.) Paris (Crosby et al., 1983) Hedwigiaceae *Braunia secunda (Hook.) Bruch et Schimp. – Station forestière de Manjakatompo, rocher, alt. 2600 m, 22/08/1971, Cremers 1705, TAN; Ankaratra, sur rocher, 27/03/1972, Cremers 1957, TAN; Ankaratra, sur rocher, basaltique, 08/1955, Bosser 8208, TAN. Bryowijkia madagassa Touw (Touw, 1993) Hedwigia ciliata (Hedw.) Ehrh. ex P. Beauv. var. ciliata (Crosby et al., 1983) Checklist of the bryophytes of Madagascar 233 Hypnaceae Ectropothecium chenagonii Renauld et Cardot (Crosby et al., 1983) Ectropothecium chenagonii var. hamatulum Renauld & Cardot (Crosby et al., 1983) End, Type – Entre Vinanitelo et Ikongo, 1892, Besson, holotype PC. Ectropothecium intertextum Renauld et Cardot (Crosby et al., 1983) Ectropothecium mauritianum Renauld et Cardot (Crosby et al., 1983) Ectropothecium occultum Renauld et Cardot (Crosby et al., 1983) Ectropothecium ovalifolium (Besch.) W.R. Buck (Crosby et al., 1983) End, Type – Zone inférieure des forêts, Nosy Be, forêt de Lokobe, Marie, herb (N.S.). Ectropothecium paillotii Renauld et Cardot (Crosby et al., 1983) Ectropothecium perrieri Thér. (Crosby et al., 1983) End, Type – Massif de l’Andringitra, alt. 1600 m, 1921, Perrier de la Bathie, holotype PC. Ectropothecium perrotii Renauld et Cardot (Crosby et al., 1983) Ectropothecium regulare (Brid.) A. Jaeger (Crosby et al., 1983) Ectropothecium seychellarum Besch. (Crosby et al., 1983) Ectropothecium tamatavense Broth. (Crosby et al., 1983) Gammiella ceylonensis (Broth.) B.C. Tan et W.R. Buck (Crosby et al., 1983) Glossadelphus guineensis (Broth. et Paris) Crosby, B.H. Allen et Magill (Crosby et al., 1983) Glossadelphus semiscabrus (Renauld et Cardot) Crosby et al. (Crosby et al., 1983) End, Type – Zone inférieure des forêts, environs d’Andevoranto, 1895, Perrot, holotype PC. Glossadelphus viridis (Renauld et Cardot) Crosby et al. (Crosby et al., 1983) End, Type – Diégo-Suarez, Montagne d’Ambre, Chénagon, holotype PC. Hypnum jutlandicum Holmen et E. Warncke (Crosby et al., 1983) Hypnum macrogynum Besch. (Crosby et al., 1983) Mittenothamnium horridulum (Broth.) Cardot (Crosby et al., 1983) Mittenothamnium lacouturei (Renauld et Cardot) Cardot (Crosby et al., 1983) End, Type – Betsileo, province d’Ambositra, 1906, Villaume, holotype BM# Mittenothamnium limosum (Besch.) Cardot (Crosby et al., 1983) Mittenothamnium madagassum (Besch.) Cardot (Crosby et al., 1983; Een, 1997) Mittenothamnium microthamnioides (Geh.) Wijk et Margad. (Crosby et al., 1983; Een, 1997) Mittenothamnium microthamnioides var. glauculum (Cardot) Wijk et Margad. (Crosby et al., 1983) End, Type – Forêt de Manongarivo, Drouhard, holotype PC# Mittenothamnium reptans (Hedw.) Cardot (Crosby et al., 1983) Phyllodon scutellifolius (Besch.) W.R. Buck (Crosby et al., 1983; O’Shea & Buck, 2001) End, Type – Nosy Komba, associé à Hookeria nossiana, 08/1879, Marie s.n., holotype BM. Phyllodon truncatulus (Welw. et Duby) W.R. Buck (Crosby et al., 1983) Rhacopilopsis trinitensis (Müll.Hal.) E. Britton ex Dixon (Crosby et al., 1983) Rhacopilopsis variegata (Welw. et Duby) M.C. Watling et O’Shea (Buck, 1993; Watling & O’Shea, 2000) Taxiphyllum taxirameum (Mitt.) M. Fleisch. (Crosby et al., 1983) 234 L. Marline et al. Vesicularia alboviridis var. rufula (Renauld et Cardot) Cardot (Crosby et al., 1983) End, Type – Zone inférieure des forêts, Diégo-Suarez, Chénagon, holotype PC. Vesicularia ayresii (Besch.) Broth. (Crosby et al., 1983) Vesicularia combae (Renauld et Cardot) Broth. (Crosby et al., 1983) End, Type – Zone inférieure des forêts, Nosy Komba, Marie, holotype PC# Vesicularia crassiramea (Renauld et Cardot) Broth. (Crosby et al., 1983) Vesicularia crassiramea var. laxifolia (Renauld et Cardot) Cardot (Crosby et al., 1983) End, Type – Zone des savanes occidentales, cercle de Maintirano, holotype PC. Vesicularia galerulata (Duby) Broth. (Crosby et al., 1983) Vesicularia galerulata var. subpiligera (Renauld et Paris) Wijk et Margad. (Crosby et al., 1983) End, Type – Zone du Plateau central, Betafo, rive droite de la Mania, 1901, Galinon, holotype PC. Vesicularia hygrobium (Besch.) Broth. (Crosby et al., 1983) Vesicularia immunda (Renauld) Cardot (Crosby et al., 1983) End, Type – Zone du Plateau central, Imerina oriental, Hildebrandt, herb (N.S.). Hypopterygiaceae Hypopterygium tamarisci (Sw. ex Sw.) Brid. ex Müll.Hal. (Crosby et al., 1983; Kruijer 1997; Kruijer, 2002) Lopidium struthiopteris (Brid.) M. Fleisch. (Crosby et al., 1983; Kruijer, 1997; Kruijer, 2002) Leptodontaceae Leptodon fuciformis (Brid.) Enroth (Crosby et al., 1983) Leskeaceae Pseudoleskea obtusiuscula Renauld et Cardot (Crosby et al., 1983) End, Type – Zone du Plateau central, Fianarantsoa, troncs d’arbres, 1904, Villaume, holotype BM!, isotype PC. Leucobryaceae Atractylocarpus madagascariensis (Thér.) Padberg et J.-P. Frahm (Crosby et al., 1983) Campylopus arctocarpus (Besch.) J.-P. Frahm subsp. madecassus (Crosby et al., 1983, Frahm, 1985) Campylopus arcuatus (Brid.) A. Jaeger (Crosby et al., 1983; Frahm, 1985) Campylopus aureonitens (Müll.Hal.) A. Jaeger (Crosby et al., 1983; Frahm, 1985) Campylopus cambouei Renauld et Cardot (Crosby et al., 1983; Frahm, 1985) Campylopus catarractilis (Müll.Hal.) Paris (Frahm, 1985) Campylopus crateris Besch. (Crosby et al., 1983; Frahm, 1985) Campylopus decaryi Thér. (Crosby et al., 1983; Frahm, 1985) Campylopus filifolius (Hornsch.) Mitt. (Crosby et al., 1983; Een, 1993) Campylopus flaccidus Renauld et Cardot (Crosby et al., 1983; Frahm, 1985) Campylopus flexuosus (Hedw.) Brid. var. flexuosus (Crosby et al., 1983; Frahm, 1985) Campylopus hildebrandtii (Müll.Hal.) A. Jaeger (Crosby et al., 1983) Campylopus jamesonii (Hook.) A. Jaeger (Crosby et al., 1983; Frahm, 1985) Campylopus julaceus (Renauld et Cardot) J.