Cryptogamie,Bryologie, 2012,33 (3): 199-255
@ 2012Adac. Tous droits r6serv6s
Checklist of the bryophytesof Madagascar
aUniversit| de La Rhunion, (JMR PVBMT,
75 avenue Ren€ Cassin, BP 7151,97 715 Saint Denis Cedex 9, France
b(Jniversity of Cape Town, Botany Department,
Private Bag X3,7701 Rondebosch, South Africa
cUniversitd d'Antananarivo, Facult| des Sciences,D1partement de Biologie
et Ecologie V4g€tales, BP 566, Antananarivo, Madagascar
dMittouri Botanical Garden Madagascar,
PO Box 3391 Antananarivo 101 Madagascar
eMus1um National d'Histoire Naturelle, UMR CNRS-7205,
D4partement de Syst4matiqueet Evolution,5T rue Cuvier,75005 Paris France
fUniversity of Sain Malaysia, School of Biological Sciences,
11800 USM Pulau Penang, Malaysia
(Received26 August 2011, accepted11 January 2012)
Abstract - This updatedchecklistof the bryophytesof Madagascarwas compiled from the
literature, herbarium and recent collections.A total of 1144speciesand infraspecifictaxa
are recorded for Madagascarcomposedof 751-mosses,390 liverworts and 3 hornworts.
28.67"/"of the speciesreported for the Island are endemics.Twenty-four taxa are newly
recorded for Madagascarand two of them, FissidenscyathaeicolaBrugg.Nann. and
Taxitheliumnepalense(Schwagr.)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 o_fAfrica. It is the world's fourth largest
island, with a total area of 590 000 k^2, the distance between Cap d'Am"bre
x Author for correspondence and reprints: marlinelova@gmail.com
Cryptogamie,Bryologie,2012,33 (3): 199-255
@ 2012Adac. Tous droits r6serv6s
Checklist of the bryophytes of Madagascar
aUniversit€ de La R1union, UMR PVBMT,
15 avenue Ren€ Cassin, BP 7151, 97 715 Saint Denis Cedex 9, France
b(Jniversity of Cape Town, Botany Department,
Private Bag X3,7701 Rondebosch, South Africa
cUniversit€ d'Antananarivo, Facult1 des Sciences,D1partement de Biologie
et Ecologie V|g1tales, BP 566, Antananarivo, Madagascar
dMittouri B otanical Garden Madagascar,
PO Box 3391 Antananarivo 101 Madagascar
eMustum National d'Histoire Naturelle, UMR CNRS-7205,
D,|partement de Syst4matiqueet Evolution, 57 rue Cuvier,75005 Paris France
fUniversity of Sain Malaysia, School of Biological Sciences,
11800 USM Pulau Penang, Malaysia
(Received26 August 2011, accepted11 January 2012)
Abstract - This updatedchecklistof the bryophytesof Madagascarwas compiled from the
literature, herbarium and recent collections.A total of 1144speciesand infraspecifictaxa
are recordedfor Madagascarcomposedof 751 mosses,390 liverworts and 3 hornworts.
28.67o/"of the speciesreported for the Island are endemics.Twenty-four taxa are newly
recorded for Madagascarand two of them, FissidenscyathaeicolaBrugg.Nann. and
Taxitheliumnepalense(Schwagr.)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 II"57' and 25"39'
latitude, 400 km from the eastern coast o_fAfrica. It is the world's fourth largest
island, with a total area of 590 000 k^2, the distance between Cap d'Anibre
* Author for correspondenceand 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 descendsmore 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 1500-2000years since the
arrival of the first humans (Harper et al., 2007). At present, only I2"/" 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 a1.,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
(800-1800m); Evergreen humid forest, lower montagne (1800-2000m); Mountain,
Schrubland (Philipio) (> 1800 m); Evergreen, sclerophyllous (Uapaca) woodland
(800-2000m) and Marshland (0-800 m)
The deciduous formations (West and South) are: Coastal forest (West);
Deciduous, seasonallydry, 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 l-9th century that colleitors first began to collect bryophytei in the
Island (Dorr, 1997). There, are more than 40 botanists that collected bryophytes
on the island during the 19rncentury. The majority of which were French botanists
and explorers. Collections through this period, by Cardot, Tixier, Cramers,
Ch6nagon, 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 specimensfor 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.
201
Checklist of the bryophytes of Madagascar
Madagascar
*
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David Du Puy & Justin Mnat
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permission of IRD).
METHODS
13,
This checklist was compiled from the literature, data from various
herbaria, and recent collections by th-e authors. This work was greatly facilitated
by the ground work of O'Shea (2006) and Wiqginton 9909) for their respective
checkhsls of mosses. liverworts and hornworts for sub-Saharan Africa.
L.i|darline 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
(200e).
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
Synopsisof the generaand families of Liverworts, Hornworts and Mosses
DIVISION MARCHANTIOPHYTA
(LIVERWORTS)
CLASS MARCHANTIOPSIDA
SubclassMarchantiidae
ORDER MARCHANTIALES
Family Aytoniaceae
Asterella (2)
Plagiochasma (1)
Family Cyathodiaceae
Cyathodium (I)
Family Dumortieraceae
Dumortiera (1.)
Family Marchantiaceae
Marchantia (2)
Family Ricciaceae
Riccia (5)
Family Targioniaceae
Targionia (2)
Checklist of the bryophytes of Madagascar
CLASS JUNGERMANNIOPSIDA
SubclassPelliidae
ORDER FOSSOMBRONIALES
Family Fossombroniaceae
Fossombronia (I)
ORDER PALLAVICINIALES
Family Pallaviciniaceae
Pallavicinia (I)
Symphyogyna (2)
Symphyogynopsis (1.)
SubclassMetzgeriidae
ORDER PLEUROZIALES
Family Pleuroziaceae
Pleurozia (l\
ORDER METZGERIALES
Family Aneuraceae
A neura (I)
Riccardia (9)
Family Metzgeriaceae
Merzgeria (6)
SubclassJungermanniidae
ORDER PORELLALES
Family Frullaniaceae
Frullania (38)
Family Lejeuneaceae
Acanthocoleus (2)
Acrolejeunea (3)
Archilejeunea (4)
Brachiolejeunea(I)
Bryopterls (1)
Caudalejeunea (4)
Ceratolejeunea (6)
Cheilolejeunea (I4)
Cololejeunea(54)
Colura (9)
Diplasiolejeunea (26)
Drepanolejeunea(9)
Frullanoides (1.\
Haplolejeunea (I)
Lejeunea (22)
Lepidolejeunea (1,)
Leptolejeunea (2)
Lopholejeunea (I2)
Marchesinia (2)
Mastigolejeunea (2)
Metalejeunea(I)
Microlejeunea (5)
Odontolejeunea(l)
Otolejeunea(3)
Prionolejeunea (I)
Ptychanthrzs(1)
Pycnolejeunea(I)
Schiffne r io I ej eun ea (6)
Symbiezidium (2)
Taxilejeunea (3)
Thysananthus(l)
Xylolejeunea (1,)
Family Porellaceae
Porella (5)
Family Radulaceae
Radula (ll\
ORDER JUNGERMANNIALES
Family Adelanthaceae
Adelanthus (2)
Family Arnelliaceae
Gongylanthus (I)
Family Balantiopsidaceae
Isotachis (I)
Family Calypogeiaceae
Calypogeia (7)
Mnioloma (I)
Family Cephaloziaceae
Cephalozia (l)
Iwatsukia (I)
Family Cephaloziellaceae
Amphicephalozia (I)
Cephaloziella (2)
Cylindrocolea (2)
Kymatocalyx (3)
Family Herbertaceae
Herbertus (2)
Familly Jamesoniellaceae
Cryprochila (1)
Cuspidatula (I)
Denotarisia (I)
Syzygiello (3)
Family Jungermanniaceae
Gottschelia (l)
Itlotoscyphus(1)
Solenostoma(4)
Family Lepidoziaceae
Amazoopsls (1)
Bazzania (ll)
Kurzia (2)
Lepidozia (2)
Telaranea(3)
Family Lophocoleaceae
Chiloscyphus (8)
Clasmatocolea(1)
Conoscyphus (I)
Heteroscyphus (2)
Leptoscyphus (2)
203
204
L.Marline et al.
Family Mastigophoraceae
Mastigophora (I)
Family Plagiochilaceae
Plagiochila (2I)
Family Scapaniaceae
Anastrophyllum (I)
Andrewsianthus (I)
Plicanthus (2)
Family Schistochilaceae
Gottschea(I)
Schistochila (2)
DrvrsroN ANTHOCBROTOPHYTA(HORNWORTS)
CLASS ANTHOCEROTOPSIDA
ORDER ANTHOCEROTALES
Family Anthocerotaceae
Anthoceros (1)
Dendroceros (I)
ORDER NOTOTHYLADALES
Family Notothyladaceae
(1)
Phaeoceros
DrvrsroN BRYOPHYTA (MOSS)
SuperclassI
CLASS SPHAGNOPSIDA
ORDER SPHAGNALES
Family Sphagnaceae
Sphagnum(1.6)
SuperclassII
CLASS ANDREAEOPSIDA
ORDER ANDREAEALES
Family Andreaeaceae
Andreaea(2)
SuperclassIII
CLASS POLYTRICHOPSIDA
ORDER POLYTRICHALES
Family Polytrichaceae
Atrichum (1)
Pogonatum(4)
Polytrichastrum(I)
Polytrichum (5)
CLASS BRYOPSIDA
SUBCLASS FUNARIIDAE
ORDER GIGASPERMALES
Family Gigaspermaceae
Gigaspermum(l)
Oedipodiella(I)
ORDER FUNARIALES
Family Funariaceae
Entosthodon(4)
Funaria (5)
Physcomitrium(2)
SUBCLASSDICRANIDAE
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 (I)
Distichium (I)
Ditrichum (4)
Garckea (1,)
Pleuridium (I)
Familv Dicranaceae
Aniiothecium (I)
Aongstroemia (2)
Dicranella (6)
Dicranoloma (3)
Dicranum (1.)
Holomitrium (ll)
Leptotrichella (2)
Leucoloma (55)
Microcampylopus (I)
Pseudephemerum(I)
Family Erpodiaceae
Erpodium (2)
Family Eustichiaceae
Eusrichia (1)
Checklist of the bryophytes of Madagascar
Family Fissidentaceae
Fissidens (40)
Family Leucobryaceae
Atractylocarpus (l)
Campylopus (23)
Leucobryum (19)
Ochrobryum (I)
Family Rhachitheciaceae
Rhachithecium (I)
Family Rhabdoweisiaceae
Oreoweisia (l)
ORDER POTTIALES
Family Pottiaceae
Anoectangium (3)
Barbula (6)
B ry o ery throphy llum (I)
Chionoloma (l)
Didymodon (l)
Gymnostomiella (1)
Gymnostomum (1)
Hymenostylium (l)
Hyophila (7)
Leptodontium (3)
Splachnobryum (2)
Streptopogon (2)
Tortella (I)
Tortula (4)
Trichostomum (6)
Weisiopsis (I)
Weissia(I)
Family Serpotortellaceae
Serpotortella (2)
SUBCLASS BRYIDAE
Super order Bryanae
ORDER SPLACHNALES
Family Meesiaceae
Leptobryum (l)
Family Splachnaceae
Brachymitrion (1,)
Tayloria (4)
ORDER BRYALES
Family Bryaceae
Anomobryum (2)
Brachymenium (12)
Bryum (26)
Mniobryum (I)
Rhodobryum (3)
Family Mniaceae
Epipterygium (l)
Mielichhoferia (2)
Plagiomnium (I)
Pohlia (4)
Pseudopohlia (I)
Schizymenium (2)
ORDER BARTRAMIALES
Family Bartramiaceae
Bartramia (2)
Breutelia (8)
Philonotis (7)
ORDER ORTHOTRICHALES
Family Orthotrichaceae
Cardotielh @)
Groutiella (l)
Macrocoma (l)
Macromitrium (24)
Schlotheimia (31)
Ulota (I)
Zygodon (4)
ORDER HEDWIGIALES
Family Hedwigiaceae
Braunia (I)
Bryowijkia (I)
Hedwigia (1,)
Family Rhacocarpaceae
Rhacocarpus (I)
ORDER RHIZOGONIALES
Family Rhizogoniaceae
Pyrrhobryum (1)
Seper order Hypnanae
ORDER HYPNODENDRALES
Family Racopilaceae
Powellia (2)
Racopilum (I0)
ORDER HOOKERIALES
Family Daltoniaceae
Adelothecium (1.)
Calyptrochaeta (I)
Daltonia (5)
Distichophyllum (2)
Family Hypopterygiaceae
Hypopterygium (I)
Lopidium (l)
Family Leucomiaceae
Leucomium (l)
Family Pilotrichaceae
Callicostella (I3)
Cyclodictyon (3)
Hookeriopsrs (1)
Lepidopilidium (5)
Lepidopilum (3)
ORDER HYPNALES
Family Amblystegiaceae
Amblystegium (2)
Hygrohypnum (I)
205
206
L.I|y4arline et al.
Family Anomodontaceae
Herpetineuron (I)
Family Brachytheciaceae
Aerolindigia (I)
Brachythecium (2)
Eurhynchium (2)
Helicodontium (I)
Palamocladium (I)
Rhynchostegiella (l)
Rhynchostegium (5)
Schimperella (l)
Squamidium (I)
Family Catagoniaceae
Catagonium (I)
Family Cryphaeaceae
Cryphaea (2)
Schoenobryum (l)
Family Entodontaceae
Campylodontium (I)
Entodon (5)
Erythrodontium (I)
Mesonodon (I)
Family Fabroniaceae
Fabronia (8)
Rhizofabronia (I)
Family Hypnaceae
Ectropothecium (12)
Gammiella (I)
Glossadelphus (3)
Hypnum (2)
Mittenothamnium (7)
Phyllodon (2)
Rhacopilopsis (2)
Taxiphyllum (I)
Vesicularia (9)
Family Leptodontaceae
Leptodon (I)
Family Leskeaceae
Pseudoleskea(I)
Family Leucodontaceae
Leucodon (2)
Pterogoniadelphus (I)
Pterogonium (2)
Family Meteoriaceae
Aerobryopsis (1)
Barbella (l)
Floribundaria (4)
Papillaria (5)
Trachypodopsis (I)
Trachypus (2)
Family Neckeraceae
Homaliodendron (2)
Neckera (2)
Neckeropsis (4)
Orthostichella (5)
Pilotrichelh @)
Pinnatella (I)
Porotrichum (5)
Thamnobryum (l)
Family Phyllogoniaceae
Phyllogonium (l)
Family Prionodontaceae
Prionodon (1)
Family Pterigynandraceae
Trachyphyllum (2)
Family Pterobryaceae
Calyptothecium (2)
HildebrandtieUa (3)
Jaegerina (2)
Orthostichidium (I)
Orthostichopsis (5)
Pterobryopsrs (1)
Renauldia (3)
Family Pylaisiadelphaceae
Isopterygium (14)
Taxithelium (7)
Wijkia @)
Family Regmatodontaceae
Regmatodon (I)
Family Rigodiaceae
Rigodium (l)
Family Rutenbergiaceae
Rutenbergia (5)
Family Sematophyllaceae
Acanthorrhynchium (I)
Acroporium (5)
Clastobryophilum (1)
Macrohymenium (4)
Meiothecium (3)
Radulina (I)
Rhap hido rrhynchium (3)
Sematophyllum (11,)
Trichosteleum (I0)
Warburgiella (I)
Family Stereophyllaceae
Entodontopsls (2)
Stereophyllum (1,)
Family Symphyodontaceae
Symphyodon (1,)
Family Thuidiaceae
Pelekium (7)
Rauiella (l)
Thuidium (3)
Checklist of the bryophytes of Madagascar
207
Liverworts reported from Madagascar
Adelanthaceae
Adelanthus decipiens (Hook.) Mitt. (P6cs & Geissler,2002)
Adelanthus lindenbergianus (Lehm.) Mitt. (Grolle, 1995)
Aneuraceae
*Aneura pseudopinguis (Herzog) P6cs - Ambavaniasy, rdserve de Vohimana, bois
mort oblique en d6composition, alt. 849 ffi, S 18"55'43.6" E 48"29'56.6",
1510412010,Reeb, Andriamananatena & Bidault CRAE 106, TAN, det.
Reeb & Andriamananatena; Ambavaniasy, r6serve de Vohimana, for€t
ombrophile, tronc vertical d'arbre vivant, alt. 830 m, S 18'55'09.33" E
48'30'49.0", L610412010,,
Reeb, Andriamananatena & Bidault CRAE120
TAN, det. Reeb & Andriamananatena; Anjangoveratra, forOt du
Makirovana, forOt ombrophile dense orientale, au dessus d'arbre mort
horizontal, alt. 432 m, S L4"10'I3.9" E 49"57'12.9", 04105120L0,Reeb
CR220, TAN, det. Reeb.
Riccqrdia am&zonfca (Spruce) Schiffn. ex Gradst. (P6cs & Geissler,2002)
Riccardi& erosa (Steph.) E.W. Jones (P6cs & Geissler,2002)
Riccardia longispica (Steph.) Pearson (Grolle,1995; P6cs & 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 caespitansSteph.) (Grolle, 1995)
Riccardia sp. (Aneura corbieri Steph.) (Grolle ,1995)
Riccardia sp. (Aneura dismleri Steph.) (Grolle ,1995)
Arnelliaceae
Gongylanthus ericetorum (Raddi) Nees (Grolle, 1995)
Aytoniaceae
Asterella coronats (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, I995;P6cs & Geissler,2002)
Calypogeiaceae
Calypogeia annabonensis Steph. (P6cs & Geissler,2002)
Calypogeia arguta Nees & Mont. (Grolle, 1995)
Calypogeia bidentula (F. Weber) Nees (Grolle, 1995)
*Calypogeiafissa (L.) Raddi - Ankaratra, sur sol, alt. 2000 m,2710311972,
Cremers
1947, TAN.
Calypogeia longifulfa Steph. (Grolle, 1995; P6cs,2005)
Calypogeia mqscarenensis Bischl. (Grolle, 1995)
Calypogeia miuostipula (Steph.) Steph. (Grolle, 1995)
Mniolomafuscum (Lehm.) R.M. Schust. (Grolle, 1995;P6cs & Geissler,2002)
L.Marhne et al.
Cephaloziaceae
Cephalozia connivens (Dicks.) Lindb. subsp.fissa Vdria (Grolle, 1995; P6cs, 1995;
P6cs & Geissler,20D2)
Iwatsukia jishibae (Steph.) N. Kitag. (P6cs, 1995;P6cs & Geissler,2002)
Cephaloziellaceae
Amphicephalozia geisslerae P6cs & Ydi.a (P6cs & Y6ia,200I; P6cs & Geissler,
2002) End, Type - Antsiranana, Manongarivo Special Reserve, NW
Madagascar, alt. 1300-1350m, 1010311999,
Geissler 19708/3, holotype G,
paratype EGR, G, PRC.