-P. Frahm subsp. arbogastii (Crosby et al., 1983; Frahm, 1985) Checklist of the bryophytes of Madagascar 235 Campylopus nanophyllus Müll.Hal. ex Broth. (Crosby et al., 1983) Campylopus nivalis (Brid.) Brid. var. nivalis (Frahm, 1985) Campylopus perpusillus Mitt. (Crosby et al., 1983; Frahm, 1985) Campylopus pilifer Brid. subsp. pilifer (Crosby et al., 1983; Frahm, 1985) Campylopus pseudobicolor Müll.Hal. ex Renauld & Cardot (Crosby et al., 1983; Frahm, 1985) Campylopus robillardei Besch. (Crosby et al., 1983; Frahm, 1985; Pócs & Geissler, 2002) Campylopus schmidii (Müll.Hal.) A. Jaeger (Crosby et al., 1983; Frahm, 1985) Campylopus smaragdinus (Brid.) A. Jaeger (Crosby et al., 1983; Frahm, 1985) Campylopus thwaitesii (Mitt.) A. Jaeger (Crosby et al., 1983; Frahm, 1985) Campylopus trachyblepharon (Renauld et Cardot) J.-P. Frahm subsp. comatus (Crosby et al., 1983; Frahm, 1985) Leucobryum acutifolium (Mitt.) Cardot (Crosby et al., 1983) Leucobryum boivinianum Besch. (Crosby et al., 1983) Leucobryum boryanum Besch. (Crosby et al., 1983) Leucobryum boryanum var. minus Cardot (Crosby et al., 1983) End, Type – Pays Betsileo, sineloco, Montaul, herb (N.S.). Leucobryum cambouei Cardot (Crosby et al., 1983) End, Type – Camboué, herb (N.S.). Leucobryum comorense Müll.Hal. (Crosby et al., 1983; Pócs & Geissler, 2002) Leucobryum cucullatum Broth. (Crosby et al., 1983) Leucobryum cucullatum var. rutenbergii (Geh.) Cardot (Crosby et al., 1983) Leucobryum galinonii Cardot et Paris (Crosby et al., 1983) End, Type – Betafo, rive droite de la Mania, Galinon, holotype PC# Leucobryum heterodictyon Besch. (Bescherelle, 1891; Crosby et al., 1983) End, Type –Sainte Marie, Marie Edouard Auguste, holotype PC. Leucobryum heterodictyon var. microphyllum (Cardot) Wijk et Margad. (Crosby et al., 1983) End, Type – Ile Sainte Marie, Marie, holotype PC# Leucobryum isleanum Besch. (Crosby et al., 1983) Leucobryum isleanum var. molle (Müll.Hal.) Cardot (Crosby et al., 1983) Leucobryum madagassum Besch. (Crosby et al., 1983) Leucobryum mayottense Cardot (Crosby et al., 1983) Leucobryum parvulum Cardot (Crosby et al., 1983, Pócs & Geissler, 2002) End, Type – holotype PC. Leucobryum perrotii Renauld et Cardot (Crosby et al., 1983, Pócs & Geissler, 2002) Leucobryum pseudomadagassum Cardot (Crosby et al., 1983) End, Type – Zone supérieure des forêts, Imerina, Andrangoloaka, Hildebrandt, holotype PC. Leucobryum sanctae-mariae Cardot (Crosby et al., 1983, Pócs & Geissler, 2002) End, Type – Ile Sainte Marie, Marie, holotype PC. Ochrobryum sakalavum Cardot et Paris (Crosby et al., 1983; Allen, 1992; Pócs & Geissler, 2002) End, Type – cercle de Maintirano, poste d’Ampoza, près du village de Tahotofoly, sur bois pourri, holotype PC! Leucodontaceae Leucodon canariensis (Brid.) Schwaegr. (Crosby et al., 1983) Leucodon rutenbergii Müll.Hal. ex Geh. (Crosby et al., 1983) 236 L. Marline et al. Pterogoniadelphus assimilis (Müll.Hal.) Ochyra et Zijlstra (Crosby et al., 1983) Pterogonium gracile (Hedw.) Sm. (Crosby et al., 1983) Pterogonium gracile var. madagassum (Geh.) M. Fleisch. (Crosby et al., 1983) Leucomiaceae Leucomium strumosum (Hornsch.) Mitt. (Crosby et al., 1983; Allen, 1987; Pócs & Geissler, 2002) Meteoriaceae Aerobryopsis capensis (Müll.Hal.) M. Fleisch. (Crosby et al., 1983) Barbella capillicaulis (Renauld et Cardot) Cardot var. capillicaulis (Crosby et al., 1983) Floribundaria floribunda (Dozy et Molk.) M. Fleisch. (Crosby et al., 1983) Floribundaria floribunda var. madagassa (Renauld et Cardot) Renauld et Cardot (Crosby et al., 1983) End, Type – Forêt d’Ambohimitombo, Forsyth Major, FH; Fianarantsoa, plateau d’Ikongo, entre Savondronina et Ranomafana, et entre Vinanitelo et Ikongo, Besson, holotype (N.S.). Floribundaria floribundula (Müll.Hal.) M. Fleisch. (Crosby et al., 1983) Floribundaria vaginans (Welw. et Duby) Broth. (Crosby et al., 1983) Papillaria africana (Müll.Hal.) A. Jaeger (Crosby et al., 1983) Papillaria africana var. gracilis (Renauld et Cardot) Wijk et Margad. (Crosby et al., 1983) End, Type – Diégo-Suarez, 1890, Chenagon; Entre Vinanintelo et Ikongo, 1892, Besson. Betsileo, sine loco, Rev. Montaut; Entre Savondronina et Ranomafana, 1892, Besson, holotype (N.S.). Papillaria borchgrevinkii Kiaer (Crosby et al., 1983) Papillaria flaccidula Cardot (Crosby et al., 1983) End, Type – Zone moyenne des forêts, forêt d’Analamazaotra, Camboué, herb (N.S.). Papillaria perichaetialis (Hampe) A. Jaeger (Crosby et al., 1983) End, Type – Analamazaotra, 1871, Borchgrewink 17, holotype BM. Trachypodopsis serrulata (P. Beauv.) M. Fleisch. var. serrulata (Van Zanten, 1959; Crosby et al., 1983; Pócs & Geissler, 2002) Trachypus appendiculatus (Renauld et Cardot) Broth. (Van Zanten, 1959; Crosby et al., 1983) Trachypus bicolor var. viridulus (Mitt.) Zanten (Van Zanten, 1959; Crosby et al., 1983) Meesiaceae Leptobryum pyriforme (Hedw.) Wilson (Crosby et al., 1983) Mniaceae Epipterygium diversifolium Renauld et Cardot (Crosby et al., 1983) End, Type – Zone des savanes occidentales, cercle de Maintirano, 1901, Collecteur indigène, holotype PC. Mielichhoferia cardotii A.J. Shaw et H.A. Crum (Crosby et al., 1983) End, Type – Zone de plateau central, Ambohipo, Camboué, herb (N.S.). Mielichhoferia madagassa var. gracilis Thér. (Crosby et al., 1983) End, Type – Mont Angavokely, alt. 1800 m, 1924, Drouhard, holotype PC. Plagiomnium rhynchophorum (Hook.) T.J. Kop. var. rhynchophorum (Koponen, 1981; Crosby et al., 1983) Pohlia baronii Wijk et Margad. (Crosby et al., 1983) Pohlia elongata Hedw. var. elongata (Crosby et al., 1983; Delgadillo, 1995) Checklist of the bryophytes of Madagascar 237 Pohlia heribaudii var. madagassa (Renauld et Cardot) Wijk et Margad. (Crosby et al., 1983) Pohlia lacouturei (Thér.) O’Shea (Crosby et al., 1983) End, Type – Pays des Betsileo, sur gneiss en décomposition, Villaume & Lacouture, herb (N.S.). Pseudopohlia microstoma (Harv.) Mizush. (Crosby et al., 1983) Schizymenium commixtum (Thér.) A.J. Shaw (Crosby et al., 1983) End, Type – Mont Tsaratanana, forêt à sous-bois herbacé, alt. 1200-2400 m, Perrier de la Bathie, holotype PC. Schizymenium madagassum (Renauld et Cardot ex Cardot) A.J. Shaw (Crosby et al., 1983) End, Type – Mont Angavokely, Est Antananarivo, alt. 1800 m, 1924, Drouhard, holotype PC. Neckeraceae Homaliodendron exiguum (Bosch et Sande Lac.) M. Fleisch. (Crosby et al., 1983) Homaliodendron piniforme (Brid.) Enroth (Crosby et al., 1983; Enroth, 1993) Neckera remota Bruch et Schimp. ex Müll.Hal. (Crosby et al., 1983) Neckera valentiniana Besch. (Crosby et al., 1983) Neckeropsis boiviniana (Besch.) Cardot (Crosby et al., 1983) *Neckeropsis disticha (Hedw.) Kindb. – Brickaville, large trees next to shallow rive, S 18˚55’ E 48˚34’, alt. 450 m, 23/10/1972, M.R. & C.A Crosby 6754, TAN. Neckeropsis lepineana (Mont.) M. Fleisch. (Crosby et al., 1983) Neckeropsis madecassa (Besch.) M. Fleisch. (Crosby et al., 1983) Orthostichidium pentastichum (Brid.) B.H. Allen et Magill. (Crosby et al., 1983; Allen & Magill, 2007) Orthostichella rigida (Müll.Hal.) B.H. Allen et Magill. (Allen & Magill, 2007) Orthostichella versicolor (Müll.Hal.) B.H. Allen et Magill. (Crosby et al., 1983; Allen & Magill, 2007) Orthostichella longinervis (Renauld et Cardot) B.H. Allen et Magill. (Allen & Magill, 2007) Orthostichella welwitschii (Duby) B.H. Allen et Magill. (Allen & Magill, 2007) Pilotrichella flexilis (Hedw.) Aongstr. (Crosby et al., 1983; Allen & Magill, 2003) Pilotrichella flexilis f. nudiramulosa (Müll.Hal.) B.H. Allen et Magill (Allen & Magill, 2003) Pilotrichella mascarenica (Müll.Hal.) A. Jaeger (Crosby et al., 1983; Allen & Magill, 2003) Pilotrichella pseudo-imbricata (Müll.Hal.) A. Jaeger (Crosby et al., 1983) Pinnatella minuta (Mitt.) Broth. (Crosby et al., 1983; Enroth, 1994) Porotrichum elongatum (Welw. et Duby) A. Gepp (De Sloover, 1983) Porotrichum madagassum Kiaer ex Besch. (De Sloover, 1983) Porotrichum stipitatum (Mitt.) W.R. Buck (De Sloover, 1983) Porotrichum usagarum Mitt. (De Sloover, 1983) Porotrichum variifolioides (De Sloover) Enroth (De Sloover, 1983) Thamnobryum malgachum (Cardot) O’Shea (Crosby et al., 1983) End, Type – Zone moyenne des forêts, forêt de Manongarivo, 1909, Drouhard, holotype PC. Orthotrichaceae Cardotiella appendiculata (Renauld et Cardot) Vitt (Crosby et al., 1983; Goffinet, 1996) 238 L. Marline et al. Cardotiella elimbata (Thér.) Goffinet (Crosby et al., 1983; Goffinet, 1996) End, Type – Farafangana, sur souche d’arbre, en forêt, Decary, holotype PC, isotype S. Cardotiella renauldii (Broth.) Vitt. (Crosby et al., 1983; Goffinet, 1996) End, Type – holotype PC. Cardotiella subappendiculata (Broth.) Vitt (Crosby et al., 1983) Groutiella laxotorquata (Besch.) Wijk et Margad. (Crosby et al., 1983) Macrocoma tenuis (Hook. et Grev.) Vitt subsp. tenuis (Crosby et al., 1983) Macromitrium adelphinum Cardot (Crosby et al., 1983) End, Type – Forêt d’Amperifery, Campenon, herb (N.S.). Macromitrium ancistrophyllum Cardot (Crosby et al., 1983) End, Type – Zone supérieure des forêts, Ankeramadinika, Borgen 202, holotype BM. Macromitrium anomodictyon Cardot (Crosby et al., 1983) End, Type – District d’Andramasina, entre Tsinjoarivo & Ambohimasina, dans un petit bois, sur branche d’arbre, alt. 1700 m, 02/12/1912, Viguier 1932, holotype PC. Macromitrium calocalyx Geh. (Crosby et al., 1983) End, Type – Zone moyenne des forêts, forêt à l’est d’Ambatondrazaka, Rutenberg, herb (N.S.). Macromitrium fasciculare Mitt. (Crosby et al., 1983) Macromitrium fasciculare var. angustifolium P. de la Varde (Crosby et al., 1983) End, Type – Sommet oriental du Massif de Marojejy, à l’Ouest de la Haute Mantenina, affluent de la Lokoho, gneiss et quartzite, alt. 1850 m, 26/3/1949, Humbert, holotype PC. Macromitrium fimbriatum (P.Beauv.) Schwaegr, (Crosby et al., 1983) Macromitrium mauritianum Schwaegr. (Crosby et al., 1983) Macromitrium nanothecium Cardot (Crosby et al., 1983) End, Type – Zone supérieure des forêts, Andrangoloaka, Hildebrandt, holotype FH. Macromitrium nanothecium var. sublaeve Thér. (Crosby et al., 1983) End, Type – Fort-Dauphin, sur ecorce, Decary, holotype PC! Macromitrium onraedtii Bizot (Bizot, 1974; Crosby et al., 1983) End, Type – Entre Foulpointe et Andondabe, au Nord de Tamatave, sur un arbre d’une forêt sur sable, 12/05/1972, Cremers 2165, holotype PC. Macromitrium orthostichum Nees ex Schwägr. (Crosby et al., 1983) Macromitrium proximum Thér. (Crosby et al., 1983) End, Type – Mont Tsaratanana, alt. 1200 m, 04/1924, Perrier de la Bathie, holotype PC. Macromitrium rhizomatosum Müll.Hal. (Crosby et al., 1983) End, Type – Nosy Be, Pervillé 789, holotype PC. Macromitrium sanctae-mariae Renauld et Cardot (Crosby et al., 1983) End, Type – Sainte Marie, 1893, Darbould, holotype PC. Macromitrium sclerodictyon Cardot (Crosby et al., 1983) Macromitrium semipapillosum Thér. et P. de la Varde (Crosby et al., 1983) End, Type – 1922, Carrougeau, holotype PC. Macromitrium serpens (Bruch ex Hook. & Grev.) Brid. (Crosby et al., 1983) Macromitrium soulae Renauld et Cardot (Crosby et al., 1983) End, Type – Ambositra, Betsileo, 1890, Soula, holotype PC. Macromitrium subpungens Hampe ex Müll.Hal. (Crosby et al., 1983) Macromitrium subpungens var. madagassum Cardot (Crosby et al., 1983) End, Type – Zone inférieure des forêts, Maroantsetra dans la baie d’Antongil, Mathieu, herb (N.S.). Checklist of the bryophytes of Madagascar 239 Macromitrium subtortum (Hook. et Grev.) Schwaegr. (Crosby et al., 1983) End, Type – unknown. Macromitrium sulcatum (Hook.) Brid. var. sulcatum (Crosby et al., 1983; Ellis & Wilbraham, 2008) Macromitrium urceolatulum Müll.Hal. ex Geh. (Crosby et al., 1983) Schlotheimia acutifolia Renauld et Paris (Crosby et al., 1983) End, Type – Forêt d’Ambohitantely, 1901, Collecteur indigène, holotype PC. Schlotheimia acutifolia var. angustifolia Thér. (Crosby et al., 1983) End, Type – Mont Tsaratanana, forêt à sous bois herbacé, alt. 1200-2400 m, Perrier de la Bathie, holotype PC. Schlotheimia badiella Besch. (Crosby et al., 1983) Schlotheimia badiella var. helicophylla Besch. (Crosby et al., 1983) Schlotheimia boiviniana Besch. (Crosby et al., 1983) Schlotheimia conica Renauld et Cardot (Crosby et al., 1983) End, Type – Environs de la forêt d’Ambatovory, alt. 1400 m, 1889, Camboué, holotype PC. Schlotheimia excorrugata Müll.Hal. ex Cardot (Crosby et al., 1983) Schlotheimia