Cephaloziella kiaeri (Austin) S.W. Arnell (Grolle, 1995; P6cs & Geissler,2002)
Cephaloziella sp. (Grolle, 1995)
Cylindrocolea atroviridis (Sim) V6ria (Grolle, 1995)
Cylindrocolea madagascariensis (Steph.) R.M. Schust. (Grolle, 1995) End, Type Imerina, Hildebrandt, herb (N.S.).
Kymatocalyx dominicensis (Spruce) V6ria (Gradstein & Vriria,1999)
Kymatocalyx madagascariensl's (Steph.) Gradst. & V6ila (Grolle, 1995; P6cs,
1995;Gradstein &Y6i,a,1999; P6cs & Geissler,2}D2)
Kymatocalyx rhizomatica (Herzog) Gradst. &Ydi'a (P6cs & Geissler,2002)
Cyathodiaceae
Cyathodium africanum Mitt (P6cs & Geissler, 2002)
Dumortieraceae
*Dumortiera hirsura (Sw.) Nees - Anjozorobe, forOt ombrophile d'altitude, sur
des terres humides, 0211959,Bosser 12648,TAN; Marojejy National Park,
Talus terreux ombrag6, alt. 690 m,S 1,4"26'07.8"E 49"45'49.9",2711112009,
Ah-Peng 349, det. Ah-Peng; Ambavaniasy, r6serve de Vohimana, forOt
ombrophile d'altitude, au pied d'une cascade, alt.849 m, S 1-8"55'43.6"E
48"29' 56.6"1510412010,Reeb, Andriamananatena & Bidault CRAEQ3,
TAN, det. Reeb; Anjangoveratra, forOt du Makirovana, sur falaise
suintante.alt. 508 m. S 14"10'17.4"8 49"57'07.5".0710512010.
Reeb CR302.
TAN, det. Reeb.
Fossombroniaceae
Fossombronia sp. (P6cs & Geissler,2002)
Frullaniaceae
Frull anis anderssonii Aongstr. (Grolle, 1995)
Frullania angulata Mitt. var. angulata (Grolle, 1995; P6cs & Geissler,2002)
Frullanis angulata var. laciniatqDemaret et Vanden Berghen (Grolle, 1995)
Frullania apicalis Mitt. (Grolle, 1995; P6cs & Geissler,2002)
Frullania apiculata (Reinw. et al.) Nees (Grolle, 1995; P6cs, 1995; P6cs &
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 depressaMitt. (Grolle, 1995)
Frullania diptera (Lehm.) Drdge (Grolle, 1995)
Checklist of the bryophytes of Madagascar
..
\
*Frullania ecklonii (Spreng.)Gottscheet al. - Ambatovory,Tananarive,vestige
Bosser11021,TAN.
de forOt,sur un tronc d'arbre,0411957,
Frullania ericoides(Nees)Mont. (Grolle, 1995)
Frullania gabonensisVanden Berghen (Grolle, 1995;P6cs,1995)
Frullania grossiclava Steph. (Grolle, 1995; P6cs, 1995; Grolle, 2001,;P6cs &
Geissler,2002)
Frullania humbertii Vanden Berghen(Grolle, 1995)
Frullania imerinensisSteph.(Grolle, 1995;P6cs& Geissler,2}l2)
Frullania lindenbergii Lehm. (Grolle, 1995;P6cs, 1,995;Grolle, 2001; P6cs &
Geissler,2002)
Frullania longistipula Steph.(Grolle, 1995)End, Type - Imerina, Ankadivavala,
Cambou( I 62, holotype G.
Frullania loricatq Pearson(Grolle, 1995;P6cs& Geissler,2002)
Frullania loricqtq var. Iaxa Pearson(Grolle, 1995)End, Type - unknown.
Frullania nodulosa (Reinw. et al.) Gottsche et al. var. nodulosa (Grolle, 1,995;
P6cs,1995)
Frullania obscurifoliaMitt. (Grolle, 1995;P6cs,1995)
Frullania purpurea Steph.(Grolle, I995;P6cs,1995;P6cs& Geissler,2002)
Frullqnia repandistipulaSandeLac. (Grolle, 1995)
Frullunia rigida Steph.(Grolle, 1995)End, Type - unknown.
Frullania schimperi var. luciniata Vanden Berghen (Grolle, 1995)End, Type Onraedt74M8078c,holotype BR.
Frullania schimperiNees var. schimperi(Grolle,1995;P6cs& Geissler,2002)
Frullania sematavar.pertenuis(Nees)Gottscheet al. (Grolle,1995;P6cs,1995)
Frullania serrata Gottschevar. serr&ra(Grolle,1995)
Frullania socotranaMitt. (Grolle, 1995)
Frullania spongiosaSteph.(Grolle, 1995;P6cs,1995)
Frullaniu trinervis (Lehm.) Drdge (Grolle, 1995)
Frullania usambaranaSchiffn.ex Steph.(Grolle, 1995;P6cs,1995)
Frullania usambaranayar. reducts Vanden Berghen(Grolle, 1995)
Frullaniq vanden-bergheniiP6cs(Grolle, 1995;P6cs,t995)
Frullania variegataSteph.(Grolle, 1995;P6cs,1995;P6cs& Geissler,2002)
Herbertaceae
Herbertus dicranus (Taylor ex Gottsche et al.) Trevis. (Grolle, 1995;P6cs, 1995;
Grolle, 2001.;Hodgetts,2008)
Herbertusjuniperoideus (S*.) Grolle (Grolle, t995; Grolle,200L:'P6cs& Geissler,
2002;Hodgetts,2008)
Jamesoniellaceae
Cryptochila grandiflora (Lindenb. et Gottsche)Grolle (Grolle. 1995)
Cuspidatulacontracra(Reinw.et a/.) Steph.(P6cs,1995)
Denotarisia linguifulia (De Not.) Grolle (P6cs& Geissler,2002)
Syzygiellapurpurascens(Steph) Feldberget al. (Grolle, 1995;P6cs,1995;P6cs&
Geissler,2002)
Syzygiellaconueta (Gottsche)Spruce(Grolle, 1995;P6cs& Geissler,2}02)
Syzygiella geminifulia (Mitt.) Steph. (Grolle, 1995;P6cs, 1995;P6cs & Geissler,
2002)
2I0
L.Marline et al.
Jungermanniaceae
Gottschelia schizopleura (Spruce) Grolle (Grolle, 1995; P6cs, 1995; P6cs &
Geissler, 2002)
Notoscyphus lutescens (Lehm. et Lindenb.) Mitt. (Grolle, 1995; P6cs & Geissler,
2002)
Solenostoma borgenii (Gottsche) Steph. (Grolle, 1995)
Solenostoma onraedtii (Vifia) Vdila et al. (Grolle, L995)
Solenostoma renauldii (Vdna) Vdila et al. (Grolle, L995)
Solenostoma sphaerocarpum (Hook.) Steph. (Grolle, 1995)
Lejeuneaceae
Acanthocoleus abetans (Lindenb. e/ Gottsche) Kruijt (Kruijt, L988; Grolle, 1995)
Acanthocoleus madagascariensis (Steph.) Kruijt (Grolle, 1995; P6cs, I995;P6cs &
Geissler.2002)
Acrolejeunea aulacophora (Mont.) Steph. (Grolle, 1995)
Acrolejeunea emergens (Mitt.) Steph. var. emergens (Grolle, 1995; P6cs &
Geissler,2002)
Acrolejeunes pycnoclada (Taylor) Schiffn. (Grolle, 1995)
Archilejeunea alata Steph. (Grolle, L995)
Archilejeuneq brevilobula Steph. (Grolle, 1995) Bnd, Type - Oriental coast,
Sakara, hototype G.
Archilejeunea helenae P6cs et Gyarmati (P6cs & Sass-Gyarmati, 2006) End, Typ"
- Mananara-Nord Biosohere Reserve. lowland rainforest on the East
slopes of Mahavoho Hili, epiphyllous on relatively rigid leathery leaf, alt.
250-350 m, S 16"27' E 49"46.9-47.5',14-1610811998,
P6cs & Szab6 9878/
FM,holotype EGR.
Archilejeuneq nobilis (Gottsche) Steph. (Grolle, 1995) End, Type - unknown.
Brachiolejeunea phyllorhza (Nees) Kruijt e/ Gradst. (Grolle, 1995)
Bry op teris gaudichaudii G ottsche (GrolIe, I99 5; P6cs, 1995)
Caudalejeunea grolleana Gradst. (Grolle, 1995) End, Type - Nosy Be, Lokobe
natural reserve, on bark in primary forest, alt. 100 m, L5l07ll97I, Onraedt
M5119, holotype BR, isotype JE.
C audal ej eunea h anningt oni i (Mitt.) Schiffn. ( Grolle, 1995)
Caudalejeunea lewallei Vanden Berghen (Grolle, 1995)
C audal ej eunea recurv is tipul a (G ottsche) Schiffn (Grolle, 1995)
Ceratolejeunea andringitrae P6cs (P6cs, 20lla) 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. 750-1000m, S 22"13'E 47"I-2',20-2310911994,
P6cs & A. Szab6 9472/
AF, holotype EGR, isotypes MO, TAN.
Ceratolejeunea belangeriana (Gottsche) Steph. (Bonner, 1953; Grolle, 1995; P6cs
& Geissler,2002;P6cs, 20IIa)
C erato I ej eunea div ersicornua Steph. (P6cs, 20IIa)
Ceratolejeuneq cornutu (Lindenb.) Schiffn. (Grolle, 1995: P6cs, 20lla)
Ceratolejeunea papuliflora Steph. (Bonner, 1953; Grolle, 1995; P6cs, 2011a)
Ceratolejeunea saroltae P6cs (P6cs, 2011a) End, Type - Toamasina Province,
Maromizaha forest, Mossy montane rainforest with bamboo (l/asrzs sp.),
undergrowth on the summit ridge of Mt. Maromizaha, south of the
Andasibe National Park, on twigs, alt. 1080-1214m, Sarolta P6cs 9890/BJ,
2610911998,
holotype EGR, paratype EGR.
Checklist of the bryophytes of Madagascar
21,1
Cheilolejeuneacordigera(Steph.)Grolle (Grolle, 1995;P6cs& Geissler,2002)
Cheilolejeunea cordistipula (Steph.) Grolle ex E.W. Jones (Grolle, 1995;
Malombe,2007)
Cheilolejeuneqdecurciva (Sande Lac.) R.M. Schust. (Grolle, 1,995:P6cs &
Geissler,2002;Malombe, 2007)
P6cs& Geissler,2002;
Cheilolejeune&intertexta(Lindenb.)Steph.(Grolle, 1,995;
Malombe,2007)
Cheilolejeunea krakqkammae (Lindenb.) R.M. Schust. (Grolle, 1995; P6cs &
Geissler,2002;Malombe 2007)
(Gottsche)R.M. Schust.(Grolle, 1995;P6cs& Geissler,
Cheilolejeuneamontagned
2002;Malombe,2007)
Cheilolejeunes pluriplicata (Pearson) R.M. Schust. (Grolle, 1995; Malombe,
2oo7)
Cheilolejeunea pocsii E.W. Jones (Grolle, 1,995;Malombe,2007)
Cheilolejeunea serpentina (Mitt.) Mizut. (Grolle, 1,979:Grolle, 1995: P6cs, 1995;
P6cs & Geissler, 2002;Malombe, 2007)
Cheilolejeunea sunepens (Mitt.) E.W. Jones (Grolle, 1995;P6cs & Geissler,2002)
Cheilolejeunea trapezia (Nees) Kachroo e/ R.M. Schust. (Malombe,2007)
Cheilolejeunea triforia (Reinw. et al.) Mizut. (Grolle, 1995; P6cs & Geissler,2002)
Cheilolejeunea usambarana (Steph.) Grolle (Malomb e, 2007)
Cheilolejeunea xanthocarpa (Lehm. & Lindenb.) Malombe (P6cs, 1,995;
Malombe ,2007)
Cololejeunea adnata Tixier (Tixier, 1985; Grolle, 1995) End, Type - Tamatave,
Tixier 7637,
Foulpointe, forOt d'Analalava, alt. 100 m, 0511211,975,
holotype PC.
Cololejeunea andapaniq Tixier (Tixier, 1985; Grolle, 1995) End, Type - Di6goSuarez, massif de l'Anjanaharibe, Ouest d'Andapa, Haute Andramota,
bassin de la rividre Lokoho, forOt ombrophile sur gneiss et granite,
Humbert, Capuron & Cour l/E,holotype PC.
alt. 700 m,1011211950,
Cololejeunea androphylla var. madecqssa Tixier (Grolle, 1995) End, Type Antananarive, Mandialaza, ah.1200 m, Tixier 7670,holotype PC.
Cololejeunea ankaiana Tixier (Tixier, 1985; Grolle, 1995) End, Type - Route de
Tamatave, Moramanga - Anosibeanala, alt. 1000 m,I8l4lI978, Tixier s.n.,
holotype PC.
Cololejeune& appressa (A. Evans) Benedix (Grolle, 1,995;P6cs & Geissler,2002)
Cololejeunea auriculsta (E.W. Jones) R.M. Schust. (Tixier, 1985; Grolle, f995;
P6cs & Geissler.2002)
Cololejeunea bidentula (Steph.) E.W. Jones (Grolle, 1995)
Cololejeunea bolombensis (Steph.) Vanden Berghen (Tixier, 1985; Grolle, 1995;
P6cs & Geissler,2}l2)
Cololejeunea borbonica Tixier (Grolle, 1995)
Cololejeunea csmusii Tixier (Tixier, 1985; Grolle, 1995) End, Type
Tixier
Antananarivo, Ambatoloana, Andasibe, aII. 7200 m, 10111,11,976,
9593,holotype PC.
Massif de
Cololejeunea capuronii Tixier (Grolle, 1995) End, Type
l'Anjanaharibe, ) l'Ouest d'Andapa, Andramonta (pente et sommet
nord), alt.500-700 m, Humbert, Capuron & Cours s.rz.,holotype PC.
Cololejeunea cardiocarpa (Mont.) A. Evans (Tixier, 1985;Grolle, 1995)
Cololejeunea cristara (Steph.) R.M.Schust. (Tixier, 1985; Grolle, 1995)
2r2
L.Marline et al.
Cololejeuneacuneata(Lehm. etLindenb.) Herzog (Grolle,1995; P6cs,1995)
Cololejeuneadecemplicata(Steph.)Tixier (Grolle, 1995)
Cololejeunea deroinii Tixter (Tixier, 1993; Grolle, 1995) End, Type - Di6goSuarez (Antsiranana),Massif du Marojejy, rive gauche de la rivibre
Manantenina,alt. 350m, Deroin & Badrl5, holotypePC.
CololejeuneadiplasiolejeuneoidesTixier (Tixier, 1985;Grolle, 1995)End, Type Tamatave,P6rinet,forOtd'Antaniditra,alt. 950m,61711977,
Tixier 10637,
holotypePC.
Cololejeuneadisciflora Tixier (Grolle, 1995)End, Type - Antananarivo, Carion,
Angavobe,alt. 1600m, Tixier10834,holotypePC.
Cololejeunea distalopapillata (8.W. Jones) R.M. Schust.(Tixier, 1985; Grolle,
1995;P6cs& Geissler,2002)
Cololejeunea dolichodonta Tixier (Tixier, 1985; Grolle, 1995) End, Type
Fianarantsoa,Ambatofinandrahana,
alt. 1650m, 211311978,
Tixier 11394,
holotypePC.
CololejeuneaduvignaudiiE.W. Jones(Tixier, 1985;Grolle, 1995;P6cs& Geissler,
2002)
Cololejeuneaduvignqudii var.papillqta Tixier (Grolle, 1995)
CololejeuneaelegansSteph.(Tixier, 1985;Grolle, 1995;P6cs& Geissler,2002)
Cololejeuneafalcatq var. madecassaTixier (Grolle, 1995)End, Type - Tamatave,
P6rinet,aIt.9I7 m, Tixier10160,holotypePC.
Cololejeuneafloccosa(Lehm. et Lindenb.) Schiffn.(Grolle, 1995)
Cololejeuneafloccos& var. trivittata Tixier (Grolle, 1995)
Cololejeuneafurcilobulata(Berrie etE.W. Jones)R.M. Schust.(Grolle, 1995)
Cololejeuneahildebrandii (Austin) Steph.(Tixier, 1985;Grolle, 1995)
Cololejeunealeloutrei (E.W. Jones)R.M. Schust.(Grolle, 1995;P6cs& Geissler,
2002)
Cololejeunealeloutrei var. microlobulata Tixier (Tixier, 1985;Grolle, 1995)End,
Type - Tananarive,Ambatolampy,stationde Manjakatompo,alt. 1700m,
2711,011976,
Tixier 9385,holotypePC.
Cololejeuneuleloutrei var. ugulurica P6cs exTixier (Grolle, 1995)
Cololejeunealemuriana Tixier (Tixier, 1985;Grolle, 1995)
Cololejeunea marginata (Lehm. er Lindenb.) Schiffn. (Grolle, 1995; P6cs &
Geissler.2002)
Cololejeuneamicroscopica(Taylor) Schiffn var. exigua (A.Evans) Bernecker e/
Pocs(Grolle, 1995;P6cs& Bernecker,2009)
Cololejeuneaminutissima (Srn.)Schiffn.(Grolle, 1995)
CololejeuneamoramangaeTixier (Grolle. 1995:P6cs& Bernecker,2009)
Cololejeuneamocambiquensl's
S.W.Arnell(Grolle, 1995)
Cololejeuneaobliqza (Nees e/ Mont.) Schiffn. (Grolle, 1995;P6cs & Geissler,
2002)
Cololejeuneaobtusifulia (E.W. Jones) Tixier (Grolle, 1995;P6cs & Geissler,
2oo2)
Cololejeuneaperaffinis var. setulatu Benedix (P6cs& Geissler,2002)
Cololejeuneaplagiochiliana Tixier (Grolle, 1995) End, Type - Pdrinet, Benoist
s.n.,holotypePC.
Cololejeuneaplatyneura(Spruce)A. Evans(Grolle, L995;P6cs& Geissler,2002)
Cololejeunea pr&eruptorum Tixier (Tixier, 1985; Grolle, 1995) End, Type Mandialaza, for€,tde I'Angavo, 19I l2l I975, Tixier 7670, holotypePC.
Checklist of the bryophytes of Madagascar
2r3
Cololejeuneapteroporum Tixier (Tixier, 1985;Grolle, 1995)End, Type - Di6goTixier 7463,holotype
alt. 900m, 0611111975,
Suarez,stationde roussettes,
PC.