fornicata Duby (Crosby et al., 1983) Schlotheimia fornicata var. apiculata P. de la Varde (Crosby et al., 1983) End, Type – Pentes orientale du massif de Marojejy, à l’ouest de la rivière Manantenina, affluent de la Lokoho, sylve à lichens sur gneiss et quartzite, alt. 1400 m, 15/12/1948, Humbert, holotype PC. Schlotheimia fornicata var. microtheca P. de la Varde (Crosby et al., 1983) End, Type – Vallée de la Lokoho Nord-Est, mont Beondroka, au nord de Maroambihy, rochers de la crête (gneiss et quartzite), alt. 1400 m, 17/03/1949, Humbert, holotype PC. Schlotheimia foveolata Renauld et Cardot (Crosby et al., 1983) End, Type – Zone inférieure des forêts, Ile de Sainte Marie sur les bords de la rivière Fitarihana, 1893, Darbould, holotype PC. Schlotheimia foveolata var. torquescens Cardot (Crosby et al., 1983) End, Type – Zone du Plateau central, Imerina, environ de Tananarive, Camboué & Sibree, holotype BM. Schlotheimia gauthieri Thér. (Crosby et al., 1983) Schlotheimia malacophylla Besch. (Crosby et al., 1983) Schlotheimia microcarpa Geh. (Crosby et al., 1983) Schlotheimia microphylla Besch. (Crosby et al., 1983) Schlotheimia nossi-beana Müll.Hal. (Crosby et al., 1983) End, Type – Nosy Be, 1841, Pervillé, holotype PC. Schlotheimia nossi-beana var. linealis (Geh.) Thér. (Crosby et al., 1983) End, Type – Nosy Be, Boivin, holotype PC. Schlotheimia pauli Renauld et Cardot (Crosby et al., 1983) End, Type – Antsahambavy, 1902, Dalincourt, holotype PC. Schlotheimia perrieri Thér. (Crosby et al., 1983) End, Type – Mont Tsaratanana, alt. 1200m, 4/1924, Perrier de la Bathie, holotype PC. Schlotheimia perrotii Renauld et Cardot (Crosby et al., 1983) End, Type – Andevoranto, 1895, Perrot, holotype PC. Schlotheimia perrotii var. breviseta Thér. (Crosby et al., 1983) End, Type – Andevoranto, herb (N.S.). Schlotheimia robillardii Duby (Crosby et al., 1983) Schlotheimia robusta Thér. (Crosby et al., 1983) 240 L. Marline et al. Schlotheimia semidiaphana (Renauld et Cardot) Cardot (Crosby et al., 1983) End, Type – Plateau d’Ikongo, 1892, Besson 294, holotype PC. Schlotheimia squarrosa Brid. (Crosby et al., 1983; Delgadillo, 1995) Schlotheimia subfornicata Thér. (Crosby et al., 1983) Schlotheimia tenuiseta Geh. (Crosby et al., 1983) End, Type – Ambatondrazaka, herb (N.S.). Schlotheimia trichophora Renauld et Cardot (Crosby et al., 1983) End, Type – Entre Ankeramadinika et la forêt d’Analamazaotra, 1891, Camboué & Campenon, holotype PC. Schlotheimia trichophora var. stenophylla (Renauld et Cardot) Thér. (Crosby et al., 1983) End, Type – Zone du Plateau central, pays de Betsileo, Villaume, holotype BM. Schlotheimia trypanoclada Geh. (Crosby et al., 1983) End, Type – Zone du Plateau central, Alabe, près de Marotandrano, Rutenberg, holotype PC. Ulota fulva Brid. (Crosby et al., 1983) Zygodon laxifolius Thér. (Crosby et al., 1983) End, Type – Massif d’Andringitra, alt. 1600 m, 1921, Perrier de la Bathie, holotype PC. Zygodon leptocarpus Thér. (Crosby et al., 1983) End, Type – Massif d’Andringitra, alt. 1600 m, 1921, Perrier de la Bathie, holotype PC. Zygodon reinwardtii var. madacassus Cardot (Crosby et al., 1983) End, Type – NW Madagascar, Pervillé, holotype PC. Zygodon reinwardtii (Hornsch.) A. Braun var. reinwardtii (Crosby et al., 1983) Phyllogoniaceae Phyllogonium viscosum (P. Beauv.) Mitt. (Crosby et al., 1983) Pilotrichaceae Callicostella africana Mitt. (Crosby et al., 1983) Callicostella brevipes (Broth.) Broth. (Crosby et al., 1983) Callicostella constricta (Müll.Hal.) Kindb. (Crosby et al., 1983) Callicostella fissidentella (Besch.) Kindb. (Crosby et al., 1983) Callicostella lacerans (Müll.Hal.) A. Jaeger (Crosby et al., 1983) Callicostella leptocladula (Broth.) Broth. (Crosby et al., 1983) Callicostella papillata var. brevifolia M. Fleisch. (Crosby et al., 1983) Callicostella papillata (Mont.) Mitt. var. papillata (Crosby et al., 1983; Pócs & Geissler, 2002) Callicostella parvocellulata Mitt. (Crosby et al., 1983) Callicostella perrotii (Renauld et Cardot ex Paris) Broth. (Crosby et al., 1983) Callicostella pusilla Demaret et P. de la Varde (Crosby et al., 1983) Callicostella seychellensis (Besch.) Kindb. (Crosby et al., 1983; Pócs & Geissler, 2002) Callicostella tristis (Müll.Hal.) Broth. (Pócs & Geissler, 2002) Cyclodictyon aubertii (P. Beauv.) Kuntze (Crosby et al., 1983) Cyclodictyon laetevirens (Hook. et Taylor) Mitt. (Crosby et al., 1983) Cyclodictyon vallis-gratiae (Müll.Hal.) Kuntze (Crosby et al., 1983; Pócs & Geissler, 2002) Hookeriopsis diversifolia (Renauld et Cardot) Broth. ex Cardot in Grandid. (Crosby et al., 1983) Lepidopilidium attenuatum Thér. (Crosby et al., 1983) End, Type – Forêt d’Andasibe, sur l’Onive, 02/1925, Perrier de la Bathie, holotype PC. Checklist of the bryophytes of Madagascar 241 Lepidopilidium chenagonii (Renauld et Cardot) Cardot (Crosby et al., 1983) End, Type – Zone inférieure des forêts, montagne d’Ambre, près DiégoSuarez, Chénagon, herb (N.S.). Lepidopilidium corbieri (Renauld et Cardot) Cardot (Crosby et al., 1983) End, Type – Zone inférieure des forêts, montagne d’Ambre, Leloutre, holotype CHE. Lepidopilidium parvulum Cardot (Crosby et al., 1983) End, Type – NW, Pervillé, herb (N.S.). Lepidopilidium subrevolutum (Renauld et Cardot) Cardot (Crosby et al., 1983) Lepidopilum hirsutum var. tuberculatum Thér. (Crosby et al., 1983) End, Type – Andasibe, sur l’Onive, forêt à sous-bois herbacé, alt. 800-1400 m, 02/1925, Perrier de la Bathie, holotype PC. Lepidopilum lastii Mitt. (Crosby et al., 1983; Churchill, 1988) Lepidopilum verrucipes Cardot (Crosby et al., 1983; Churchill, 1988) Polytrichaceae Atrichum androgynum (Müll.Hal.) A. Jaeger (Crosby et al., 1983; De Sloover, 1986) Pogonatum belangeri (Müll.Hal.) A. Jaeger (Hyvönen, 1989; Pócs & Geissler, 2002) Pogonatum capense (Hampe) A. Jaeger (Crosby et al., 1983; Hyvönen, 1989; De Sloover, 1986; Pócs & Geissler, 2002) Pogonatum convolutum (Hedw.) P. Beauv. (Crosby et al., 1983; Hyvönen, 1989; De Sloover, 1986; Pócs & Geissler, 2002) Pogonatum gracilifolium Besch. (Crosby et al., 1983) Polytrichastrum formosum (Hedw.) G.L. Sm. (Crosby et al., 1983) Polytrichum commune Hedw. var. commune (Crosby et al., 1983; De Sloover, 1986) Polytrichum glabrum Schrad. (Crosby et al., 1983) Polytrichum juniperellum Geh. (Crosby et al., 1983) End, Type – Plateau central, Ambohimaro, Ambatomainty près Tananarive, Rutenberg, herb (N.S.). Polytrichum subformosum var. parvisetum (Müll.Hal.) Cardot (Crosby et al., 1983) End, Type – Plateau central, Imerina, bords du lac d’Ambohipo, Camboué, PC. Analamainty, Campenon, PC, holotype (N.S.). Polytrichum subpilosum P. Beauv. (Crosby et al., 1983; De Sloover, 1986) Pottiaceae Anoectangium impressum Hampe (Crosby et al., 1983) Anoectangium madagassum Renauld et Paris (Crosby et al., 1983) End, Type – Pays des Betsileo, Villaume, herb (N.S.). Anoectangium rufoviride Besch. (Crosby et al., 1983) Barbula bolleana (Müll.Hal.) Broth. (Crosby et al., 1983) Barbula eubryum Müll.Hal. (O’Shea et al., 1997) Barbula indica (Hook.) Spreng. var. indica (Crosby et al., 1983) Barbula jananicus O’Shea (O’Shea, 2008) End, Type – Janjanina, carton de Mahaiza, herb (N.S.). Barbula omissa Thér. (Sim & Dixon, 1922) Barbula subobtusa Thér. (Crosby et al., 1983) End, Type – Vondrozo, forêt sur rocher humide au bord d’une cascade, 11/09/1924, Decary, holotype PC. Bryoerythrophyllum campylocarpum (Müll.Hal.) H. A. Crum (Sollman, 2009) 242 L. Marline et al. Chionoloma bombayense (Müll.Hal.) P. Sollm. (Crosby et al., 1983) Didymodon subrevolutus (Hampe) Broth. (Crosby et al., 1983) End, Type – 1869, Borgen, holotype PC. Gymnostomiella tuberculosa (Renauld et Paris) Arts et P. Sollm. (Crosby et al., 1983) Gymnostomum madagascariense O’Shea (O’Shea, 2008) End, Type – Desvaux, herb (N.S.). Hymenostylium subcrispulum Thér. (Crosby et al., 1983) End, Type – Mont Tsaratanana, forêt à sous bois herbacé, alt. 1200-2400 m, 04/1924, Perrier de la Bathie, holotype PC# Hyophila acuminata Broth. et P. de la Varde (Crosby et al., 1983) Hyophila angustifolia Paris et Renauld (Crosby et al., 1983) End, Type – Rive de la Mania, 08/1901, Galinon, holotype PC. Hyophila involuta (Hook.) A. Jaeger (Crosby et al., 1983) Hyophila lanceolata Renauld et Cardot (Crosby et al., 1983) Hyophila parietalis Cardot (Crosby et al., 1983) End, Type – Zone des savanes occidentales, Marovoay, Mathieu, herb (N.S.). Hyophila potieri var. atroviridis Renauld et Cardot (Crosby et al., 1983) End, Type – Plateau central, Ambohibeloma, Imerina, Camboué, holotype PC# Hyophila sakalavensis Paris et Renauld (Crosby et al., 1983) Leptodontium flexifolium (With.) Hampe (Sollman, 2006) Leptodontium pungens (Mitt.) Kindb. (De Sloover, 1987) Leptodontium viticulosoides (P. Beauv.) Wijk et Margad. var. viticulosoides (Zander, 1972; Crosby et al., 1983; De Sloover, 1987; Delgadillo, 1995) Splachnobryum gracile Besch. (Crosby et al., 1983) Splachnobryum obtusum (Brid.) Müll.Hal. (Crosby et al., 1983; Arts, 1996; Arts, 2001) Streptopogon calymperes Müll.Hal. ex Geh. (Crosby et al., 1983) Streptopogon erythrodontus (Taylor) Wilson (Crosby et al., 1983; Delgadillo, 1995) Tortella humilis (Hedw.) Jenn. (Crosby et al., 1983) Tortula madagassa Dixon (Crosby et al., 1983) End, Type – unknown. Tortula muralis var. madagascariensis Thér. (Crosby et al., 1983) End, Type – Andrahomana, sur rocher gréseux, Decary, herb (N.S.). Tortula omissa Thér. (Crosby et al., 1983) End, Type – Imerimandroso, sur écorce, associé à Bryum truncorum Brid. Decary, holotype PC. Tortula subrufa Cardot (Crosby et al., 1983) End, Type – Fianarantsoa, 1905, Villaume s.n., holotype PC, isotype PC. Trichostomum brachydontium Bruch var. brachydontium (Crosby et al., 1983) Trichostomum pennequinii Renauld et Paris (Crosby et al., 1983) End, Type – Cercle des Bara, secteur d’Ivondro, 1900, Lespagnol et Croll, holotype PC. Trichostomum perrieri Thér. (Crosby et al., 1983) End, Type – Massif granitique de l’Andringitra, alt. 1600 m, 1921, Perrier de la Bathie, holotype PC. Trichostomum sporaphyllum (Renauld et Cardot) Cardot (Crosby et al., 1983) End, Type – Ambohibeloma, 1906, Camboué, holotype PC. Trichostomum tenuirostre (Hook. et Taylor) Lindb. var. tenuirostre (Crosby et al., 1983) Checklist of the bryophytes of Madagascar 243 Trichostomum villaumei Thér. (Crosby et al., 1983) End, Type – Fianarantsoa, 1905, Villaume, holotype PC. Weisiopsis plicata (Mitt.) Broth. (Crosby et al., 1983) Weissia ayresii Schimp. (Crosby et al., 1983) Prionodontaceae Prionodon ciliatus Besch. (Crosby et al., 1983) Pterobryaceae Calyptothecium bescherellei (Renauld) Broth. (Crosby et al., 1983) End, Type – Mont Ankaratra, Borgen, holotype BM! Calyptothecium planifrons (Renauld et Paris) Argent (Argent, 1973; Crosby et al., 1983; Delgadillo, 1995) Hildebrandtiella endotrichelloides Müll.Hal. (Crosby et al., 1983) Hildebrandtiella longiseta Renauld et Cardot (Crosby et al., 1983) End, Type – Diégo-Suarez, 1890, Chénagon, holotype PC. Hildebrandtiella rotundifolia Renauld (Crosby et al., 1983) Jaegerina solitaria var. nossi-beana Besch. (Crosby et al., 1983) End, Type – Zone inférieure des forêts, Nosy Be, Pervillé, herb (N.S.). Jaegerina stolonifera (Müll.Hal.) Müll.Hal. subsp. stolonifera (Argent, 1973; Crosby et al., 1983) Orthostichidium involutifolium (Mitt.) Broth. subsp. involutifolium (Crosby et al., 1983) Orthostichopsis longinervis (Renauld et Cardot) Broth. (Crosby et al., 1983) Orthostichopsis madagassa Dixon (Crosby et al., 1983) End, Type – unknown. Orthostichopsis scaberula (Renauld et Cardot) M. Fleisch (Crosby et al., 1983) End, Type – Zone moyenne des forêts, entre Vinanitelo et Ikongo, sur les rameaux grêles des arbres, 1892, Besson, holotype PC# Orthostichopsis subimbricata (Hampe) Broth. (Argent, 1973; Crosby et al., 1983) Orthostichopsis subimbricata var. flageyi (Renauld et Cardot) Cardot (Crosby et al., 1983) End, Type – Zone inférieure des forêts, Diégo-Suarez, Chenagon; Zone supérieure, Andrangoloaka, Sikora, herb (N.S.). Pterobryopsis longissima Dixon (Crosby et al., 1983) End, Type – unknown. Renauldia