Cololejeunearaduliloba Steph.(Tixier, 1985;Grolle, 1995)
Cololejeunea saltuum Tixier (Tixier, 1985; Grolle, 1995) End, Type
Tixier
Fianarantsoa,
Ranomafana,forOtdu pk 24, alt.1200m,3011211976,
10038,holotypePC.
Cololejeunea sambiroana Tixier (Tixier, 1985; Grolle, 1995) End, Type
Tixier 10744,
Sambirano, forOt d'Anjozobe, alt. 300m, 2210811977,
holotypePC.
CololejeuneaselangorensisTixier (Tixier, 1985;Grolle, 1995)
Cololejeunea subinflata Tixier (Tixier, 1985; Grolle, 1995) End, Type
Tixier 8158,
Antananarivo,Manjakandriana,lac Mantasoa,L91041I979,
holotypePC.
Cololejeunea succineaTixier (Tixier, L985;Tixier, I979a) End, Type - Di6goSuarez,Montagned'Ambre, station des Roussettes,forOt de montagne,
Tixier 7395,holotype PC.
0311,111975,
Cololejeunea tamataven^sisTixier (Tixier, L985; Grolle, 1995) End, Type
Tixier 7230,
Foulpointe, for€t d'Analalava, alt. 100 m, 0511211975,
holotype PC.
CololejeuneatqnzaniqeP6cs(Grolle, 1995;P6cs& Geissler,2002)
Cololejeuneavidaliana Tixier (Grolle, 1995)
Cololejeuneazenkeri(Steph.)E.W.Jones(P6cs& Geissler,2002)
Colura benoistii Jovet-Ast (Grolle, 1995)
Colura bicornis Jovet-Ast (Grolle, 1995;P6cs, 1995) End, Typ" - Route de
Moramangad Anosibe,pk 29, for€t mar6cageuse,
sur feuille de palmier,
Millot & Bosser6319,herb (N.S.)
1.510911953,
Colura digitalis (Mitt.) Steph.(P6cs,1995;P6cs& Geissler,2002)
Colura heimii Jovet-Ast(Grolle, 1995)
Colura humbertiiJovet-Ast(Grolle, 1995)
Colura inornata Jovet-Ast (Grolle, 1995: P6cs, 1995) End, Type - Route de
sur feuille de palmier,
Moramangad Anosibe, pk 29, forOt mardcageuse,
L510911953,
Millot & Bosser6323,herb (N.S.)
Colura obesaJovet-Ast(Grolle, 1995)
Colura obvolutaJovet-Ast(Jovet-Ast,1983;Grolle, 1995)
Colura tenuicornl's(A. Evans)Steph.(Grolle, 1995)
Diplasiolejeunea andringitrae P6cs et Schafer-Verwimp (P6cs & SchaeferVerwimp, 2006) End, Type - Monts d'Andringitra, Nature Reserve,
montane rainforest, West side of Korokoro River, around Camp II,
P6cs et al., 9472/AM,
alt. 750-1000m, S 22'13' E 47"1.-2',20-231911994,
holotypeEGR.
DiplasiolejeuneaaulaeE.W. Jones(Grolle, 1995)
Diplasiolejeuneaauriculata Tixier (Grolle, 1995)End, Type - Fianarantsoa,for6t
de Tsarafidy,Tixier 8908,holotypePC.
Diplasiolejeuneacavifolia Steph.(Grolle, 1995;P6cs& Geissler,2002)
Diplasiolejeunea cobrensisGottsche ex Steph. (Grolle, 1.995;P6cs & Geissler,
2002\
DiplasiolejeuneacobrensisP6cssubsp.antsirananae(P5cs,2001;P6cs& Geissler,
2002) End, Type
Province d'Antsiranana, R6serve Spdciale de
2t4
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',311999,Geissler19549/8,,holotype G.
D ip I asio I ej eunea comorensis Tixier ( Grolle, I99 5)
Diplasiolejeunea cornuta Steph. (Grolle, 1995; P6cs, 1995)
Antananarivo,
Diplasiolejeunea ensifera Tixier (Grolle, 1995) End, Type
Andasibe, alt. 1400 m, Tixier 9626, holotype PC.
Fianarantsoa,
Diplasiolejeunea hamqta Tixier (Grolle, 1995) End, Type
Vohiparara, Tixier 11373,holotype PC.
Diplasiolejeunea insignis Tixier (Grolle, 1995) End, Type - Tananarive, Carion,
Angavokely, alt. 1800 m,2710111977,Tixier 10258,holotype PC.
Tamatave,
Diplasiolejeunea jonesii Tixier (Grolle, 1995) End, Type
Antaniditra, Vohiparara, alt. 1100 m,3011211976,Tixier 10157, holotype
PC.
Diplasiolejeunea kraussiana (Lindenb.) Steph. (Grolle, 1995)
Diplasiolejeunea lemuriana Tixier (Grolle, 1995) End, Type - Fianarantsoa,
Ranomafana, alt. 7200 m, 30lI2lI97 6, Tixier 10171,holotype PC.
Diplasiolej eunea magnistipulata Tixier (Tixier, 1985; Grolle, 1995)
Diplasiolejeunea ornatu P6cs er Schiifer-Verwimp (P6cs & Schaefer-Verwimp,
2006) End, Type - Province deToamasina, Mananara - Nord Biosphere
Reserve, Lowland rain forest on the NW slope of Behafotra Hill, alt.250Pbcs & Szab6 9877/CH, holotype
300 m, S 16"27.2'E 49"47.7',1610811998,
EGR.
Diplasiolejeunea palustriuraz Tixier (Grolle, 1995) End, Type - Tamatave, P6rinet,
Antanidi tr a, aIt. 1000 m, 06I 07I I97 7, Tixier I 0636, holotype PC.
Diplasiolejeunea phyllarthonii Tixier (Grolle, 1995) End, Type - Tamatave,
Manohitaly (route du Chrome), alt. 1200 m, 03/0311978,Tixier 11632,
holotype PC.
Diplasiolejeunea ramicola Tixier (Grolle, 1995) End, Type - Antananarivo,
Ambatoloana, 1400 m, Tixier 9629, holotype PC.
Diplasiolejeunea ranomafanae P6cs & Schiifer-Verwimp (P6cs & 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.L0'E 47"24.44',3010712004,P6cs
04I 28/AK, holotype EGR.
Diplasiolej eunea runssoren^sisSteph. (Grolle. 1995)
D ip I asio lej eunea subcornuta Trxter (Grolle, 1995)
Diplasiolejeunea symoensii Vanden Berghen (Grolle, 1995)
Diplasiolej eunea utriculuta Steph. (Grolle, 1995)
Diplasiolejeunea villaumei Steph. (Grolle, 1995)
Diplasiolejeunea zakiae Tixier (Tixier, 1977; P6cs & Geissler,2002)
Drepanolejeunea cambouena Steph. (Grolle, 1995; P6cs, 201Ib)
Drepanolejeunea cultrella (Mitt.) Steph. (P6cs & Geissler,2002)
Drepanolejeunea geisslerae P6cs (P6cs, 200I; P6cs & Geissler,,2002) Bnd, Type Antsiranana, R6serve Sp6ciale de Manongarivo, NW Madagascar,
abundant on palm stem, alt. 950-1050 m, N 13'59'E 48"26',031L999,
Geissler 19559,holotype G.
Drepanolejeunea madagascariensis (Steph.) Grolle (Grolle, 2001; P6cs &
Geissler,2002)
Checklist of the bryophytes of Madagascar
Drepanolejeunea mqscarend (S.W. Arnell) R.L. Zhu er Grolle (Zht
21.5
& Grolle,
2oo3)
5; P6cs,20Ilb)
D repanoI ejeunea p entadactyIa (Mont.) Steph. (Mizutani, 1.97
Drepanolejeuneaphysaefulia (Gottsche)Steph.(Grolle, 1995;P6cs,1,995;P6cs&
Geissler,2002)
Drepanolejeuneapocsii Grolle (P6cs& Geissler,2002)
Drepanolejeuneatrematodes(Nees)Bischl. (Grolle, 1995;P6cs& Geissler,2002)
Frullanoides tristis(Steph.)Van Slageren(Grolle, L995)
Haplolejeuneqsticta Grolle (Grolle, 1995;P6cs& Geissler,2002)
Lejeuneaulata Gottsche(Grolle, 1995;P6cs& Geissler,2002)
End, Type P6cs& Geissler,2002)
Lejeuneaalatavar. patriciaeP6cs(P6cs,2001.;
Provinced'Antsiranana,RdserveSpdcialede Manongarivo,on the rain
forest covered north ridge leading to the 1,869m high summit of
Geissler
Manongarivomassif,epiphylous,N L3'59'E 48"26',1210311,999,
19856/1,holotype G.
LejeuneaamaniensltE.W. Jones(Grolle, 1995)
P6cs,20I7b)
LejeuneaanisophyllqMont. (Grolle, 1985;P6cs& Geissler,2002;
LejeuneacapensisGottsche(Grolle, 1995)
LejeuneaconfusaE.W. Jones(P6cs& Geissler,2}12)
Lejeuneaecarinatq(Steph.)Steph.(Grolle, 1995)
LejeuneaecklonianaLindenb.(Grolle,1995;P6cs& Geissler,2002)
Lejeuneaflava(S*.) Nees(Grolle, 1995)
Lej euneaflav ovirensAongstr. (Groll e, 1995;P6cs& Geissler, 2002)
Lejeuneaisophylla E.W. Jones(Grolle, 1995)
Lejeuneajulacea Steph. (Grolle, 1995;P6cs,2011b) End, Type - Forsyth-Major
B 473,holotypeG.
LejeunealeucosisBesch.& Spruce(Grolle, 1995;P6cs,20I0a)
Lejeunea masoqlae P6cs (P6cs, 2010a) End, Type - West side of Masoala
Peninsula,aboveAntongil Bay, montanerainforeston the summit ridge
E 50"05" 0910911994,
SE of Ambanizanavillage,alt. 660-720m, s L5"37',
P6cs9448/AN,holotype EGR, isotypeMO, TAN.
LejeuneaobtusatuGottsche(Grolle, 1995)
Lejeuneapapilionacea Steph.(Grolle, 1995;Zhu & Grolle, 2001)
Lejeuneaphyllobola Neese/ Mont. (P6cs,1995)
Lejeuneatubularis (Spreng.)Gottscheet al. (Grolle, 1995;P6cs& Geissler,2002)
LejeuneatuberculosaSteph.(Grolle, 1995;P6cs& Geissler,2002)
Lejeuneavillaumei (Steph.)Grolle (Grolle, 1995;P6cs& Geissler.2002)
Lejeunea vojtkoi P6cs (P6cs,2010a)End, Type - 10 km North of Fianarantsoa,
m, S 2L'15'B 47"14',
on decayingwood in montaneforest, alt. 1230-1360
2210911,994,
Vojtk6 9461/AY,holorypeEGR.
Lejeuneavulgarifurmis Gottsche(Grolle, 1995;P6cs,2010a)
Lepidolejeunea bidentula (Steph.) R.M. Schust. var. bidentula (Piippo, 1986;
Grolle,1995;Zhu& So,2001)
Leptolejeuneaepiphylla (Mitt.) Steph.(P6cs& Geissler,2002)
*Leptolejeuneavitrea (Nees) Schiffn.- Anjozorobe, 6piphylle, alt. 300 m, 221071
1977,Tixier10743,TAN.
Lopholejeune&borbonicaSteph.(Grolle 1995,P6cs& Geissler,2002)
LopholejeuneserugataB. Thiers (Grolle, 1995)End, Type - 1896,Forsyth-Major,
holotypeG.
2 t6
L.Marline et al.
Lopholejeunea eulopha (Taylor) Schiffn. (Grolle, 1995; P6cs & Geissler, 2002;
Zhu & Gradstein,2005)
Lopholejeunea grandicrisla Steph. (Grolle, 1995; P6cs & Geissler,2002)
Lopholejeunea leioptera Gyarmati (Sass-Gyarmati,2001.;P6cs & Geissler,2002)
End, Type - Fianarantsoa, alt. 1590 ffi, S 22"11'00" E 46'53'00", 1.91091
1994, Magill & P6cs 12855, holotype MO.
Lopholejeunea lepidoscypha Kiaer et Pearson (Grolle, 1995) End, Type
Ankaratra,1877 , Borgen s.n., holotype BM.
Lopholejeunea multilacera Steph. (Grolle 1995)
Lopholejeanea nigricans (Lindenb.) Schiffn. (Grolle, 200I; Zhu & Gradstein,
200s)
Lopholejeuneq onraedtii Vanden Berghen (Grolle, 2001) End, Type - Route de
Moramanga,Tamatave, L7 km from Moramanga, forOt primaire humide
et d6grad€e, aIt.800 m, 0111970,Onraedt 70M2030, holotype BR.
Lopholejeunea subfusca (Nees) Schiffn. (Grolle, 1995; P6cs & Geissler,2002;Zhu
& Gradstein,2005)
Lopholejeunea tixieriana Vanden Berghen (Grolle, 1995) End, Type
Antanidetra, 0711977,Tixier 10720, holotype BR.
Lopholejeunea utriculata Steph. (Grolle, 1995) End, Type - unknown.
Marchesinia excavata (Mitt ) Schiffn. (P6cs & 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)
Microlejeuneafissistipula Steph. (Grolle, 1995; P6cs & Geissler,2002) End, Type
- unknown.
Microlejeunea inflata Steph. (Grolle, 1995; P6cs & Geissler,2002)
Microlejeunea oblongistipula (Gottsche) Pearson (Grolle, 1995)
Microlejeunea papulosa (Gottsche) Pearson (Grolle, 1995) Bnd, Type - unknown.
Odontolejeunea lunulata (F. Weber) Schiffn. (Grolle, 1995; P6cs, 1.995;P6cs &
Geissler,2002)
Otolejeunea moniliuta Grolle (Grolle, 1985; Grolle, 1995) End, Type - Partie
nord du massif du Morojejy, alt. 1450 m, Guillaumet 4133/7,holotype JE,
isotypes PC, BR.
Otolejeuneq rabenorii Tixier (Grolle, 1995) End, Type - Toamasina, route de
Lakato, dpiphylle en forOt dense d gramin€,es,0210611979,
Tixier 12591,
holotype PC.
Otolejeunea subana P6cs (P6cs, 2004) End, Type - Toamasina Province, Mantadia
Forest Reserve, North of Andasibe (P6rinet), submontane rainforest, alt.
1030 m, S 18"52'20" E 48"28'26", 0211.011994,
P6cs & Szab6 9485/R,
holotype EGR.
Prionolejeunea grato (Gottsche) Schiffn. (Grolle, 1995; P6cs & Geissler,2002)
Ptychanthus africanus Steph. (Grolle, L995;Ahonen et a\.,2005)
Pycnolejeunea contigua (Nees) Grolle (Grolle, 1,995:He, 1999)
Schiffneriolejeunea madagascariensis (Steph.) Gradst. (Grolle, 1995) End, Type unknown.
Schiffneriolejeunea pappeana (Nees) Gradst. (Grolle, 1995; P6cs & Geissler,
2002\
Checklist of the bryophytes of Madagascar
2I 7
Schiffneriolejeunea pappeana var. bidentata Gradst. etYanden Berghen (Grolle,
lees)
Schiffneriolejeunea pappeana var. integra Gradst. et Yanden Berghen (Grolle,
\.
lees)
Schiffneriolejeuneaparviloba (Steph.)Gradst. (P6cs& Geissler,2002)
Schiffneriolejeuneapolycarpa (Nees) Gradst. (Grolle, 1995; P6cs & Geissler
2002)
Symbiezidium barbiflorum (Lindenb. et Gottsche) A. Evans (Sass-Gyarmati&
P6cs,2002)
Symbiezidium madagascarienseSteph. (Gradstein & Van Beek, 1985; Grolle,
1995;P6cs,1995)
Taxilejeunea conformis (Mont. er Nees) Steph. (Grolle, 1995;P6cs & Geissler
2oo2)
Taxi lejeuneafurcicornuta Grolle ( Grolle, 1995)
Taxilejeuneasikorqe Steph.(Grolle, 1995)End, Type - unknown.
Thysananthus spathulistipus (Reinw. et al.) Lindenb. (Grolle, 1995; P6cs &
Ah-Peng
Geissler,2002)- Marojejy NationalPark, alt. 650m,2411112009,
349, det.Ah-Peng,Bardat.
Xylolejeuneagrolleana (P6cs)X.-L.He et GroIIe (P6cs,1999;He & Grolle, 200I)
Lepidoziaceae
Amazoopsisdiplopoda (Pocs)J.J.Engele/ G.L.S.Merr. (Grolle, 1995;P6cs,1995)
Bazzania curvidensSteph.(Grolle, 1995)End, Type - Alakaty forest, 3011111,889,
Perrot,herb (N.S.).
Bazzania deciduifulia Onr. (Onraedt, 1977;Grolle, 1995)End, Type - Province
de Fianarantsoa,forOt d'Ambatofitorahana,Onraedt 7I M 5364, holotype
NAM, isotypesBR, JE.
(Lehm. etLindenb.) Trevis. (Grolle, 1995)
Bazzania decrescens
Bazzania decrescens
subsp.pumila (Mitt.) P6cs(P6cs1995)
var. ambahatraeP6cs (P6cs& Geissler,2002)End, Type Bazzania decrescen^s
Manongarivo massif, alt. 1300-1400m, P6cs & GeisslerPG 19742
holotypeEGR.
Bazzaniam&scarena(Steph.)Herzog(Grolle, 1995)
Bazzania nitida (F. Weber) Grolle (Grolle, 1995;P6cs, 1995;P6cs & Geissler
2002)
Bazzania orbanii P6cs (P6cs, 2010b) End, Type - SW Madagascar,Toliara
Province,IsaloNationalPark, "Piscinenaturelle",a deeppool of a brook
in a sandstonegorge, shaded by Pandanuspulcher trees, on soil,
alt. 800m, Orbdn 9455/J,holotype EGR, isotypesMO, TAN.
Bazzaniapemotana(Steph.)E.W.Jones(Jones,1975;Grolle, 1995)
Bazzaniapraerupta (Reinw. et al.) Trevis. (Grolle, 1995;P6cs,1995)
Bazzaniu reflexa (Gottsche)Steph.(Grolle, 1995)End, Type - unknown.
Kurzia capillaries (Sw.) Grolle subsp. capillaris (P6cs, 1995;P6cs & Geissler
2002)
Kurziq capillaris (Renauld ex Steph.)P6cssubsp.stephanii (Grolle, 1995;P6cs,
1995;P6cs& Geissler,2002)
Lepidozia africana Steph.(Grolle, 1995)
Lepidozia succidaMitt. (Grolle, 1,995;P6cs,1995)
Telaranea bischleriana P 6cs(P6cs,2006)
21,8
L.MarIine et al.