hildebrandtielloides Müll.Hal. (Argent, 1973; Crosby et al., 1983) Renauldia patentissima (Hampe) Broth. (Argent, 1973; Crosby et al., 1983) End, Type –1892, Besson, holotype PC. Renauldia patentissima var. mutica Thér. (Crosby et al., 1983) End, Type – Montagne d’Ambre, alt. 1000 m, 9/1926, Perrier de la Bathie, holotype PC. Pterygynandraceae Trachyphyllum borgenii (Renauld et Cardot) Broth. (Buck, 1979; Crosby et al., 1983; O’Shea, 1999) End, Type – Mont Ankaratra, lectotype BM, isolectotypes H, NY Trachyphyllum inflexum (Harv.) A. Gepp (Buck, 1979; Crosby et al., 1983) Ptychomitriaceae Ptychomitrium crispatum (Hedw.) A. Jaeger (Crosby et al., 1983) Ptychomitrium deltorii (Thér.) Broth. (Crosby et al., 1983) 244 L. Marline et al. Pylaisiadelphaceae Isopterygium ambreanum Renauld et Cardot (Crosby et al., 1983) End, Type – Diégo-Suarez, sur les écorces, 1890, Chénagon, holotype PC. Isopterygium antsihanakense Cardot (Crosby et al., 1983) End, Type – Zone moyenne des forêts, pays d’Antsihanaka, Perrot, herb (N.S.). Isopterygium argillicola (Renauld et Cardot) Broth. (Crosby et al., 1983) End, Type – Entre Tananarive et Betafo, Caussèque, holotype PC. Isopterygium argyroleucum Besch. (Crosby et al., 1983) Isopterygium austrodenticulatum (Renauld et Cardot) Broth. (Crosby et al., 1983) End, Type – Andriba, 1897, Dorr, holotype PC. Isopterygium combae Besch. (Crosby et al., 1983) End, Type – Zone inférieure des forêts, Nosy Be et Nosy Komba, Marie, herb (N.S.). Isopterygium gracile Renauld et Cardot (Crosby et al., 1983) Isopterygium intortum (P. Beauv.) A. Jaeger (Crosby et al., 1983) Isopterygium intortum var. chenagonii Renauld et Cardot (Crosby et al., 1983) End, Type – Zone inférieure des forêts, Diégo-Suarez, Chénagon, herb (N.S.). Isopterygium leptoblastum (Müll.Hal.) A. Jaeger (Crosby et al., 1983) Isopterygium luteonitens (Paris) Paris (Crosby et al., 1983) End, Type – Zone du Plateau central, bords gramineux du lac d’Ambohipo, Camboué, herb (N.S.). Isopterygium meylanii Cardot (Crosby et al., 1983) End, Type – Zone du Plateau central, Imerina, Villaume, holotype G. Isopterygium podorrhizum Renauld et Cardot (Crosby et al., 1983) End, Type – Zone du Plateau central, Ambatomanga, dans un gazon d’Ectropothecium regulare, alt. 1200-1300 m, 1894, Talazac, herb (N.S.). Isopterygium subleptoblastum Müll.Hal. (Crosby et al., 1983) Taxithelium argyrophyllum Renauld et Cardot (Crosby et al., 1983) End, Type – Zone inférieure des forêts, sur les troncs pourrissants à Diégo-Suarez, Chénagon, holotype PC. Taxithelium confusum Cardot (Crosby et al., 1983) End, Type – Ile Sainte Marie, Boivin, herb (N.S.). Taxithelium glaucophyllum Besch. (Crosby et al., 1983) End, Type – Zone inférieure des forêts, Nosy Be, forêt de Lokobe, Marie, herb (N.S.). Taxithelium hirtellum Paris et Renauld (Crosby et al., 1983) End, Type – Zone des savanes orientales, Maintirano, Paris 66, holotype PC. *Taxithelium nepalense (Schwägr.) Broth. – Forêt d’Ankara, Sambirano, sur sol, sur calcaire, alt. 200 m, 21/08/1977, Tixier 10746, TAN. Taxithelium nossianum Besch. (Crosby et al., 1983) Taxithelium planulum Besch. (Crosby et al., 1983) Wijkia bessonii (Renauld et Cardot) H.A. Crum (Crosby et al., 1983) End, Type – Zone moyenne des forets, plateau d’Ikongo, Besson, herb (N.S.). Wijkia madagassa (Thér.) H.A. Crum (Crosby et al., 1983) End, Type – Vallée de l’Anony, sous-bois, 05/08/1921, Decary, holotyp PC. Wijkia rutenbergii (Müll.Hal.) H.A. Crum (Crosby et al., 1983) End, Type – Massif d’Ankaizina, herb (N.S.). Wijkia rutenbergii var. eflagellaris (Cardot) H.A. Crum (Crosby et al., 1983) Checklist of the bryophytes of Madagascar 245 Racopilaceae Powellia elliptica (Renauld) Zanten (Crosby et al., 1983; Zanten, 2003) End, Type – Entre Fianarantsoa et Mananjary, 1880, Besson, holotype PC. Powellia pocsii Zanten (Zanten, 2003) Racopilum africanum Mitt. (Crosby et al., 1983) Racopilum cardotii Renauld (Crosby et al., 1983) End, Type – Zone inférieure des forêts, Diégo-Suarez, 1890, Chénagon, PC; Andevoranto, 1896, Perrot, PC; holotype (N.S.). Racopilum madagassum Renauld (Crosby et al., 1983) Racopilum madagassum var. grandistipulaceum Thér. et Renauld (Crosby et al., 1983) End, Type – Zone du Plateau central, pays Betsileo, Villaume, holotype BM# Racopilum marginatum Dixon (Crosby et al., 1983) Racopilum mauritianum Müll.Hal. ex Besch. (Crosby et al., 1983) Racopilum microdictyon Besch. (Crosby et al., 1983) Racopilum perrieri Thér. (Crosby et al., 1983) End, Type – Entre la Sahambana et l’Ihosy, Forêt d’Analamanara, alt. 1300 m, 06/1919, Perrier de la Bathie, holotype PC. Racopilum plicatum Renauld et Cardot (Crosby et al., 1983) Racopilum tomentosum (Hedw.) Brid. (Crosby et al., 1983) Regmatodontaceae Regmatodon orthostegius Mont. (Crosby et al., 1983; Eakin, 1998) Rhabdoweisiaceae Oreoweisia erosa (Müll.Hal.) Kindb. (Crosby et al., 1983) Rhachitheciaceae Rhachithecium perpusillum (Thwaites et Mitt.) Broth. (Crosby et al., 1983) Rhacocarpaceae Rhacocarpus purpurascens (Brid.) Paris (Crosby et al., 1983; Frahm, 1996; Delgadillo, 1995) Rhizogoniaceae Pyrrhobryum spiniforme (Hedw.) Mitt. (Crosby et al., 1983; Frahm et al., 2003; Delgadillo, 1995) Rigodiaceae Rigodium toxarion (Schwaegr.) A. Jaeger var. toxarion (Crosby et al., 1983; Zomlefer, 1993) Rutenbergiaceae Rutenbergia borbonica Besch. (Crosby et al., 1983) Rutenbergia cirrata Renauld et Cardot (Crosby et al., 1983) End, Type – Forêt d’Analamazoatra, Camboué 295, holotype PC. Rutenbergia limbata (Hampe) Besch. (Crosby et al., 1983) End, Type – Forêt d’Analamazaotra, Borchgrevink, herb (N.S.). Rutenbergia madagassa Geh. et Hampe (Crosby et al., 1983) End, Type – Ambatondrazaka, herb (N.S.). 246 L. Marline et al. Rutenbergia madagassa var. flagelliformis Renauld et Cardot (Crosby et al., 1983) End, Type – Zone du Plateau central, Ambohimahatsara près d’Ambositra, 1892, Berthieu, holotype PC. Sematophyllaceae Acanthorrhynchium papillatum (Harv.) M. Fleisch. (Crosby et al., 1983; Tixier, 1995) Acroporium depressum Thér. (Crosby et al., 1983) End, Type – Mont Tsaratanana, forêt à sous-bois herbacé, alt. 1200-2400 m, Perrier de la Bathie, holotype PC. Acroporium laevifolium (Renauld) Broth. (Crosby et al., 1983) End, Type – Ankeramadinika, zone supérieure des forêts, 11/1897, Dorr, holotype PC. Acroporium megasporum (Duby) M. Fleisch. (Crosby et al., 1983) Acroporium megasporum var. densum (Renauld et Cardot) Wijk et Margad. (Crosby et al., 1983) End, Type – Forêt entre Fianarantsoa et Mananjary, 1892, Benson, holotype PC. Acroporium stellatum (Renauld et Cardot) Broth. (Crosby et al., 1983) End, Type – Zone inférieure des forêts, forêt à l’Est de Mahambo, environ d’Andevorante, au milieu de touffes de Schlotheimia perroti, 1890-1896, Perrot, holotype PC. Clastobryophilum bogoricum (Bosch et Sande Lac.) M. Fleisch. (Crosby et al., 1983) Macrohymenium acidodon (Mont.) Dozy et Molk. (Eakin, 1976; Crosby et al., 1983; Een, 1997) Macrohymenium muelleri Dozy et Molk. (Eakin, 1976) Macrohymenium rufum Müll.Hal. (Eakin, 1976) Macrohymenium strictum Bosch. et Sande Lac. (Eakin, 1976) Meiothecium crassirete (Renauld et Cardot) Cardot (Crosby et al., 1983) End, Type – holotype PC. Meiothecium madagascariense (Brid.) Broth. (Crosby et al., 1983; Een, 1997) Meiothecium undulatum (Renauld et Cardot) O’Shea (Crosby et al., 1983) End, Type – Zone moyenne des forêts, forêt d’Analamazaotra, Camboué, herb (N.S.). Radulina borbonica (Bel.) W.R. Buck (Crosby et al., 1983) Rhaphidorrhynchium angustissimum (Geh.) Broth. (Crosby et al., 1983) End, Type – Forêt de Vondrozona, Rutenberg, herb (N.S.). Rhaphidorrhynchium cambouei (Renauld et Cardot) Broth. (Crosby et al., 1983) End, Type – Zone du Plateau central, Ankadivavala, Campenon, herb (N.S.). Rhaphidorrhynchium rubricaule (Besch.) Broth. (Crosby et al., 1983) Sematophyllum coppeyi (Cardot) Broth. (Crosby et al., 1983) End, Type – Forêt de Manongarivo, 1909, Drouhard, holotype PC. Sematophyllum dixonii (Thér.) O’Shea (Crosby et al., 1983) End, Type – Vallée de l’Anony, Imerimandroso, Ambatondrazaka, Amparafaravola, 300 km NE, alt. 600 m, 05/08/1921, Decary & Waterlot, holotype PC. Sematophyllum dubium (Renauld) Broth. (Crosby et al., 1983) End, Type – Ankeramadinika, 11/1897, Dorr, holotype PC. Sematophyllum nanopyxis (Geh.) O’Shea (Crosby et al., 1983) End, Type – Zone des forêts, forêt de Vondrozona, Rutenberg, herb (N.S.). Checklist of the bryophytes of Madagascar 247 Sematophyllum obtusifolium (Renauld et Cardot) Broth (Crosby et al., 1983) Sematophyllum rigescens (Cardot) O’Shea (Crosby et al., 1983) End, Type – Zone inférieure des forêts, forêt de Fito, district de Tamatave, Perrot, holotype FH# Sematophyllum sinuosulum (Besch.) Broth. (Crosby et al., 1983) Sematophyllum sinuosulum var. latifolium (Cardot) Thér. (Crosby et al., 1983) End, Type – Zone du Plateau central, province d’Imamo, Camboué, holotype G# Sematophyllum subpinnatum (Brid.) E. Britton (Crosby et al., 1983) Sematophyllum subrugifolium Dixon (Crosby et al., 1983) End, Type – unknown. Sematophyllum subscabrellum Renauld et Cardot (Crosby et al., 1983) End, Type – Andevoranto, au milieu d’autres mousses, 1895, Perrot, holotype PC. Trichosteleum debettei var. laevisetum Cardot (Crosby et al., 1983) End, Type – unknown. Trichosteleum heterophyllum Bizot et Onr. (Crosby et al., 1983) End, Type – Nord Maromandia, au Sud de Diégo-Suarez, sur sol caillouteux, au bord d’un marigot permanent, sous forêt, alt. 50 m, 13/08/1973, Cremers 2735, holotype BR, isotypes BR, TAN, PC. Trichosteleum leviusculum Renauld et Cardot (Crosby et al., 1983) Trichosteleum microdontum (Besch.) Renauld (Crosby et al., 1983) Trichosteleum microdontum var. megapterum Renauld et Cardot (Crosby et al., 1983) End, Type – Zone moyenne des forêts, forêt d’Analamazaotra, Camboué & Campenon, herb (N.S.). Trichosteleum permixtum Cardot (Crosby et al., 1983) End, Type – Diégo-Suarez, Chénagon, holotype PC. Trichosteleum perrotii (Renauld et Cardot) Renauld et Cardot (Crosby et al., 1983) Trichosteleum perrotii var. eurydictyon Renauld et Cardot (Crosby et al., 1983) End, Type – Zone inférieure des forêts, île de Sainte-Marie, Mahavelona ou Foulpointe, Darbould, herb (N.S.). Trichosteleum pervilleanum (Müll.Hal. ex Geh.) W.R. Buck (Crosby et al., 1983) Trichosteleum trachypyxis (Geh.) Renauld et Cardot (Crosby et al., 1983) End, Type – Zone moyenne des forêts, forêt à l’est d’Ambatondrazaka, Rutenberg, herb (N.S.). Warburgiella leptorrhyncha (A. Jaeger) Broth. (Crosby et al., 1983) Serpotortellaceae Serpotortella chenagonii (Renauld et Cardot) W.D. Reese et R.H. Zander (Crosby et al., 1983) Serpotortella cyrtophylla (Besch.) W.D. Reese et R.H. Zander (Reese & Zander, 1987) Sphagnaceae Sphagnum capense Hornsch. (Eddy, 1985) Sphagnum ceylonicum Mitt. ex Warnst. (Eddy, 1985) Sphagnum cuspidatulum Müll.Hal. (Eddy, 1985) Sphagnum cuspidatum Hoffm. var. cuspidatum (Eddy, 1985) *Sphagnum davidii Warnst. – Massif de l’Ankaratra, S 19˚20’ E 47˚20’, alt. 2100 m, 24/10/1972, M.R. & C.A. Crosby 6636, TAN. 248 L. Marline et al. Sphagnum ericetorum Brid. (Eddy, 1985) Sphagnum magellanicum (Warnst.) A. Eddy subsp. grandirete (Eddy, 1985) End, Type – Andevoranto, Androrangobe, 1901, holotype PC. Sphagnum perichaetiale Hampe (Eddy, 1985) Sphagnum perrieri Thér. (Crosby et al., 1983) End, Type – Massif de l’Andringitra, alt. 1600-2500 m, 1921, Perrier de la Bathie, holotype PC. Sphagnum planifolium var. angustilimbatum (Warnst.) A. Eddy (Eddy, 1985) Sphagnum planifolium Müll.Hal. var. planifolium (Eddy, 1985) Sphagnum rutenbergii Müll.Hal. (Eddy, 1985) Sphagnum strictum (Müll.Hal.) A. Eddy subsp. pappeanum (Eddy, 1985) Sphagnum truncatum var. bordasii (Besch.) A. Eddy (Eddy, 1985) Sphagnum tumidulum Besch. var. tumidulum (Eddy, 1985) Sphagnum violascens Müll.Hal. (Eddy, 1985) Splachnaceae Brachymitrion moritzianum (Müll.Hal.) A.K. Kop. (Koponen & Weber, 1972; Crosby et al., 1983) Tayloria indica Mitt. (Crosby et al., 1983) Tayloria isleana (Besch.) Broth. (Crosby et al., 1983) Tayloria orthodonta (P.Beauv.) Wijk et Margad. (Crosby