Telaraneacoactilis (Spruce)J.J.Engel er G.L.S.Merr. (Engel & MerrilI,2004)
Telaraneanematodes(Gottscheex Austin) M. Howe (Grolle ,1995)
r\
Lophocoleaceae
Chiloscyp-huscoadunatus(S*.) J.J. Engel er R.M. Schust.(Grolle, 1,995;P5cs &
Geissler,2002)
Chiloscyphusconcretus(Mont.) J.J. Engel e/ R.M. Schust.(Grolle, 1995;P6cs &
Geissler,2002)
Chiloscyphusdifformis (Nees)J.J.Engel e/ R.M. Schust.(P6cs& Geissler,,2002)
Chiloscyphuslucidus (Spreng.ex Lehm.) J.J.Engel e/ R.M. Schust.(Grolle, 1995;
P6cs& Geissler,2}l2)
(Gottsche) J.J. Fngel e/ R.M. Schust. (Grolle,
Chiloscyphus madagascariensr's
1995)End, Type - unknown.
Chiloscyphusmartianus (Nees) J.J. Engel er R.M. Schust.(Grolle, 1995;P6cs &
Geissler,2002)
Chiloscyphusmuricatus (Lehm.) J.J. Engel e/ R.M. Schust.(Grolle, 1995:P6cs&
Geissler,2002)
ChiloscyphusregularisSteph.(Grolle, 1995)
Clasmatocoleavermicularls(Lehm.) Grolle (Engel, 1980)
Conoscyphustrapezioides(SandeLac.) Schiffn. (Grolle, 1995;P6cs,1995;P6cs&
Geissler,2002)
Heteroscyphusdubius (Gottsche)Schiffn.(Grolle, 1995)
Heteroscyphussplendens(Lehm. etLindenb.) Grolle (P6cs,1995;P6cs& Geissler,
2002)
Leptoscyphusexpanszs(Lehm.) Grolle (Grolle, 1995)
Leptoscyphusinfuscatus(Mitt.) E.W. Jones(P6cs& Geissler,2002)
Marchantiaceae
Marchantia dehilis Goebel (Grolle, 1995)
M archantia pappeanaLehm. subsp.p appeana ( Grolle, I995)
Mastigophoraceae
Mastigophora diclados (Brid.';;exF. Weber) Nees (Grolle, 1995;P6cs& Geissler,
,:
2002)
Metzgeriaceae
Metzgeriacons&nguineqSchiffn.(So,2004)
Metzgeriafurcata (L.) Dumort. (So, 2004)
MetzgerialeptoneuraSpruce(Grolle,1995;So,2004)
MetzgeriamadagassaSteph.(Grolle,1995;So,2004)
Metzgerianudifrons Steph.(So,2004)
Metzgeria quadrifaria Steph.(So, 2004)
Pallaviciniaceae
*Psllavicinia lyellii (Hook.) - Ambavaniasy,r6servede Vohimana,forOtombrophile
d'altitude, jonction sol-racine au pied d'une micro falaise, alt. 870m,
E 48'30'05.7",1,410412010,
Reeb,Andriamananatena
S L8'55'19.5"
& Bidault
CRAE 40 & CRAE 48, TAN, det. Reeb & Andriamananatena;Ambavaniasy,r6servede Vohimana,bord de cheminvers relaisdu naturaliste,terre
tass6e,alt.790 ffi, S 18"55'14.95"
E 48"30'49.58",1610412010
Reeb,Andriamananatena
& Bidault CRAE 155,TAN, det Reeb& Andriamananatena.
Checklist of the bryophytes of Madagascar
2t9
Symphyogyna brasiliensis Nees e/ Mont. (Grolle, 1995)
*Symphyogyna aff. podophylla (Thunb.) Mont. e/ Nees. - Ambavaniasy, rdserve
de Vohimana, grande cascade microfalaise d 3 m du bord, face verticale,
Reeb, AndriamananaIt.849 m, S 18"55'43.6" E 48"29'56.6",1,510412010,
atena & Bidault CRAE 87. TAN. det. Reeb & Andriamananatena.
Symphyogynopsis gottscheanc (Mont. er Nees) Grolle (Grolle, 1995)
Plagiochilaceae
Pl agi ochil a angusta Lindenb. (Grolle, 1995; P6cs & Geissler, 2002)
Plagiochila barteri Mitt. var. barterf (Grolle , 1995; P6cs & Geissler ,2002)
Plagiochila boivinfi Steph. (Grolle, 1995)
Plagiochila boryana Gottsche ex Steph. (Grolle, 1995)
Plagiochila cambuena Steph. (Grolle, L995)
Plagiochila drepanophylla Sande Lac. (Grolle, 1995)
Plagiochilafracta P6cs,nom. inval. (P6cs & Geissler,2002; Heinrichs et a|2005)
Plagiochila granditexta Steph. (Grolle, 1995) End, Type - Hildebrandt, herb
(N.s.).
Plagiochila incerta Gottsche (Grolle, 1995; P6cs & Geissler,2002)
Plagiochila integenima Steph. (Grolle, 1995)
Plagiochila kiaeri Gottsche (Grolle, 1995; P6cs & Geissler,2002)
Plagiochila paucidentats Mont. e/ Gottsche (Grolle,1995; P6cs & Geissler,2002)
Plagiochila pectinataWilld. exLindenb. (Grolle,,1.995;P6cs & Geissler,2002)
Plagiochila punctata (Taylor) Taylor (Grolle, L995; P6cs &. Geissler, 2002;
Heinrichs et a1.,2005)
Plagiochila repanda (Schwaegr.) Lindenb. (Grolle, 1995)
Plagiochila repanda var. perrotana (Steph.) Vanden Berghen (Grolle, 1995)
Plagiochil a rodriguezii Steph. (Grolle, I99 5)
Plagiochila squamulosa var. crispulo-caudata (Gottsche) Vanden Berghen
(Grolle, 1995)
Plagiochila squamulosavar. sinuosa (Mitt.) Vanden Berghen (Grolle, 1995)
Plagiochila stricta Lindenb. (Lindner et a1.,2004)
Plagiochila terebrans Nees e/ Mont. exLindenb. (Grolle,1995; P6cs & Geissler,
2002)
Pleuroziaceae
Pleurozia gigantea (F. Weber) Lindb. (P6cs & Geissler.2002)
Porellaceae
Porella abyssinica var. hoehnelii (Steph.) P6cs (Grolle, 1995)
Porells cucullistipula Sreph. (Grolle, 1995)
Porella madagascariensis (Nees er Mont.) Trevis. (Grolle, 1995)
Porella montqntid (Steph.) E.W. Jones (Grolle, 1995)
Porella triquetra (Steph.) E.W. Jones (Grolle, 1995)
Radulaceae
Radula ankdinensis Gottsche ex Steph. (Grolle, 1995; Grolle, 200I; P6cs &
Geissler,2002)
Radulq appressa Mitt. (Grolle, 1995;P6cs, 1995;P6cs & Geissler,2002)
Raduls boryana (F. Weber) Mont. (Grolle, 1995: Grolle, 200I; P6cs & Geissler,
2002)
Radula comorensis Steph. (P6cs & Geissler,2002)
L.Marline et al.
\
Radulafulvifulia (Hook.f. etTaylor) Gottscheet al. (Grolle,200I)
Radula madagascariensisGottsche (Yamada, 1975; Grolle, 1995; P6cs, 1995;
Grolle, 2001)
Radula marojezicaE.W. Jones(Grolle, 1995)End, Type - ProvinceAntsiranana,
Parc National de Marojejy, P6cs 90114/BN,holotype E, isotype and
paratypeEGR.
Radula quadrata Gottsche(Grolle, 1995)
Radula silvestrisGottsche(Grolle, 1995)
Radula stenocalyxMont. (Grolle, 1995;P6cs,1995;Grolle, 2001;P6cs& Geissler,
2002)
Rsdula voluta Taylor er Gottsche et al. (Grolle, 1995;P6cs, 1995;Grolle, 200I;
P6cs& Geissler,2}D2)
Ricciaceae
Riccia atromarginata var.jovet-astiae Rauh & Buchloh (Grolle, L995)
Riccia caver.tosaHoffm. (Grolle, 1995)
Riccia uinita Taylor (Grolle, 1995)
Riccia percsonii SultanKhan (Perold,1995)
*Riccia purpurascensLehm. - Makirovana, Talus vertical en bord de rizidre,
Ambodimanga au pied de Makirovana, alt. 4l m, S 14"II'13.6 8
49"59'09.2",0310512010,
ReebCR 203,det. Reeb & Bardat.
Scapaniaceae
Anastrophyllum piligerum (Nees) Steph. (Grolle, 1995; P6cs, L995; P6cs &
Geissler,2002)
Andrewsianthusaberrans(Neeser Mont.) Grolle (P6cs,1995)
Plicanthus birmensis(Steph.)R.M. Schust.(Grolle, 1995)
Plicanthus hirtellus (F. Weber) R.M. Schust.(Grolle, 1995;P6cs& Geissler,,2002)
Schistochilaceae
Gottscheaneesii (Mont.) R.M. Schust.(Grolle, 1995;P6cs,1995;P6cs& Geissler,
2002)
Schistochila alata (Lehm.) Schiffn. (Grolle, 1995;P6cs, 1995;P6cs & Geissler,
2002)
Schistochilapiligera Steph.(Grolle, 1995;P6cs,1995;P6cs& Geissler,2002)End,
Type - Cambou1,herb(N.S.).
Targioniaceae
Targionia hypophylla L. (Grolle, 1995)
*Targionia dioica Schiffn. - Biosphere Reserveand National Park, MananaraNord, Toamasina, alt.220m, 1710811998,
P6cs& Szab69878/CY,EGR.
Hornworts reported from Madagascar
Anthocerotaceae
Anthoceros sp. (Grolle, L995)
*Dendroceros cf. africanzs Steph. - R6serve Sp6ciale Angavokely, alt. 1750 m,
II12009. Bardat JB MAD 147. det. Bardat.
Notothyladaceae
Phaeoceros carolinianzs (Michx.) Prosk. (Grolle, 1995)
Checklist of the bryophytes of Madagascar
Mosses reported
22r
from Madagascar
Amblystegiaceae
Amblystegium chalaropelma Mtill.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. er W.Guembel (Crosby et al.,
1e83)
Hygrohypnum caussequei(Renauld & Cardot) Cardot (Crosby et a1.,1983) End,
Type - Entre Tananarive et Betafo, 1894, Causs)que, holotype PC.
Andreaeaceae
Andreaeq subulata Harv. (Crosby et a1.,1983)
Andreaea tsarstananae Th6r. (Crosby et al., 1983)
Anomodontaceae
Herpetineuron toccoqe (Sull. e/ Lesq.) Cardot (Touw, 1993)
Archidiaceae
*Archidium ohioense Schimp. ex Miill.Hal. - Beza mahafaly, Betioky, Riverine
forest adjacent to Sakamena river, open ground on forest floor, alt. 1-30m,
S 23"39' B 44"38',18/05/1987,Phillipson 1818,TAN.
*Archidium subulatum MiJILHal. - Beloha Tsihombe, dalle de gneiss, L01041I971.,
Cornet & Cremers 2014, TAN.
Bartramiaceae
Bartramia gigantea Bory (Crosby et a1.,1983)
Bartramia longifulia Hook. (Crosby et a1.,1983)
Breutelia dffiacta Mitt. (Crosby et a1.,1983)
Breutelia gnaphalea P. Beauv. (Crosby et a1.,1983)
Breutelia madagassq Th6r. var. madagassa (Crosby et al., 1983) End, Type Mont Tsaratanana, alt. 1200 m,0411924,Perrier de la Bathie, holotype PC.
Breutelia miuodonra (Mitt.) Broth. (Crosby et a1.,1983;Virtanen,1997)
Breutelia penieri Th6r. (Crosby et a1.,1983)
Breutelia sciuroides Th6r. (Crosby et a1.,1983)
Breutelia stuhlmanniiBroth. (D" Sloover, 1975;Crosby et a1.,1983)
Breuteliq viguieri Cardot (Crosby et al., 1983) End, Type - Ambatolampy,
ruisselet prds du sommet du Tsiafajavona, alt.2575 m,2S|II|I9I2,Viguier
1688ter, holotype PC.
Philonotis byssifurmr'sMiill.Hal. (Crosby et a1.,1983)End, Type -Zone inf6rieure
des forOts,Nosy Be, Hildebrandt, holotype PC.
Philonotis hastuta (Duby) Wijk etMargad. (Crosby et a1.,1983)
Philonotis mauritiana Aongstr. (Crosby et a1.,1983)
Philonotis mauritiana var. stricla Renauld et Cardot (Crosby et al., 1983) End,
Plateau central, environs de Tananarive, Cambou6; Entre
Type
Tananarive et Betafo, CaussAque. 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 Th6r. (Crosby et a1.,1983)
Philonotis sparsifulia (Hampe) A.Jaeger (Crosby et a1.,1983)
L.Marline et al.
Brachytheciaceae
Aerolindigia capillacea (Hornsch.) M. Menzel (Crosby et a1.,1983)
Brachythecium borgenii (Hampe) A. Jaeger (Crosby et a1.,1983)
Brachythecium decurrensCardot (Crosby et a1.,1983)
Eurhynchium spinulinerve Kiaer ex Cardot (Crosby et a1.,1983)
Eurhynchium striatum (Schreb. ex Hedw.) Schimp. (Crosby et a1.,1983)
Helicodontium fabroniopsr's Mi.ill. Hal. (Crosby et al., 1983) End, Type - Zone
sup6rieure des forOts, troncs d'arbres d Andrangoloaka, Sikora, herb
(N.s.).
Palamocludium
r9e7)
ldskeoides (Hook.) E. Britton (Crosby et al., 1983; Hofmann,
Rhynchostegiella miuotheca (Renauld et Cardot) Broth. (Crosby et a1.,1983)
Rhynchostegium angustifolium Renauld & Cardot (Crosby et a1.,1983)End, Type
Zone moyenne des forOts, forOt d'Analamazaotra, Cambou1 &
Campenon,herb (N.S.).
Rhynchostegium comorae (Mi.ill.Hal.) A.Jaeger (Crosby et a1.,1983)
Rhynchostegium distans Besch. (Crosby et a1.,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 Car dot (Crosby et a1.,,1983)
Schimperella rhynchostegioides Thdr. (Crosby et al., 1983) End, Type - Mont
Tsaratanana, alt. 1200 m,0411924,,Perrier de la Bathie 176 p.p., lectotype
PC.
Squamidium brqsiliense (Hornsch.) Broth. (Crosby et a1.,1983; Allen & Crosby,
1986;Delgadillo, 1995)
Bruchiaceae
Trematodon curvicollis Cardot (Crosby et al., 1983) End, Type
Entre
Savondronina et Ranomaf ana, B esso n, holotype PC.
Trematodon decaryi Th6r. (Crosby et a1.,1983)End, Type - Vallde de I'Anony, sur
talus humide, 0510811921,
Decary, holotype PC.
Trematodon lacunosus Renauld & Cardot (Crosby et a1.,1983)End, Type - Entre
Savondronina et Ranomaf ana,\892, Besson,PC.
Trematodon longicol/is Michx. (Crosby et a1.,1983)
Tremqtodon platybasis Miill.Hal. (Crosby et al., 1983) End, Type - Alasora,
Ambohipo, Tananarive, alt. 1250m, Cambou1,holotype PC.
Trematodon reticulalus Mtill.Hal. ex Geh. (Crosby et a1.,1983) End, Type - Zone
moyenne des forOts,d I'Est d'Ambatondrazaka, Rutemberg,herb (N.S.).
Trematodon viguieri Cardot (Crosby et a1.,1983)
Bryaceae
Anomobryum filifurme var. madagassum Renauld et Cardot (Crosby et a1.,1983)
End, Type -Zone de plateau central, Imerina, Cambou6, herb (N.S.).
Anomobryum julaceum (P. Gaertn., B. Mey. er Scherb.) Schimp. var. julaceum
(Ochi, 1972;Crosby et a1.,1983)
Brachymenium acuminatum Harv. (Ochi, 1972; Crosby et al., 1983; Delgadillo,
rees)
Brachymenium apiculatum P. Beauv. (Crosby et al., 1983)
Brachymenium capitulatun (Mitt.) Kindb. (Crosby et a1.,1983)
Checklist of the bryophytes of Madagascar
223
Brachymenium exile (Dozy e/ Molk.) Bosch e/ SandeLac. (Ochi, 1972;Crosby
Delgadillo,1995)
et a1.,1983;
Brachymeniumleptophyllum (Bruch et Schimp.ex Mtill.Hal.) Bruch & Schimp.ex
A. Jaeger(Crosbyet a1.,1983)
(Ochi, 1972;Crosbyet a1.,1983)End, Type BrachymeniummadagassumHampe
Plateaucentral, 1869,Borgen,holotype PC.
P6cs& Geissler,
Hook. (Ochi, 1972;Crosbyet a1.,1983;
Brachymeniumnepalense
2oo2)
Besch.(Ochi, 1972;Crosbyet a1.,1983)
Brachymeniumnigrescens
Brachymeniumnotqrisii (Mitt.) A.J. Shaw(Crosbyet a1.,1983)
BrachymeniumphilonotulaBroth. (Ochi, 1972;Crosbyet a1.,1983)
Tixier, 1995)
Brachymeniumpulchrum Hook. (Ochi, 1972;Crosbyet a1.,1983;
Brachymeniumrigidum Broth. etParrs(Crosbyet a|.,1983)
Bryum albo-marginatum Cardot (Ochi, 1972;Crosby et al., 1983) End, Type Mont Tsaratanana.alt. 1200-2400m, 411924,Perrier de Ia Bathie s. n.,
holotype PC#
Bryum alpinulum var. levieri Renauldet Cardot(Crosbyet a1.,1983)End, Type Zone inf6rieure desforOts,for0t de Fito, Perrot, holotype FI#
Bryum alpinum Huds. ex With. var. alpinum (Ochi, 1972;Ochi, 1,973;Crosby
e ta L . , 1 9 8 3 )
Bryum apiculaturr Schwaegr.(Ochi,1972;Crosbyet a1.,1983)
Bryum arachnoideumMiill.Hat. (Ochi, 1972;Crosbyet a1.,1983)
Bryum argenteumHedw. var. argenteum(Ochi, 1972;Crosbyet a1.,1983;P6cs&
Geissler,2002)
Brid. (Ochi, 1972;Crosbyet a1.,1983)
Bryum aubertii (Schwaegr.)