et al., 1983; Pócs & Geissler, 2002) Tayloria solitaria (Hedw.) T.J. Kop et W. Weber (Crosby et al., 1983; Pócs & Geissler, 2002) Stereophyllaceae Entodontopsis nitens (Mitt.) W.R. Buck et Ireland (Crosby et al., 1983) Entodontopsis tenuinervis (Broth.) W.R. Buck et Ireland (Crosby et al., 1983) Stereophyllum radiculosum (Hook.) Mitt. (Crosby et al., 1983) Symphyodontaceae Symphyodon pygmaeus (Broth.) S. He et Snider (Crosby et al., 1983; He & Snider, 2000) Thuidiaceae Pelekium chenagonii (Müll.Hal. ex Renauld et Cardot) Touw (Touw, 1976; Crosby et al., 1983) Pelekium gratum (P. Beauv.) Touw (Touw, 1976; Crosby et al., 1983; Touw, 2001) Pelekium intricatum (A. Jaeger) Touw (Pócs & Geissler, 2002) Pelekium investe (Mitt.) Touw (Touw, 1976; Crosby et al., 1983; Touw, 2001; Pócs & Geissler, 2002) Pelekium ramusculosum (Mitt.) Touw (Touw, 1976; Crosby et al., 1983) Pelekium varians (Welw. et Duby) Touw (Touw, 1976; Crosby et al., 1983) Pelekium versicolor (Müll.Hal.) Touw (Touw, 1976; Crosby et al., 1983; Touw, 2001) Rauiella praelonga (Schimp. ex Besch.) Wijk et Margad. (Touw, 1976; Crosby et al., 1983) Thuidium aculeoserratum Renauld et Cardot (Touw, 1976; Crosby et al., 1983) Thuidium assimile (Mitt.) A. Jaeger (Touw, 1976; Crosby et al., 1983) Thuidium pseudoinvolvens (Müll.Hal.) A. Jaeger (Touw, 1976; Crosby et al., 1983) Checklist of the bryophytes of Madagascar 249 Note on Riccardia: According to Grolle 1995, three species of the genus Aneura: A. caespitans Steph., A corbieri Steph and A. dismieri Steph. were placed on the name of Riccardia sp. The attribution of those species to the genus Riccardia has not been revised. But since the revision of the genus Riccardia from Africa and Madagascar is now in a progress, most of the morphological and reproductive characters of those three species seem to fit with those of Riccardia. We decided to keep the name used by Grolle (1995) and Wigginton (2009). CONCLUSION In total 1144 bryophyte species and infraspecific taxa, including 751 mosses, 390 liverworts and 3 hornworts are reported from Madagascar (Table 1). The following families: Dicranaceae (83 species and infraspecific taxa), Orthotrichaceae (66 species and infraspecific taxa) and Bryaceae (44 species and infraspecific taxa) are the most speciose for mosses and the Lejeuneaceae (203 species and infraspecific taxa), Frullaniaceae (38 species and infraspecific taxa) and Plagiochilaceae (21 species and infraspecific taxa) for liverworts. The lack of reported hornwort taxa undoubtedly represents a significant bias in historic collecting effort especially since more than 5 hornwort taxa are reported for Réunion Island, a landmass both a fraction of the age and size of Madagascar (Ah-Peng et al., 2010). Future floristic exploration in Madagascar should target such little known groups as they are likely to harbour a wealth of unknown diversity. In this checklist, 13 mosses, 10 liverworts and one species of hornwort are newly recorded for Madagascar. The majority of these new records are reported from collections in the National Herbarium of Madagascar (TAN) and from recent collections by the authors. Two species: Fissidens cyathaeicola Brugg.Nann. and Taxithelium nepalense (Schwägr.) Broth. are reported for the first time for the Sub-Saharan Africa region. The bryoflora of Madagascar displays remarkably high levels of endemism, 33.82% of mosses and 18.97% of liverworts are unique to the island, providing ample support for its status as a global hotspot of biodiversity. The moss Table 1. Summary of the bryoflora of Madagascar Mosses Liverworts Hornworts Total Taxa 751 390 3 1144 Genera 186 86 3 275 Families 59 30 2 91 Endemics 254 74 0 328 33.82% 18.97% Endemicity (%) 28.67% 250 L. Marline et al. families with the largest number of endemic species are the Dicranaceae (41/83), Orthotrichaceae (38/66) and Sematophyllaceae (22/40). Of the 74 endemic liverwort species recorded on the island, 59 belong to the family Lejeuneaceae, of which the most important is the epiphyllous genus Cololejeunea which has radiated throughout Madagascar and the neighbouring islands (Pócs, 1997). The number of endemics should be regarded with caution as many of the species were described a long time ago and have yet to form part of a recent revision: this figure will need to be continuously updated. To help in further research e.g. taxonomic revision, we listed the type specimens of endemic species including specimen information and host herbarium. Unfortunately a few proportion (18) of type specimens are poorly detailed, often providing only “Madagascar” as the type location, we did not manage to identify the host herbarium of 56 endemic species. Since much of the original bryological exploration of Madagascar took place during the late 1800’s (Dorr, 1997) it is not known what proportion, if any, of the recorded taxa have been lost due to anthropogenic destruction of habitats for farming and logging. Many of the sites which were frequented by early bryologists have been lost due to such activities. New inventories of intact and unexplored habitats on the island should prove invaluable for verifying many of the old records presented in this checklist and will undoubtedly reveal many new and interesting records for the island. Much work remains to be done to improve knowledge on the diversity and distribution of bryophytes in Madagascar. For example, collections of bryophytes in the National Herbarium of Madagascar (TAN) need to be revised and fully identified. More fieldwork to collect bryophytes in poorly investigated areas should be planned, especially in areas such as the dry forests. Acknowledgments. The first author was supported by a grant from the French Government (BGF). The authors are grateful to the BRYOLAT project (French Foundation for Research in Biodiversity), FEDER POE 3.21, MOVECLIM (ANR Net Biome) for financial support. The last author thanks the Claude Leon Harris Fellowship for supporting her research. 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