Bryum billardierei Schwaegr.var. billardierei (Ochi, 1972; Mohamed, 1979;
Crosbyet a1.,1983)
Bryum caespiticiumHedw.var. caespiticium(P6cs& Geissler,2002)
Bryum capillare Hedw. var. capillsre (Ochi, 1972;Crosbyet a1.,1983)
Bryum coronatunr Schwaegr.(Ochi, 1972;Crosby et a1.,,1983;P6cs & Geissler,
2002)
Bryum dichotomumHedw.(Crosbyet a1.,1983)
Crosbyet a1.,1983)
Brid. (Mohamed,1979;
Bryum erythrocaulon(Schwiiegr.)
Bryum huillenseWelw. etDuby (Ochi, 1972;Crosbyet al.,1983;P6cs& Geissler,
2oo2)
Bryum imerinqnusO'Shea(O'Shea,2008)
Bryum leptospeironMi.ill.Hal. (Crosbyet a1.,1983)
Crosbyet a1.,1983)
Bryum megalacrionSchwaegr.(Ocht,,1972;
var. wichurae(Broth.) Mohamed (P6cs& Geissler,2}02)
Bryum neelgheriense
Delgadillo,1995)
(Ochi,1972;Crosbyet a1.,1983;
BryumparadoxumSchwaegr.
centre,alt. 2000m,
Type
ForOt
du
End,
(Crosby
a1.,1983)
et
Bryum penieri Th6r.
Perrier de la Bathie12127p.p,holotype PC.
Bryum pseudotriquetrum (Hedw.) P. Gaertn., B. Mey. et Scherb. var.
pseudotriquetrum(Crosbyet al., 1983)
Bryum purpureonigrumDuby (Crosbyet a1.,1983)End, Type - unknown.
Bryum
savondroninicusO'Shea(Crosbyet a1.,1983;O'Shea,2008)End, Type BetweenSavondroninaand Ranomafana,Besson,herb(N.S.).
Hampe (Ochi, 1972;Crosbyet a1.,1983)
Bryum subargenteum
Bryum viguieri Th6r. (Ochi, 1972;Crosbyet a1.,1983)
224
L.Marline et al.
Bryum voeltzkowiiBroth. (Ochi, 1972;Mohamed,1979;Crosbyet a1.,1983)
Mniobryum perlaxum Thdr. et P. de la Varde (Crosby et a1.,1983)End, Type Totohaza,0511931,,
Grainville,holotypePC.
Rhodobryumcommerconii(Schwaegr.)
Brid. (Crosbyet a1.,1983;
P6cs& Geissler,
2002)
Rhodobryumgiganteum(Schwaegr.)
Paris(Crosbyet a1.,1983;Ochi, 1972)
Rhodobryumontariense(Kindb.) Paris(Crosbyet a1.,1983)
Calymperaceae
Calymperesafzelii Sw.(Crosbyet a1.,1983;
P6cs& Geissler,2}D2)
CalympereserosumMi.ill.Hal.(Crosbyet a1.,1983)
Calympereshispidun Renauldet Cardot(Crosbyet a1.,1983;
Ellis, 1988)
Calymperesloucoubense
Besch.(Crosbyet a1.,1983)End, Type - Zone inf6rieure
desforOts,Nosy Be, Lokobe,1879,Marie, holotypeBM.
Calymperespalisolii Schwaegr.(Crosbyet a1.,1983)
pallidum Mitt. (Crosbyet a1.,1983;
Calymperes
Delgadillo,1-995)
*Calymperes sqnctae-mariae var. mayottense Besch. - Mananjary, tronc de
Pandanus, 14107
11972,Cornet& Cremers2250, TAN.
(Sull.)Mitt. (Crosbyet a1.,1983;
Calymperestaiten^se
Orbdn, 1995)
CalymperestenerumMiill.Hal. (Crosbyet a1.,1983)
Calymperesvenezuelanum(Mitt.) Broth. exPittier (Orbrin, 2000)
Leucophaneshildebrandtii Miill.Hal. (Crosbyet a1.,1983;
SalazarAllen, 1993)
Leucophanesrenauldii Cardot (Crosbyet a1.,1983;SalazarAllen, 1,993;P6cs&
Geissler,2002)
Leucophanesrodriguezii Mtill.Hal. (SalazarAllen, L993)
Mitthyridium fasciculatum (Hook. et Grev.) H. Rob. var. fasciculatum (Reese,
reer)
Mitthyridium subflavum (Renauld& Cardot) H. Rob. (Mtiller, 1897;Crosbyet al.,
1e83)
Octoblepharumafricanum(Broth.) Cardot (Crosbyet a1.,1983)
Octoblepharumalbidum Hedw. (Crosbyet a1.,1983;
P6cs& Geissler,2002)
Syrrhopodonafricanus (Mitt.) Parisvar. africanus (Crosbyet a1.,1983)
Syrrhopodonafricanus var. mandrakensis(Tixier) W.D. Reese(Tixier, 1992;P6cs
& Geissler,Z}D?)
Syrrhopodon albidus (E.B. Bartram) L.T. Ellis subsp. integrifulizs (P6cs &
Geissler,2002)
Synhopodon apertifuIius Besch.(Ellis, 1995)
SyrrhopodonarmatissimusW.D. Reese(Reese,1991.;
Orb6n, 1995)
Syrrhopodon armatus (BizoI et Onr.) Orbr4nand W.D. Reese subsp. insularus
(Orbiin, 1977)
Syrrhopodon asperMitt. (Een, 1993)
SyrrhopodoncuneifuliusTh6r. (Orbiln, 1981;Crosbyet a1.,1983;P6cs& Geissler,
2002) End, Type - Massif de l'Ankaizina, alt. 1200m,,0411923,
Decary,
holotypePC.
Syrrhopodon dimorphophyI lus L.T . Ellis (Ellis, 2005)
Syrrhopodongardneri (Hook.) Schwaegr.(Orbr4n,1981;Crosbyet a1.,1983;Ellis,
2002;P6cs& Geissler,2}}2)
SyrrhopodongaudichaudiiMont. (Orbr4n,1981;Crosbyet a1.,1983;
Ellis, 2005)
Syrrhopodongraminifulius Renauldet Cardot(Orbdn,1981;Crosbyet a1.,1983)
Checklist of the bryophytes of Madagascar
225
et Catdot in Renauld (Orb6n,1978;Orbdn'
Syrrhopodonhispidocosta-tus^Renauld
2003;Ellis, 2005)
& Geissler,2002;Etlis,
1981;cior6iri a1.,1983;P6cs
a1.,1983)
et
SyrrhopodoninvolutusSchwaegr.(Orb6n,1981;Crosby
al',1983)
iyut ipodon mauritianusMiill.Hai.ex Aongstr.(Orb6n, L98L;Crosbyet
a\..1983)
et
(Crosby
iyohipodon parasiticus(Sw.ex Brid.) Besch.
iyohipodon pottioides Orbdn (Orbr{n,1995)
(orb6n, L981;
iyotipodon proltfer yy. acanthoneuros(Miill.Hal.) Mult.Hal'
2003)
Ellis,
CrosbYet a1.,1983;
et al., 1983;
syrrhopodon proLifrr lg!*aegr. var. proliftr (otb6n, 1981';Crosby
Delsadillo.1995)
SyrrhopodoiputcherW.b. Reese(Orb6n,1995)iyohipodon-spiralis Renau\det Cardot (Ellis, 2005)
iyrrlripodon tixieri W.D. Reese-(Tixier, I?97)^ Eld; I{p"^,- Ranomafana'
Fianarantsoa,route de Mananjary, pk 24, corticole, alt' 900 m, Tixier
MO'
PC, ParatYPe
10152,holotYPe
2005)
(Ellis'
Ellis
L.T.
Syrrhopodonvardet
Catagoniaceae
Catigonium nitens(Brid.) Cardotsubsp.nitens(Crosbyet al',1983)
Cryphaeaceae
Cryphaeaiamesonii Taytor (Crosbyet al',1983;|uo, 200t)
cryphaea rutenbergitMiill.Hal. (Crosbyet al.,,t.ltz; Rao, 2001)
O'Shea,2003)
Schoenobryumcoicavifulium (Griff.) Manuel (Crosbyet a1.,1983;
Daltoniaceae
Delgadillo,1-995)
Hampe (Crosbyet a1.,1983;
Adelotheciumbogotense
et
calyptrochaetaiplenioides (Brid.) crosby (crosby al-,1983)
Dultonia angustifoliaDozy e/ Molk. vat. angustifutia(Crosbyet al',1983)
Daltonia cardotiiBizot et Ont. (Crosbyet al',1983)
des
Daltoniaforsythii cardot (crgs.by.et a!.,1983) E.4, typ",^zone moy€nne
Maior,
Forsyth
1895,
m,
1350
ald.
forets, foret d'Ambohimitombo,
holotype PC.
Daltonia latimarginataBesch.(Crosbyet al', 1983)
Renauld (Crosbyet al',1983)
Daltonia latimarginata var.ma,d'agassa
*Distichophytlui m&sc&renicum
Besch.- Marojejy.\ut,t^o:9lPark, scattered"littt
forest, soil bank above the stream,s 14"26'E 49'45" alt. 1050m, Magill,
P6cs& La Farge9971,TAN'
Distichophyttum rakotomariae Crosby-(.Cros}y,t al-, .I983;^louw, L993)Ftd'
Type -- lvf"rt"gne d'Ambre,"Di6go-Suarez,alt. 900-1100m, M'R' &
MO, isotypesPC, TAN'
i.A Crotby 6883,holotype
Dicranaceae
Anisothecium madagassum Thdr. (Crosby et al., 1983) ETd,-Typ" - Yo1tagne
Perr:ierde La Bathie,holotype
d'Ambre. forOt, alt. 1000-1i00m,1011926,
PC.
(Crosby et al., L983;
Aongstroemia
filtfurmit (P. Beauv.) Wijk et Margad.
Delgadillo,1995)
Aongstroemiajulacea (Hook.) Mitt. (crosby et a1.,1983)
Dicianellq auoclada Cardot(Crosbyet al',1983)
226
1;
L.Marlineetal.
Dicranella borbonica var. madagascariensis
Th6r. (Crosbyet a1.,1983)End, Type
- Montagne d'Ambre, For€t du versant oriental-,1899,Girod, holotyire
PC.
Dicranella madagassaRenauld (Crosbyet a1.,I983)
D icrunella p ervi ll eanaB esch.(Crosby et al., 1983)
Dicranella polii Renauldet Cardot(Crosbyet a1.,1983)
Dicranella subsubulata(Hampeex Mtill.Hal.) A. Jaeger(Crosbyet a1.,1983)
Dicranoloma billardierei (Brid. ex Anon.) paris (Crosbyet a1.,19g3)
Dicranoloma bll^lyrdieri var. patentifolium (Renauld er Paris) Th6r. (Crosby
e ta 1 . , 1 9 8 3 )
Dicranoloma billardieri var. scopareolum(Mill.Hal.) Th6r. (Crosby et a1.,1983)
Dicranum johnstonii Mitt. (P6cs& Geissler,2}l2)
Holomitrium borbonicumBesch.(Crosbyet a1.,1983)
Holomitrium borbonicumvar.hamatum(Mult.Hal.)Th6r. (Crosbyet a1.,1983)
Holomitrium camptocarpum,P. de la-Varde (Crosby et al., 19g3) End, Type Pentes orientales du massif de Marojejy, d I'Ouest de la rivibre
Manantenina, affluent _d9 l? Lokoho, syive a Hchens sur gneiss et
quartzite,alt. 1400m,I5lI2ll948, Humbert, holotype pC.
Holomitrium cylindraceam var. cucttllatum (Besch.) Wijk et Margad. (Crosby
eta1.,1983)
Holomitrium cylin{a99yyt (P. Beauv.) Wijk & Margad. var. cylindraceum
(Crosbyet a1.,1983)
Holomitrium cylindraceum var. obtusifuIium (Besch.)Wijk et Margad. (Crosby
et a1.,1983)
Holomitrium gracilisetumThdr.(Crosbyet a1.,1983)End, Type - Ankaizina, alt.
1200-1500
m,0411933,
Decary,holotypepC..
Holomitrium hilli.eri Th6r. - (Crosby et al., 1983) End, Type
For€t 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 lepervancheiTh6r. (Crosbyet a1.,1983)
Holomitrium rhyparodictyon p.
4: la Varde (Crosby et a1.,19g3)End, Type Pente orientale du massif de Marojejy, sylve i lichens'et vdg€iaiion
f'ricoide des sommets,affluent de la Lokoho, alt. 1500 m. I7l12lIg48.
Humbert, holotype PC.
Holomitrium subglqQoylm Thdr. (Crosby et al., 1983) End, Type - Ifanadina,
for6t, herb (N.S.).
Leptotrichella minuta (Hampe) Ochyra (Crosbyet a1.,1983)
Leptotrichella pallidiseta (Brid.) ochyra (Crosby et al., 19g3) End, Type unknown.
Leucoloma albocinctum Renauldet Cardot (Crosbyet a1.,I983;La Farge,2002c)
Leucoloma allorgei^TlgL-(!19suy et-a1.,1983;La Farge,2002c)End, Type Befotaka,201081
1926,D ecary,holotypepC.
Leucoloma ambreanumRenauld-etCardot (Crosby et a1.,1983;La Farge,2002c)
Ftd, Typg Montagned'Ambre, prdsde Di6go-Suarez,1890,ehenogoi,
holotypePC.
Leucolomabifidum (Brid.) Brid. (Crosbyet a1.,1983)
Leucoloma boivinianum Besch.var. boivinianum (Crosbyet at.,I9g3)
Checklist of the bryophytes of Madagascar
Leucolomaboivini&numvar.brevifoliumRenauld(Crosbyet a1.,1983)End, Type
- Betsileo, entre Savondroninaet Ranomafana,1892,Besson,holotype
PC.
Leucoloma brevioperculatumDixon (Crosby et al., 1983;La Farge,2002c) End,
Thompson30, holotypeBM.
Type - Voh6mar,181,4,
La Farge,2002c)
Leucolomabrotheri Renauld(Crosbyet a1.,1983;
*Leucoloma caespitulans(Mi.ill.Hal.) A. Jaeger- Montagne des Frangais,bois
mort, alt. 400m,2510811973,
Cremers2657,TAN.
Leucoloma candidurnBroth. (Crosbyet a1.,1983;La Farge,2002c)End, Type 1904,Voeltzkow,
holotypeBR.
SainteMarie, East Madagascar,
Leucoloma capillifulium Renauld (Crosby et al., 1983; La Farge, 2002c) End,
Type
Sainte Marie, Sahasifotro,troncs d'arbres, 1894, Darbould,
lectotypePC.
Leucoloma chrysobasilare (Mi.ill.Ha1.)A. Jaeger var. chrysobasilare (Crosby
e ta \ . , 1 9 8 3 )
Leucoloma cinosulumRenauld(Crosbyet a1.,1983;
La Farge,2002c)End, Type For6t d'Analam azaotra, l-89L,Cambou4 & Campenon,holotype PC.
Leucoloma crosbyi La Farge-England(La Farge-England, 1992;La Farge,2002a;
La Farge,2002c)End, Type - Di6go-Suarez,Montagne d'Ambre, Parc
National,L0 km alongtrail betweenPetit 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 Mi.ill.HaI.et Geh.) C.H. Wright (Crosby et al.,
1983;La Farge,2002b;La Farge,2002c)
LeucolomadecaryiTh6r. (Crosbyet a1.,1983;
La Farge,2002c)End, Type - Massif
Decary,holotypePC.
d'Ankaizinana,L010611923,
LeucolomadelicatulumRenauld(Crosbyet a1.,1983;
La Farge,2002c)
Leucoloma dichelymoides(Miill.Hal.) A. Jaeger (Crosby et a1.,,1983;La Farge,
2002a)
Leacolomafontinaloides Dixon (Crosby et al., 1983;La Farge, 2002a;La Farge,
2002c)End, Type - Ambohimitomba forest, Tanala, 05101,11895,
ForsythMajor 473b,,holotype BM.
LeucoI oma fuscifu lium Besch.( Crosby et al., 1983)
Leucolomafuscifulium var. crispatulum Renauld (Crosbyet al., 1983)
Leucoloma garnieri Paris & Renauld (Crosby et al., 1983;La Farge,2002c) End,
Type - Ad radices arboris "Jisy", Dictae secusrivulum "Pleurs des
Bcufs", 2410811904,
Savelli,lectotypePC.
La Farge,2002c)End,
Leucolomagarnieri f. aquaticumTh€r.(Crosbyet a1.,1983;
Perrier de la Bathie,holotype PC.
Type - Massifde I'Andringitra, 1921,,
Leucolomagrandidieri Renauldet Cardot(Crosbyet a1.,1983;
La Farge-England,
1992; La Farge, 2002a; La Farge, 2002c) End, Type
For0t
d'Analamazoatra, 1891,Cambou| & Campenon, IectotypePC.
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 - Pentesorientalesdu massifde Marojejy, d
l'ouest de la rividre Manantenina,affluent de la Lokoho sylve d lichens
sur gneisset quartzite,alt. 1400m,1,511211948,
Humbert,holotypePC.
Leucoloma incrassatumTh6r. (Crosbyet a1.,1983;La Farge,2002c)End, Type Massifde I'Ikongo,rochershumidesen forOt,Decary,holotypePC.
L.Marline et al.
Leucoloma lepervancheri Besch. (La Farge,2002a; La Farge,2002c)
Leucoloma longifulium (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 (P6rinet), Reserve ForestiEre,
montane rainforest with shady, wet habitats along stream leading to lac
vert, Canopy reaching 25 m, mostly closed, with tall palms and bamboo,
alt.920-980m, S 18"54'00"E 48"25'3",1610311990,
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 Di6go-Suarez, Tahirin-javaboahary
Marojejy, north of Manantenina and Loko River Valley, slopes of the
Marojejy massif above Mandena village, S 1,4"26'E 49"44', La FargeEngland 6298, holotype ALTA, isotypes MO, TAN.
Leucoloma membran&ceum La Farge (La Farge ,2002a)
Leucoloma ochrobqsilqre Ren. (Crosby et a1.,1983;La Farge,2002c)
Leucoloma ochrobasilare (Th6r.) La Farge subsp. Iongifulium (Crosby et al.,
1983; La Farge, 2002a;La Farge,2002c) End, Type - Mont Tsaratanana,
forOt )r sous bois herbac6, alt. 1200-2400m,0411924, Perrier de la Bathie,
holotype PC.
Leucoloma ochrobssilare Renauld subsp. ochrobasilare (Crosby et al., 1983;
La Farge,2002a)
Leucoloma oncophorellum Mi.ill.Hal. (Crosby et a1.,1983; La Farge,2002c) End,
Type - lectotype PC.
Leucoloma perrieri Th6r. (Crosby et a1.,1983;La Farge,2002c) End, Type - For0t
d I'Est de Tananarive, IIll927, Perrier de la Bathie, holotype PC, isotype
FH.
Leucoloma persecundum Besch. (Crosby et a1.,1983)
Leucoloma persecundum var. perrolii Renauld (Crosby et al., 1983; La Farge,
2002c)End, Type - Environs d'Andevoranto, 1895,Perrot,lectotype BM.
Leucoloma pumilum C.H. Wright (Crosby et a1.,1983)End, Type - Sur le Plateau
central, d Alaby, prds de Maritandrano, Rutenberg, herb (N.S.).
Leucoloma pusillum Cardot (Crosby et al., 1983; La Farge,2002c) End, Type Di6go-Suarez, zone inf6rieure des forOts, sur 6corces, Ch€nagon,
lectotype PC.
Leucoloma rutenberg;; (Geh.) C.H. Wright (Crosby et a1.,1983;La Farge,2002c)
End, Type - Ambatondrazaka, Analamazaotra, Rutenberg, lectotype PC.
Leucoloma rutenbergii var. abbreviatumRenauld (Crosby et a1.,1983) End, Type
For€t d'Alakaty, Perrot; ForOt d'Analamazaotra, Cambou€ &
Campenon;holotype (N.S.).
Leucoloma rutenbergii var. elqtum Renauld (Crosby et a1.,1983;La Farge,2002c)
Leucoloma rutenbergii var. petotii Renauld (Crosby et a1.,1983;La Farge,2002c)
End, Type - Environs d'Andevoranto, 1895,Perrot, holotype PC, isotypes
BR, FH.
Leucoloma sanctue-mariqe Besch. (Crosby et a1.,1983;La Farge,2002a; La Farge,
2002c)
Leucoloma seychellenseBesch. (Crosby et al., 1983; La Farge, 2002a; La Farge,
2002c)
Leucoloma sinuosulumMnlLHal. ex Besch. (Crosby et a1.,1983)
Checklist of the bryophytes of Madagascar
Leucoloma squatrosulum (Geh.) Wright (Crosby et a1.,1983) End, Type - ForOt
de Vondr ozona, IIll877, Rutenberg, holotype PC.
Leucoloma subchrysobasilare Renauld (Crosby et al., 1983; La Farge, 2002a;
La Farge, 2002c) End, Type - In montibus Ankaratra, 1.877,Borgen,
lectotype PC.
Leucoloma tulazaccii Renauld et Cardot (Crosby et al., L983; La Farge, 2002a;
La Farge, 2002c) End, Type - Ambondrombe, 1.894,Talazac, holotype
PC, isotypes BR, PC, S.
Leucoloma thruustum Hampe ex Besch. (Crosby et al., 1983: La Farge, 2002a;
Alamazantraskoven, Borchgrevink,
La Farge, 2002c) End, Type
lectotype BM.
Leucoloma thuretii Besch. (Crosby et a1.,1983;La Farge,2002a; La Farge,2002c)
End, Type - COte Est, prds de la cascadede Simiang, Bernier,lectolype
BM.
Leucoloma tuberculosum Renauld (Crosby et a1.,1983;La Farge,2002a;LaFarge,
2002c) End, Type - For0t d'Analamazaotra,1891, Cambout & Campenon,
holotype PC.
Leucoloma villaumei'thdr. (Crosby et al., 1983; La Farge, 2002c) End, Type Betsileo, Villaume, holotype PC.
Microcampylopus laevigatus (Thdr.) Giese & J.-P. Frahm (Crosby et al., 1983;
Giese & Frahm, 1985)
Pseudephemerumnitidum (Hedw.) Reimers (Crosby et a1.,1983)
Ditrichaceae
Ceratodon pufpureus (Reichardt) Burley subsp. convolutzs (Crosby et a1.,1983;
Burley & Pritchard, 1990; Een, 1993)
Distichium capillaceum (Hedw.) Bruch, Schimp. et W. Guembel var. capillaceum
(Crosby et a\.,1983)
Ditrichum dfficile (Duby) M. Fleisch. (Crosby et aL.,1983;Delgadillo, L995)
Ditrichum luteum Dixon et Th6r. (Crosby et al., 1983) End, Type - Mont
Tsaratanana, Perrier de Ia Bathie, holotype PC.
Ditrichum madagassum (Renauld et Cardot) Paris (Crosby et a1.,1983)End, Type
-Andrangoloaka, 1891,Sikora, holotype PC.
Ditrichum pallidurz (Hedw.) Hampe (Crosby et a1.,1983)
Garckeaflexuosa (Griff.) Margad. e/ Nork. (Crosby et a|.1983)
Pleuridium nervosum (Hook.) Mitt. (Tixier, 1.979b;Crosby et a1.,1983)
Entodontaceae
Campylodontium flavescen^svar. viride Cardot (Crosby et a1.,1983) End, Type Plateau central, environs de Tananarive, Sibree,holotype BM#
Entodon corbieri Renauld et Cardot (Crosby et a1.,1983;Tixier, 1995) End. Type Montagne d'Ambre, prds de Di6go-Suarez, Leloutre, holotype CHE#
Entodon felicis Renauld et Cardot (Crosby et a1.,1983) End, Type - Fianarantsoa
(zone du Plateau central), Fdlix, holotype PC.
Entodon felicis var. julacezs Renauld et Cardot (Crosby et a1.,1983) End, Type Plateau central, Ambohimahatsara, prds d'Ambositra, Berthieu, herb
(N.s.).
Entodon madagasszs Geh. (Crosby et al., 1983) End, Type - Ambatondrazaka,
B orgen I 00, holotype PC.
L.Marline et al.
Entodon rutenbergi;.G.eh.(cqolby et ar., 1gg3) End, Type - Mont Ankaratra,
Borgen101,holotype PC.
Erythrodsntium lacoutourei Renauld et Cardot (Crosby et a1.,1983)Bnd, Type Fianarantsoa,
1905,Villaume,holotypefiU+
Mesonodonflavescens
(Hook.) w.R. Buck (crosby et a1.,19g3)
Erpodiaceae
Erpodium beccqrii Miill.Hal. exYenturi var. beccarii (Tixier, I979b;Crosbyet al.,
1e83)
Erpodiury madagass.um
Parise/ Renauld.(Crum, 1972;Crosby et a1.,1983)End,
Typ^e.-Antsirabe,vallde de la rividre Andrants ay, Gaiinor, Pc,froi"tyfe
(N.s.).
Eustichiaceae
Eustichia longirostris(Brid.) Brid. (Crosbyet a1.,19g3)
Fabroniaceae
Fabronia.campenoniiRenauld,et
(Crosbyet a1.,1983)End, Type
'^ - For6t
d'Amperifery,alt. 1300m,?-rOg!
06/1890,Campenon,holotypepc.
,
Fabronia crassiretisRenauld et cardot (crosby et a1.,19g3)
Fabronia fastigiata Renauld et Cardot (QrosUyet a1.,1983)End, Type - Zone du
Plateaucentral,Fianarantsoa,
pC.
1891,B-esson,holotype
Fabronia fastigiata var. asperulaRenauldet Paris (Crosbyet al., 1.983)End, Type
- Zone moyennedesfor6ts,cerclemilitaire desBarii, secteui
d'Ivondio
I8lIIlI900, Croll, holorypepC.
Fabronia garnieri (Paris e/ Renaul_d)
$enluld et Paris (Crosby et al., I9g3) Bnd,
Type - Betafo,rive de la Mania, Ig0Z,holotypeFC.
Fabroniq lqchenaudii Renauld (Crolby et al., 1983) End, Type -District de
Fihaonana,3llg0\,Ab indig. Coll30, holotypepC.
Fabronia motelayi Renauld et
..cardo1 lcloppv et al., 19g3) End, Type
Tananarive,dansles jardins
de la R6sidence,sur les troncs, I5l04ligg7,
Dorr, holotypePC.
Fabronia villaumii Renauld et Cal{ot (Crosby et al., 19g3) End, Type
Fianarantsoa,1905,Villaume 722,-holotypepC.
Rhizofabrolia, persoonii v9l sphaerocarpa (Dusdn) Bizot ex Ochyra (Crosby
et a1.,1983;
Touw, 1993)
Fissidentaceae
Fissidens^angustifuliusSull. (Bruggeman-Nanne
nga,2009;Bruggeman-Nannenga
& Arts,2010)
F issidensaristifer Brugg.-Nann. (Bruggeman-Nannenga,
z00g)
Fissidens^asplenioides
Hedw. (Crosbyet a1.,1983;Bruggeman-Nannenga
& Arts,
20r0)
Fissidens_
_bogosicusMtill.Hal. in Venturi (Crosby et al., 1983; BruggemanNannenga& Arts, 2010)
F i ssi densborgenii Hampe (Bruggeman-Nanne
nga, 1993)
FissidenscrispulusBrid. (Crosbyet a1.,1983;Bruggeman-Nannenga
& Arts, Z0I0)
*FissidenscryptarurnMtill.Hal. - Tampolo
Tamatave,littoral foiest baseof tree,
eartheninsectlest, ant or termit, s 17'16,E 49"24,,2611011972,
M.R. &
C.A Crosby6700,6702,,6697,
TAN.
FissidenscryptoneuronP.de la varde (crosby et a1.,19g3)
Checklist of the bryophytes of Madagascar
231
Fissidens curyatus 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, 2411112009,Wilding 4, det.
Bruggeman-Nannenga; Marojejy National Park, on tree fern 300 m up
trail from camp Mantella, alt.465 m,24llIl09,Wilding 6, det. BruggemanNannenga.
Fissidens darntyi Schimp. (Crosby et a1.,1983)
Fissidens ellipticus Besch. (Crosby et a1.,1983)
F i ssi dens fl acci dus Mitt. (B ruggeman-Nanne nga, 2009)
Fissidens grainvilleiThdr. (Crosby et a1.,1983)End, Type - Kiangara, sur 6corces,
en trds petite quantit6, 1931,,Grainville, holotype PC.
Fissidens hookerioidesBizot et Onr. (Crosby et a1.,1983)Bnd, Type - Tananarive,
domaine des Frdres de Soavimbahoaka, sur Iat1,rite ombragde,
alt. 1400 m,0610711971,,
Onraedt 71M5137, holotype PC.
Fissidens hymenodon Besch. (Crosby et a1.,1983)
Fissidens inclusus Bizot et Dury ex P6cs (Bruggeman-Nannenga,2009)
Fissidens intramarginatus (Hampe) A. Jaeger (Bizot, I974; Crosby et al., 1983;
Bruggeman-Nannenga & Arts, 20t0)
Fissidens leucocinctusHampe (Crosby et a1.,1983;Bruggeman-Nannenga & Arts,
2010)
Fissidens madecasszsMi.ill.Hal. (Crosby et al.,1983; Bruggeman-Nannenga,2009 ;
Bruggeman-Nannenga & Arts, 2010)
Fissidens mariei (Besch.) Broth. (Crosby et a1.,,1983)End, Type - Nosy Be, forOt
de Lokobe,04lI879, Marie, holotype PC.
Fissidens motelayi Renauld et Cardot (Crosby et al.,1983) End, Type - Andriba,
1897,Dorr, holotype PC.
Fissidens ovatus Brid. (Crosby et a1.,1983;Bruggeman-Nannenga & Arts, 2010)
Fissidens pellucidu^s Hornsch. var. pellucidus (Crosby et al., 1983; Delgadillo,
1995; Bruggeman-Nannenga,2010)
Fissidens planifron^s Besch. (Crosby et al., 1983; Bruggeman-Nannenga & Arts,
2010)
Fissidens pletyneuros Renauld et Cardot (Crosby et al., 1983) End. Type Andevoranto, 1-895,Perrot, holotype PC.
Fissidens plumosus Hornsch. (Bruggeman-Nannenga & Arts, 2010)
Fissidens pocsii Brzot et Dury ex P6cs (Bruggeman-Nannenga,2009; BruggemanN a n nenga& A rts,2010)
Fissidens punctulalzs Sande Lac. (Bruggeman-Nannenga & Arts, 2010)
Fissidens ramulosus Mitt. (Crosby et al., 1983; Bruggeman-Nannenga & Arts,
20ro)
2009)
FissidensreflexusHampe (Crosbyet al., 1983;Bruggeman-Nannenga,
FissidensrosulatusP. de la Varde (Crosbyet a1.,1983)End, Type - Tananarive,
PC.
Rotereau,holotype
sur talus,2010111907,
F i ssi densrufescensHornsch. (B ruggeman-Nannenga,2009)
Fissidens sciophyllus f. sciophyllus Mitt. (Crosby et al. 1983; BruggemanNannenga, L993;Bruggeman-Nannenga
& Arts, 2010)
1891;Crosbyet a1.,1983)
Fissidensscleromitrizs(Besch.)Broth. (Bescherelle,
FissidenssematusMtill.Hal. var. senutus (Crosby et al., 1983;Delgadillo, 1995;
20L0)
Bruggeman-Nannenga,
L.Marline et al.
Fissidens submarginatus Bruch ex C. Krauss (Crosby et al., 1983; BruggemanNannenga & Arts, 2010)
Fissidens subplanifrons Bizot et Onraedt ex Brtgg.-Nann (Bruggeman-Nannenga,
200e)
Fissidens subundatus Dury (Crosby et a1.,1983)
Fissidens zollingeri Mont. (Crosby et al., 1983; Bruggeman-Nannenga & Arts,
2010)
Funariaceae
Entosthodon borbonicus Besch. (Crosby et a1.,1983)
Entosthodon marginatulus Miill. Hal. ex Geh. (Crosby et a1.,1983) End, Type Zone moyenne des forOts, forOt )r I'Est d'Ambatondrazaka, Rutenberg,
herb (N .S .).
*Entosthodon mauritianus Schimp. ex Besch. - Befotaka forOt, S 1.8"56'47"
E 47"06'03",1610811926,
Decary TT,holotype MO.
Entosthodon renauldii Th6r. (Crosby et a1.,1983) End, Type - Zone du Plateau
central, pays Bets ileo, 2l 1905, Villaume, holotvpe PC, isotype PC, BM.
Funaria decaryi Th6r. (Crosby et al., 1983) End, Type - Ambovombe, FortDauphin, Decary, holotype S.
Funaria delicatula Th6r. (Crosby et a1.,1983)End, Type - Environ de Tananarive,
Perrier de la Bathie, holotype PC.
Funaria hygrometrica var. calvescens (Schwaegr.) Kindb. (Crosby et al., 1983;
P6cs & Geissler,2}02)
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.
Funqria subleptopodqHampe (Crosby et a1.,1983)End, Type - Imerina,2l9lI847,
Borgen, holotype PC.
Physcomitrium coarctatum Cardot (Crosby et a1.,1983)
Physcomitrium dilatatum Renauld et Cardot (Crosby et al., 1983) End, Type Environ de Tananarive, in terra, Cambou6, holotype PC.
Gigaspermaceae
Gigaspermum repens (Hook.) Lindb. (Tixier, L979b; Crosby et a1.,1983)
*Oedipodiella australl's (Wager & Dixon) Dixon - Morondava, forOt, alt.20 m,
1.110411979,
Tixier 12114,TAN.
Grimmiaceae
Grimmia elatior Bruch ex Bals.-Criv. et De Not. (Crosby et a1.,1983)
Grimmia elongata Kaulf. (Crosby et a1.,1983)
Grimmia longirostris Hook. (Mufroz, 1998)
Bucklqndiella membranaceun (Mitt.) Paris (Crosby et a1.,1983)
Hedwigiaceae
*Braunia secunda (Hook.) Bruch e/ Schimp. - Station forestibre de Manjakatompo,
rocher, alt. 2600 m, 2210811971,Cremers 1705, TAN; Ankaratra, sur
rocher, 2710311972,Cremers 1957, TAN; Ankaratra, sur rocher,
basaltique, 08/1955,Bosser 8208, TAN.
Bryowijkia madagassa Touw (Touw, 1993)
Hedwigia ciliata (Hedw.) Ehrh. exP. Beauv. var. ciliara (Crosby et a1.,1983)
Checklist of the bryophytes of Madagascar
233
Hypnaceae
EctropotheciumchenagonifRenauld et Cardot (Crosbyet a1.,1983)
Ectropothecium chenagonii var. hamatulum Renauld & Cardot (Crosby et al.,
1983) End, Type - Entre Vinanitelo et Ikongo, 1892,Besson,holotype
PC.
EctropotheciumintertexturnRenauldet Cardot (Crosbyet a1.,1983)
EctropotheciummauritianumRenauldet Cardot (Crosbyet a1.,1983)
Ectropotheciumoccultum Renauld et Cardot (Crosbyet a1.,1983)
Ectropotheciumovalifulium (Besch.)W.R. Buck (Crosbyet a1.,1983)End, Type Zone inf6rieuredesforOts,Nosy Be, forOtde Lokobe,Marie, herb (N.S.).
Ectropotheciumpaillotii Renauld et Cardot (Crosbyet a1.,1983)
Ectropothecium perrieri Th6r. (Crosby et al., 1983) End, Type - Massif de
l'Andringttra, alt.1600m,I92l, Perrierde la Bathie,holotypePC.
Ectropotheciumperrotii Renauldet Cardot (Crosbyet a1.,1983)
Ectropotheciumregulare(Brid.) A. Jaeger(Crosbyet a1.,1983)
Ectropotheciumseychellarum Besch.(Crosbyet al., 1983)
Ectropotheciumtamatavense
Broth. (Crosbyet a1.,1983)
(Broth.)
B.C. Tan et W.R. Buck (Crosbyet a1.,1983)
Gammiella ceylonensis
Glossadelphusguineensl's(Broth. et Paris) Crosby,B.H. Allen et Maglll (Crosby
e ta 1 . , 1 9 8 3 )
Glossadelphussemiscabrzs(Renauld et Cardot) Crosby et al. (Crosbyet a1.,1983)
End, Type - Zone inf6rieure des for6ts, environs d'Andevoranto, 1-895,
Perrot,holotype PC.
Glossadelphusviridis (Renauld et Cardot) Crosby et al. (Crosby et a1.,1983)End,
Type - Di6go-Starez,Montagned'Ambre, ChEnagon,holotype
PC.
Hypnum jutlandicum Holmen et E. Warncke (Crosbyet a1.,1983)
(Crosbyet a1.,1983)
Hypnum macrogynumBesch.
Mittenothamniumhonidulum (Broth.) Cardot (Crosbyet a1.,1983)
Mittenothamnium lacouturei (Renauld et Cardot) Cardot (Crosby et al., 1983)
End, Type - Betsileo, province d'Ambositra, 1906, Villaume, holotype
BM#
Mittenothamniumlimosum (Besch.)Cardot (Crosbyet a1.,1983)
Mittenothamnium madagassum(Besch.)Cardot (Crosbyet al.,1983;Een, 1997)
Mittenothamniummicrothamnioides(Geh.) Wijk etMargad.(Crosbyet a1.,1983;
Een, 1997)
Mittenothamnium microthamnioides var. glauculum (Cardot) Wijk et Margad.
(Crosby et al., 1983) End, Type - For0t de Manongarivo,Drouhard,
holotype PC#
Mittenothamniumreptans(Hedw.) Cardot (Crosbyet a1.,1983)
Phyllodon scutellifolius(Besch.)W.R. Buck (Crosbyet a1.,1983;O'Shea& Buck,
200I) End, Type - Nosy Komba, associ6d Hookeria nossiana,08ll879,
Marie s.n.,holotypeBM.
Phyllodon truncatulzs(Welw. etDuby) W.R. Buck (Crosbyet a1.,1983)
Rhacopilopsistrinitensis(Miill.Hal.) E.Britton ex Dixon (Crosbyet a1.,1983)
Rhacopilopsis variegata (Welw. er Duby) M.C. Watling e/ O'Shea (Buck, 1993;
Watling & O'Shea,2000)
Taxiphyllum taxirameum(Mitt.) M. Fleisch.(Crosbyet a1.,1983)
234
L.Marline et al.
Vesicularia alboviridis var. rufula (Renauld et Cardot) Cardot (Crosby et al.,
1983) End, Type - Zone inf6rieure des forOts, Di6go-Suarez, Ch4nagon,
holotype PC.
Vesicularia ayresii (Besch.) Broth. (Crosby et a1.,1983)
Vesicularia combae (Renauld et Cardot) Broth. (Crosby et a1.,1983) End, Type Zone infdrieure des forOts,Nosy Komba, Marie, holotype PC#
Vesicularia crqssiramea (Renatld et Cardot) Broth. (Crosby et a1.,1983)
Vesicularia crassiramea var. laxifulia (Renauld et Cardot) Cardot (Crosby et al.,
1983) End, Type - Zone des savanesoccidentales,cercle de Maintirano,
holotype PC.
Vesicularia galerulata (Duby) Broth. (Crosby et a1.,1983)
Vesicularia galerulata var. subpiligera (Renauld e/ Paris) Wijk et Margad.
(Crosby et a1.,1983) End, Type - Zone du Plateau central, Betafo, rive
droite de la Mania, 1901, Galinon, holotype PC.
Vesicularia hygrobium (Besch.) Broth. (Crosby et a1.,1983)
Vesicularia immunda (Renauld) Cardot (Crosby et a1.,1983) End, Type - Zone
du Plateau central, Imerina oriental, Hildebrandt,herb (N.S.).
Hypopterygiaceae
Hypopterygium tamarisci (Sw. ex Sw.) Brid. ex Miill.Hal. (Crosby et al., 1.983:
Kruijer 1997; Kruijer, 2002)
Lopidium struthiopteris (Brid.) M. Fleisch. (Crosby et al., 1983; Kruijer, 1997;
Kruijer,2002)
Leptodontaceae
Leptodonfucifurmr's (Brid.) Enroth (Crosby et a1.,1983)
Leskeaceae
Pseudoleskea obtusiuscula Renauld et Cardot (Crosby et a1.,1983) End, Type Zone du Plateau central, Fianarantsoa, troncs d'arbres, 1904, Villaume,
holotype BM!, isotype PC.
Leucobryaceae
Atractylocarpus madagascariensis (Th6r.) 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 a1.,1983;Frahm, 1985)
Campylopus aureonitens (Mi.ill.Hal.) A. Jaeger (Crosby et a1.,1983;Frahm, 1985)
Campylopus camboueiRenauld et Cardot (Crosby et a1.,1983;Frahm, 1985)
Campylopus catatectilis (Mtill.Hal.) Paris (Frahm, 1985)
Campylopus crateris Besch. (Crosby et a1.,1983;Frahm, 1985)
Campylopus decaryf Th6r. (Crosby et a1.,1983;Frahm, 1985)
Campylopusfilifulius (Hornsch.) Mitt. (Crosby et a1.,1983;Een, 1993)
Campylopusflaccidu.s Renauld et Cardot (Crosby et a1.,1983;Frahm, 1985)
Campylopus flexuoszs (Hedw.) Brid. var. flexuoszs (Crosby et al., 1983; Frahm,
198s)
C ampy lopus hildebrandtii (Mi.i1l.Hal.) A. Jaeger (Crosby et al., 1983)
Campylopus jamesonid (Hook.) A. Jaeger (Crosby et a1.,1983;Frahm, 1985)
Campylopus julacezs (Renatld et Cardot) J.-P. Frahm subsp. arbogastii (Crosby
et a1.,1983;
Frahm, 1985)
Checklist of the bryophytes of Madagascar
235
CampylopusnanophyllusMtill.Hal. exBroth. (Crosbyet a1.,1983)
Campylopusnivalis (Brid.) Brid. var.nivalis (Frahm,L985)
Frahm,1985)
CampylopusperpusillusMitt (Crosbyet a1.,1983;
Frahm, 1985)
Campylopuspilifer Brid. subsp.pilifur (Crosbyet a1.,1983;
CampylopuspseudobicolorMtill.Hal. er Renauld & Cardot (Crosbyet a1.,1983;
Frahm,1985)
Frahm,1985;P6cs& Geissler,
CampylopusrobillardeiBesch.(Crosbyet a1.,1983;
2002)
Frahm, 1985)
Campylopusschmidit (Miill.Hal.) A. Jaeger(Crosbyet a1.,1983;
1985)
Campylopussm&ragdinus(Brid.) A. Jaeger(Crosbyet a1.,,1983;Frahm,
Frahm, 1985)
Campylopusthwaitestt(Mitt.) A. Jaeger(Crosbyet a1.,1983;
Campylopus trachyblepharon (Renauld et Cardot) J.-P. Frahm subsp.comatus
(Crosbyet a1.,1983;
Frahm,1985)
Leucobryumacutifolium (Mitt.) Cardot (Crosbyet a1.,1983)
LeucobryumboivinianumBesch.(Crosbyet a\.,1983)
(Crosbyet a1.,1983)
LeucobryumboryanumBesch.
Leucobryumboryanum var. minus Cardot (Crosbyet a1.,1983)End, Type - Pays
Betsileo,sineloco,Montaul,herb (N.S.).
LeucobryumcumboueiCardot (Crosbyet al., 1983)End, Type - Cambou6,herb
(N.s.).
Miill.Hal. (Crosbyet a1.,1983;
P6cs& Geissler,2002)
Leucobryumcomorense
LeucobryumcucullatumBroth. (Crosbyet a1.,1983)
Leucobryumcucullatumvar. rutenbergii(Geh.) Cardot (Crosbyet a1.,1983)
End, Type - Betafo,
Leucobryumgalinonii Cardot et Paris (Crosbyet a1.,,1983)
rive droite de la Mania, Galinon,holotypePC#
LeucobryumheterodictyonBesch.(Bescherelle,1891;Crosby et al., 1983) End,
Type -SainteMarie, Marie EdouardAuguste,holotypePC.
Leucobryum heterodictyonvar. miuophyllum (Cardot) Wijk etMargad. (Crosby
et a1.,1983)End, Type - Ile SainteMarie, Marie,holotypePC#
LeucobryumisleanumBesch.(Crosbyet al., 1983)
Leucobryumisleanumvar. molle (Miill.Hal.) Cardot (Crosbyet a1.,1983)
Leucobryummadagassum
Besch.(Crosbyet a1.,1983)
LeucobryummqyottenseCardot (Crosbyet a|.,1983)
Leucobryumparyulum Cardot (Crosbyet a1.,1983,P6cs& Geissler,2002)End,
Type - holotypePC.
Leucobryumperrolii Renauld et Cardot (Crosby et al., 1983,P6cs & Geissler,
2002)
Leucobryumpseudomadagassum
Cardot (Crosbyet a|.,1983) End, Type - Zone
sup6rieure des forOts, Imerina, Andrangoloaka, Hildebrandt, holotype
PC.
Leucobryumsanctae-mariaeCardot (Crosbyet al.,1983,P6cs& Geissler,2002)
End, Type - Ile SainteMarie, Marie, holotypePC.
Ochrobryumsqkalavum Cardotet Paris(Crosbyet al.,1983;Allen, 1992;P6cs&
Geissler,2002)End, Type - cerclede Maintirano,poste d'Ampoza,prds
du villagede Tahotofoly,sur bois pourri, holotypePC!
Leucodontaceae
Leucodoncanariensis(Brid.) Schwaegr.(Crosbyet a1.,1983)
LeucodonrutenbergitMtill.Hal. ex Geh. (Crosbyet a1.,1983)
L.Marhne et al.
Pterogoniadelphus assimilis (Miill.Hal.) OchyraetZljlstra (Crosby et a1.,1983)
Pterogonium gracile (Hedw.) Sm. (Crosby et al.,1983)
Pterogonium gracile var. madagassum (Geh.) M. Fleisch. (Crosby et a1.,1983)
Leucomiaceae
Leucomium strumosum (Hornsch.) Mitt. (Crosby et a1.,1983;Allen, 1,987;P6cs &
Geissler,2002)
Meteoriaceae
Aerobryopsis capensis(Mi.ill.Hal.) M. Fleisch. (Crosby et a1.,1983)
Barbella capillicazh's (Renatld et Cardot) Cardot var. capillicaulis (Crosby et al.,
1eB3)
(DozyerMolk.)M. Fleisch.(Crosbyeta1.,1983)
Floribundariafloribunda
Floribundaria floribunda var. madagassa (Ren auld et Cardot) Renauld et Cardot
(Crosby et al., 1983) End, Type - For0t d'Ambohimitombo, Forsyth
Major, FH; Fianarantsoa, plateau d'Ikongo, entre Savondronina et
Ranomafana, et entre Vinanitelo et Ikongo, Bessoru,holotype (N.S.).
F loribundaria floribundulu (Mi.ill.Ha1.) M. Fleisch. (Crosby et al., 1983)
Floribunduriu vaginans (Welw. etDuby) Broth. (Crosby et a1.,1983)
Papillaria africana (Miill.Hal.) A. Jaeger (Crosby et aL.,1983)
Papillaria african& var. gracilis (Renauld et Cardot) Wijk et Margad. (Crosby er
al.,1983) End, Type - Didgo-Srrarez,1890, Chenagon; Entre Vinanintelo
et Ikongo, 1892, Besson. Betsileo, sine loco, Rev. Montaut; Entre
Savondronina et Ranomaf ana,1892,Besson,holotype (N.S.).
Papillaria borchgrevinkii Kiaer (Crosby et a1.,1983)
Papillaria flaccidula Cardot (Crosby et a1.,1983) End, Type - Zone moyenne des
forOts,forOt d'Analamazaotra, Cambou6, herb (N.S.).
Papillaria perichaetialis (Hampe) A. Jaeger (Crosby et al., 1983) End, Type Analama zaotr a, 1,87l, B o r chgr ewin k 1 7, holotyp e B M.
Trachypodopsis serrulats (P. Beauv.) M. Fleisch. var. serculata (Yan Zanten,
1959;Crosby et a1.,1983;P6cs & Geissler,2002)
Trachypus appendiculatus (Renauld et Cardot) Broth. (Van Zanten,1959; Crosby
eta1.,1983)
Trachypus bicolor var. viridulus (Mitt.) Zanten (Van ZanIen, 1959; Crosby et al.,
1e83)
Meesiaceae
Leptobryum pyrifurme (Hedw.) Wilson (Crosby et a1.,1983)
Mniaceae
Epipterygium diversifolium Renauld et Cardot (Crosby et a1.,1983) End, Type Zone des savanes occidentales, cercle de Maintirano, 1,90I, Collecteur
indigAne, holotype PC.
Mielichhoferia cardotii A.J. Shaw etH.A. Crum (Crosby et a1.,1983)End, Type Zone de plateau central, Ambohipo, Cambou'/,herb (N.S.).
Mielichhoferia madagassa var. gracilis Thdr. (Crosby et al., 1983) End, Typ" Mont Angavokely, alt. 1800 m,1924, Drouhard, holotype PC.
Plagiomnium rhynchophorunz (Hook.) T.J. Kop. var. rhynchophorunt (Koponen,
1 981;C rosby et a\.,1983)
Pohlia baronii Wijk etMargad (Crosby et a1.,1983)
Pohlia elongata Hedw. var. elongata (Crosby et a|.,L983; Delgadillo, 1995)
Checklist of the bryophytes of Madagascar
237
Pohlia heribaudii var. madagessa (Renauld et Cardot) Wijk etMatgad. (Crosby
eta\.,1983)
Pohlia lacouturei (Th6r.) O'Shea (Crosby et al., 1983) End' Type - Pays des
Betsileo, sur gneissen d6composition, Villaume & Lacouture,herb (N.S.).
Pseudopohlia microstoma (Harv.) Mizush. (Crosby et a|.,1983)
n"{. Type Schizymenium commixtum (Th6r.) A.J. Shaw (Crosby_et a!- Il8!
Mont Tsaratanana, for0t )r sous-bois herbac6,, alt.1200-2400 m, Perrier de
la Bathie, holotyPe PC.
Schizymenium madagassun (Ren auld et Cardot ex Cardot) 4.J. Shaw (Crosby
Mont Angavokely, Est Antananarivo,
et al., 1983t End, Type
alt. 1800m,1924, Drouhard, holotype PC.
Neckeraceae
Homaliodendron exiguum (Bosch e/ Sande Lac.) M. Fleisch. (Crosby et a1.,1983)
Homaliodendron pinifurme (Brid.) Enroth (Crosby et a1.,1983;Enroth, 1993)
Neckera remota Bruch e/ Schimp. ex Mi.ill.Hal. (Crosby e/ a1.,1983)
N eckera val entiniana B esch.(Crosby et al., 1983)
Neckeropsis boivinianu (Besch.) Cardot (Crosby et a\.,1983)
*Neckeropsis disticha (Hedw.) Kindb. - Brickaville, largg trees next to shallow
rive, S L8'55' E 48"34',att. 450 m,2311011972,M.R. & C.A Crosby 6754,
TAN.
Neckeropsis lepineana (Mont.) M.Fleisch. (Crosby et a1..1983)
Neckeropsis msdecassa(Besch.) M. Fleisch. (Crosby et a|.,1983)
Orthostichidium pentestichum (Brid.) B.H. Allen et Maglll (Crosby et a1.,1983;
Allen & Magill,2007)
Orthostichella rigida (Mtill.Hal.) B.H. Allen et Magill. (Allen & Magill ,2007)
Orthostichella vercicolor (Miill.Hal.) B.H. Allen er Magill. (Crosby et al., 1983;
Allen & Magill,,2007)
Orthostichella longinervis (Renauld et Cardot) B.H. Allen et }dagtlL (Allen &
Magill, 2007)
Orthostichella welwitschii (Duby) B.H. Allen et Maglll. (Allen & Magill ,2007)
Pilotrichellaflexilis (Hedw.) Aongstr. (Crosby et a1.,1983;Allen & Magill,2003)
Pilotrichetla flexilis f. nudiramulosa (Mtill.Hal.) B.H. Allen et Magtll (Allen &
Magi l l ,2003)
Pilotrichella mqscarenica (Mtill.Hal.) A. Jaeger (Crosby et al., 1983; Allen &
Magill, 2003)
Pilotrichella pseudo-imbricata (Mid.ll.Hal.)A. Jaeger (Crosby et a1.,1983)
Pinnqtellq minuta (Mitt.) Broth. (Crosby et a1.,1983;Enroth, 1994)
Porotrichum elongatum (Welw. er Duby) A. Gepp (De Sloover, L983)
Porotrichum madagnssum Kiaer ex Besch. (De Sloover, L983)
Porotrichum stipitatum (Mitt.) W.R. Buck (De Sloover, L983)
Porotrichum usagarumMitt (D. Sloover, 1983)
Porotrichum variifolioides (De Sloover) Enroth (De Sloover, 1983)
Thamnobryum malgachum (Cardot) O'Shea (Crosby et al., 198?) Et!, TyP" :
Zone moyenne des for0ts, forOt de Manongarivo, 1909, Drouhard,
holotype PC.
0rthotrichaceae
Curdotiella appendiculata (Renauld et Cardot) Vitt (Crosby et a1.,1983; Goffinet,
1996)
L.Marline et al.
Cardotiella elimbata (Thdr.) Goffinet (Crosby et a1.,1983;Goffinet, 1996) End,
Type - Farafangana,sur souched'arbre, en for0t, Decary,holotype PC,
isotypeS.
Cardotiella renauldii (Broth.) Vitt. (Crosbyet a1.,1983;Goffinet, 1996)End, Type
- holotype PC.
Cardotiella subappendiculata(Broth.) Vitt (Crosbyer a1.,1983)
Groutiella laxotorquata (Besch.)Wijk etMargad.(Crosbyet a1.,1983)
Macrocomatenuis(Hook. et Grev.)Vitt subsp.tenuis(Crosbyet a|.,1983)
Macromitrium adelphinum Cardot (Crosby et al., 1983) End, Type - For0t
d'Amperifery,Campenon,herb (N.S.).
Macromitrium ancistrophyllum Cardot (Crosby et al., 1983) End, Type - Zone
sup6rieuredesforOts,Ankeramadinika,Borgen202,holotype BM.
Macromitrium anomodictyon Cardot (Crosby et al., 1983) End, Type - District
d'Andramasina,entre Tsinjoarivo& Ambohimasina,dansun petit bois,
sur branched'arbre,alt. L700m,02lI2l1.9I2,Viguier1932,holotypePC.
Macromitrium calocalyx Geh. (Crosby et al., 1983)End, Type - Zone moyenne
Rutenberg,herb(N.S.).
desforOts,forOti I'est d'Ambatondrazaka,
M acromitrium fasciculare Mitt. (Crosbyet al., 1983)
Macromitrium fasciculare var. angustifolium P. de la Varde (Crosby et a1.,1983)
End, Type - Sommet oriental du Massif de Marojejy, d I'Ouest de la
Haute Mantenina,affluent de la Lokoho, gneisset quartzite,alt. 1850m,
Humberr,holotypePC.
261311949,
M acromitrium fimbriatum (P.Beauv.) Schwaegr,(Crosbyet al., 1983)
(Crosbyet a1.,1983)
Macromitrium mauritianum Schwaegr.
Macromitrium nanothecium Cardot (Crosby et al., 1983) End, Type - Zone
sup6rieuredesforOts,Andrangoloaka,H ildebrandt, holotype FH.
Macromitrium nanotheciumvar.sublaeveTh6r. (Crosbyet a1.,1983)End, Type Fort-Dauphin,sur ecorce,Decary,holotypePC!
Macromitrium onraedtiiBizot (Bizot, 1974;Crosbyet a1.,1983)End, Type - Entre
Foulpointe et Andondabe,au Nord de Tamatave,sur un arbre d'une for€t
Cremers2165,holotypePC.
sur sable,1210511972,
Nees
ex Schwiigr.(Crosbyet a1.,1983)
orthosticharz
Macromitrium
Mont
proximum
Th6r. (Crosby et al., 1983) End, Type
Macromitrium
Tsaratanana,alt. 1200m,0411924,Perrier de la Bathie,holotype PC.
Macromitrium rhizomatosum Mid,Il.Hal.(Crosby et al., 1983) End, Type - Nosy
Be, PerviUe789,holotype PC.
Macromitrium sanctae-mariaeRenauld et Cardot (Crosby et a1.,1983)End, Type
- SainteMarie, l-893,Darbould,holotypePC.
Macromitrium sclerodictyonCardot (Crosbyet a|.,1983)
Macromitrium semipapillosumTh6r. etP. de la Varde (Crosbyet a1.,1983)End,
Type - 1922,Carrougeau,holotypePC.
Macromitrium serpens(Bruch ex Hook. & Grev.) Brid. (Crosbyet a1.,1983)
Macromitrium soulae Renauld et Cardot (Crosby et aI., 1983) End, Type Ambositra,Betsileo,L890,Soula,holotypePC.
Macromitrium subpungensHampe ex Miill.Hal. (Crosbyet a1.,1983)
var. madagassumCardot (Crosby et al., 1983) End,
Macromitrium subpungen^s
Type - Zone inf6rieure desfor6ts, Maroantsetradansla baie d'Antongil,
Mathieu,herb (N.S.).
Checklist of the bryophytes of Madagascar
*
239
Macromitrium subtortum (Hook. et Grev.) Schwaegr.(Crosbyet a1.,1983)End,
Type - unknown.
Macromitrium sulcatum (Hook.) Brid. var. sulcatum (Crosby et al.,1983; Ellis &
Wilbraham,2008)
Macromitrium urceolatulumldid,llHal. ex Geh. (Crosbyet a1.,1983)
Schlotheimiaacutifulia Renauldet Paris(Crosbyet a1.,1983)End, Type - For0t
d'Ambohitantely, 190L,Collecteurindigdne,holotype PC.
Schlotheimia acutifolia var. angustifulia Th6r. (Crosby et a1.,1983)End, Type Mont Tsaratanana,forOt d sousbois herbac6,alt.1200-2400m, Perrier de
la Bathie,holotype PC.
Schlotheimiabadiella Besch.(Crosbyet a1.,1983)
Schlotheimiabadiella var.helicophyllaBesch.(Crosbyet a1.,1983)
Schlotheimiaboiviniana Besch.(Crosbyet a1.,1983)
Schlotheimia conica Renauldet Cardot (Crosbyet a1.,1983)End, Type - Environs
de la forOtd'Ambatovory,alt. L400m, 1889,Cambou6,holotypePC.
Schlotheimia excorrugataMtill.Hal. ex Cardot (Crosbyet a1.,1983)
Schlotheimiafornicata Duby (Crosbyet al., 1983)
Schlotheimiafornicata var. apiculata P. de la Varde (Crosby et al., 1983) End,
Type - Pentesorientale du massifde Marojejy, d l'ouest de la rivibre
Manantenina, affluent de la Lokoho, sylve e Hchens sur gneiss et
quaftzite,alt. 1400m,,1,511211948,
Humbert, holotype PC.
Schlotheimiafornicata var. microthecaP. de la Varde (Crosbyet a1.,1983)End,
Type - Vall6e de la Lokoho Nord-Est, mont Beondroka, au nord
de Maroambihy, rochers de la crOte(gneisset quartzite), alt. 1400m,
I710311949,
Humberf,holotypePC.
SchlotheimiafoveolataRenauldet Cardot(Crosbyet a1.,1983)End, Type -Zone
inf6rieure des for€ts. Ile de Sainte Marie sur les bords de la rivibre
Fitarihana,1893,Darbould,holotypePC.
Schlotheimiafoveolata var. torquescensCardot (Crosby et a1.,1983)End, Type Zone du Plateau central, Imerina, environ de Tananarive, Cambout &
Sibree,holotypeBM.
Schlotheimiagauthieri Th6r. (Crosbyet a1.,1983)
Schlotheimia malacophylla Besch.(Crosbyet al., 1983)
Schlotheimia microcarpa Geh. (Crosbyet al., 1983)
Schlotheimia miuophylla Besch.(Crosbyet al., 1983)
Schlotheimia nossi-beanaMtill.Hal. (Crosby et a1.,1983) End, Type - Nosy Be,
184'1,,
Pervill4,holotype PC.
Schlotheimia nossi-beanavar. lineqlis (Geh.) Th6r. (Crosby et al., 1983) End,
Type - Nosy Be, Boivin, holotypePC.
Schlotheimia pauli Renauld et Cardot (Crosby et al., 1983) End, Type
Antsahambavy,1902,Dalincourr,holotypePC.
Schlotheimiaperrieri Th6r. (Crosbyet a1.,1983)End, Type - Mont Tsaratanana,
Perrier de la Bathie,holotype PC.
alt. 1200m,411924,
Schlotheimia perrotii Renauld et Cardot (Crosby et al., 1983) End, Type Andevoranto,1895,Perrot,holotypePC.
Schlotheimia perrotii var. breviseta Th6r. (Crosby et al., 1983) End, Type Andevoranto,herb (N.S.).
Schlotheimiarobillardii Duby (Crosbyet a1.,1983)
SchlotheimisrobustaTh6r. (Crosbyet a1.,1983)
L.Marline et al.
Schlotheimiq semidiaphana (Renauld et Cardot) Cardot (Crosby et a1.,1983) End,
Type - Plateau d'Ikongo ,1892, Besson 294, holotype PC.
Schlotheimia squar osa Brid. (Crosby et al.,1983; Delgadillo,1995)
Schlo theimia subfornicata Th6r . (Crosby et al., 1983)
Schlotheimia tenuiseta Geh. (Crosby et a1.,1983) End, Type - Ambatondrazaka,
herb (N.S.).
Schlotheimia trichophora Renauld et Cardot (Crosby et al., 1983) End, Type Entre Ankeramadinika et Ia for0t d'Analamazaotra, 1891, Cambou| &
Campenon,holotype PC.
Schlotheimia trichophora var. stenophyl/a (Renauld et Cardot) Thdr. (Crosby er
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, prds de Marotandrano, Rutenberg, holotype PC.
Ulota fulva Brid. (Crosby et al., 1983)
Zygodon laxifulius Th6r. (Crosby et a1.,1983) End, Type - Massif d'Andringitra,
alt. 1600 m,l92l, Perrier de la Bathie, holotype PC.
Zygodon leptocarpusThdr. (Crosby et a1.,1983)End, Type - Massif d'AndringiIra,
alt. 1600 m,I92l, Perrier de la Bathie, holotype PC.
Zygodon reinwardtii var. madacassus Cardot (Crosby et a1.,1983) End, Type NW Madagascar, Pervill1, holotype PC.
Zygodon reinwardtii (Hornsch.) A. Braun var. reinwardtii (Crosby et a1.,1983)
Phyllogoniaceae
Phyllogonium viscosum (P. Beauv.) Mitt. (Crosby et a1.,1983)
Pilotrichaceae
C allicostell a africana Mitt. (Crosby et al., 1983)
Callicostella brevipes (Broth.) Broth. (Crosby et a1.,1983)
Callicostella constricta (Mnlrl.Hal.) Kindb. (Crosby et a1.,1983)
Callicostella fissidentella (Besch.) Kindb. (Crosby et al., 1983)
Callicostella lacerans (Mtill.Hal.) A. Jaeger (Crosby et a1.,1983)
Callicostella leptocladula (Broth.) Broth. (Crosby et a1.,1983)
Callicostella papillatavar. brevifuIia M. Fleisch. (Crosby et a1.,1983)
Callicostella papillata (}/.ont.) Mitt. var. papillata (Crosby et a1.,1983; P6cs &
Geissler, 2002)
Callicostella parvocellulata Mitt. (Crosby et a1.,1983)
Callicostella petotii (Renauld et Cardot ex Paris) Broth. (Crosby et a1.,,1983)
Callicostella pusilla Demaret etP. de la Varde (Crosby et a1.,1983)
Callicostella seychellensis(Besch.) Kindb. (Crosby et a1.,1983; P6cs & Geissler,
2002)
Callicostella tristis (Mtill.Hal.) Broth. (P6cs& Geissler,2002)
Cyclodictyonaubertii (P. Beauv.)Kuntze (Crosbyet a1.,1983)
Cyclodictyon laetevirens(Hook. etTaylor) Mitt. (Crosbyet a1.,1983)
Cyclodictyon vallis-gratiae (Mill.Hal.) Kuntze (Crosby et al., 1983; P6cs &.
Geissler,2002)
Hookeriopsis divercifulia (Renauld et Cardot) Broth. ex Cardot in Grandid.
(Crosbyeta1.,1983)
Lepidopilidium attenuatum Thdr. (Crosby et al., 1983) End, Type - ForOt
d'Andasibe,sur I'Onive, 0211,925,
Perrier de la Bathie,holotype PC.
Checklist of the bryophytes of Madagascar
241
Lepidopilidium chenagonff (Renauld et Cardot) Cardot (Crosby et a1.,1983)End,
Type - Zone inf6rieure des forOts, montagne d'Ambre, prbs Di6goSuarez, Ch4nagon,herb (N.S.).
Lepidopilidium corbieri (Renauld et Cardot) Cardot (Crosby et al., 1983) End,
Type -Zone inf6rieure des forOts,montagne d'Ambre, Leloutre, holotype
CHE.
Lepidopilidium parvulum Cardot (Crosby et a1..1983) End, Type - NW, Pervill1,
h e rb (N .S .).
Lepidopilidium subrevolutum (Renauld et Cardot) Cardot (Crosby et a1.,1983)
Lepidopilum hircutum var. tuberculatumTh6r. (Crosby et a1.,1983) End, Type Andasibe, sur I'Onive, forOt )r sous-boisherbac€,,alt.800-1400 m,0211925,
Perrier de la Bathie, holotype PC.
Lepidopilum lastii Mitt. (Crosby et a1.,1983;Churchill, 1988)
Lepidopilum vetucipes Cardot (Crosby et a1.,1983;Churchill, 1988)
Polytrichaceae
Atrichum androgynum (Mnl.l.Hal.) A. Jaeger (Crosby et al., L983; De Sloover,
1e86)
Pogonatum belangeri (Mi.i11.Hal.) A. Jaeger (Hyvonen, L989; P6cs & Geissler,
2oo2)
Pogonatum capense (Hampe) A. Jaeger (Crosby et al., 1983; Hyvonen, L989;
De Sloover, L986;P6cs & Geissler,2002)
Pogonatum convolutum (Hedw.) P. Beauv. (Crosby et a1.,1983; Hyvdnen, 1989;
De Sloover, L986;P6cs & Geissler,2002)
Pogonatum gracilifulium Besch. (Crosby et a1.,,1983)
Polytrichastrumformosum (Hedw.) G.L. Sm. (Crosby et a1.,1983)
Polytrichum commune Hedw. var. commune (Crosby et al., 1983; De Sloover,
1e86)
Polytrichum glabrurn Schrad. (Crosby et a1.,1983)
Polytrichum juniperellum Geh. (Crosby et a1.,1983) End, Type - Plateau central,
Ambohimaro, Ambatomainty prds Tananarive, Rutenberg, herb (N.S.).
Polytrichum subformosum var. parvisetum (M{ill.Hal.) Cardot (Crosby et al.,
1983) End, Type - Plateau central, Imerina, bords du lac d'Ambohipo,
Cambou6, PC. Analamatnty, Campenon, PC, holotype (N.S.).
Polytrichum subpilosum P. Beauv. (Crosby et a|.,1983; De Sloover, 1986)
Pottiaceae
Anoectangium impressum Hampe (Crosby et al., 1983)
Anoectangium madagassum Renauld e/ Paris (Crosby et al., 1983) End, Type Pays des Betsileo,VillaLtme, herb (N.S.).
Anoectangium rufoviride Besch. (Crosby et a\.,1983)
Barbulq bolleana (Mi.i1l.Hal.)Broth. (Crosby et a|.,1983)
Barbulq eubryum Mi.ill.Hal. (O'Shea et a1.,1997)
Barbula indica (Hook.) Spreng. var. indica (Crosby et a1.,1983)
Barbula jananicu,s O'Shea (O'Shea, 2008) End, Type - Janjanina, carton de
Mahaiza, herb (N.S.).
Barbula omissa Th6r. (Sim & Dixon, 1922)
Barbula subobtusa Thdr. (Crosby et al., 1983) End, Type - Vondrozo, forOt sur
rocher humide au bord d'une cascade,lIl09lI924, Decary, holotype PC.
Bry oery throphy llum campy locarpum (MUll.Hal.) H. A. Crum (Sollman, 2009)
242
L.MarIine et al.
Chionoloma bomb¥se(M{ill.Hal.) P. Sollm. (Crosby et a1.,1983)
Didymodon subrevolutus (Hampe) Broth. (Crosby et a1.,1983) End, Type - 1869,
Borgen, holotype PC.
Gymnostomiella taberculosa (Renauld et Paris) Arts er P. Sollm. (Crosby et al.,
1e83)
Gymnostomum madagascariense O'Shea (O'Shea, 2008) End, Type - Desvaux,
herb (N.S.).
Hymenostylium subuispulum Thdr. (Crosby et al., 1983) End, Type - Mont
Tsaratanana, for€t d sous bois herbac6, alt. 1200-2400m,0411924, Perrier
de la Bathie, holotype PC#
Hyophila acuminata Broth. etP. de la Varde (Crosby et a1.,1983)
Hyophila angustifoliaParis e/ Renauld (Crosby et a1.,1983) End, Type - Rive de
la Mania, 08/L901-,Galinon, holotype PC.
Hyophila involata (Hook.) A. Jaeger (Crosby et a1.,1983)
IIyophila lanceolata Renauld et Cardot (Crosby et a1.,1983)
Hyophila parietalis Cardot (Crosby et a1.,1983) End, Type - Zone des savanes
occidentales,Marovoay, Mathieu, herb (N.S.).
Hyophila potieri var. atroviridis Renauld et Cardot (Crosby et aI., 1983) End,
Type - Plateau central, Ambohibeloma, Imerina, Cambou6, holotype
PC#
Hyophila sakalavensl's Paris e/ Renauld (Crosby et a1.,1983)
Leptodontium flexifulium (With.) Hampe (Sollman ,2006)
Leptodontium pungens (Mitt.) Kindb. (De Sloover, L987)
Leptodontium viticulosoides (P. Beauv.) Wijk et Margad. var. viticulosoides
(Zander, 1972;Crosby et aI., 1983;De Sloov er, 1987; Delgadill o, 1995)
Splachnobryum gracile Besch. (Crosby et a1.,1983)
Splachnobryum obtusum (Brid.) Miill.Hal. (Crosby et al., 1983; Arts, 1996; Arts,
200r)
StreptopogoncalymperesMi.ill.Hal.ex Geh. (Crosbyet a1.,1983)
Streptopogon erythrodontus (Taylor) Wilson (Crosby et al., 1983; Delgadillo,
lees)
Tortella humilis (Hedw.)Jenn.(Crosbyet a1.,1983)
Tortula madagassaDixon (Crosbyet a1.,1983)End, Type - unknown.
Tortula muralis var. madagascariensl's
Th6r. (Crosby et al., 1983) End, Type Andrahomana,sur rochergr6seux,Decary,herb (N.S.).
Tortula omissaTh6r. (Crosbyet a1.,1983)End, Type - Imerimandroso,sur 6corce,
associ6d,Bryum truncorumBrid. Decary,holotype PC.
Tortula subrufa Cardot (Crosby et al., 1983) End, Type - Fianarantsoa,1905,
Villaumes.n.,holotypePC, isotypePC.
TrichostomumbrachydontiumBruch var. brachydontium(Crosbyet a1.,1983)
Trichostomumpennequinii Renauld et Paris (Crosby et al., 1983) End, Type Cercle des Bara, secteurd'Ivondro, l-900,Lespagnolet Croll, holotype
PC.
Trichostomumpetieri Th6r. (Crosby et a1.,1983)End, Type - Massif granitique
de l'Andringitra,alt. 1600m,I92I, Perrierde la Bathie,holotypePC.
Trichostomum sporaphyllum (Renauld et Cardot) Cardot (Crosby et al., 1983)
End, Type - Ambohibeloma,1906,Cambou6,holotypePC.
Trichostomumtenuirostre(Hook. etTaylor) Lindb. var. tenuirostre(Crosby et al.,
1983)
Checklist of the bryophytes of Madagascar
243
Trichostomum villaumei Thdr. (Crosby et al., 1983) End, Type - Fianarantsoa,
1905, Villaume, holotype PC.
Weisiopsisplicata (Mitt.) Broth. (Crosby et a1.,1983)
Weissia ayresii Schimp. (Crosby et a1.,1983)
Prionodontaceae
Prionodon ciliatus Besch. (Crosby et a1.,1983)
Pterobryaceae
Calyptothecium bescherellei (Renauld) Broth. (Crosby et a1.,1983) Bnd, Type Mont Ankaratra, Borgen, holotype BM!
Calyptothecium planifron^s (Renauld er Paris) Argent (Argent, 1973: Crosby et al.,
1983;Delgadillo, 1995)
Hildebrandtiella endotrichelloides Miill.Hal. (Crosby et al., 1983)
Hildebrandtiellq longiseta Renauld et Cardot (Crosby et al., 1983) End, Type Di6go-Suarez, 1890, Ch€nagon,holotype PC.
Hildebrandtiella rotundifolia Renauld (Crosby et al., 1983)
Jaegerina solitaria var. nossi-beana Besch. (Crosby et a\.,1983) End, Type -Zone
inf6rieure des forOts,Nosy Be, Pervill€,herb (N.S.).
Jaegerina stolonifera (Mtill.Hal.) Mtill.Hal. subsp. stolonifera (Argent, 1973;
C ro sby et a1.,1983)
Orthostichidium involutifulium (Mitt.) Broth. subsp. involutifulium (Crosby
eta1.,,1983)
,
' .,:
Orthostichopsis longinervis (Renau\d et Cardot) Broth. (Crosby et a\.,1981; ::t'
,n.,
Orthostichopsis madagassa Dixon (Crosby et a1.,1983)End, Type - unknown.-d
Orthostichopsis scaberula (Renauld et Cardot) M. Fleisch (Crosby et al., 1983)
l(
End, Type - Zone moyenne des forOts,entre Vinanitelo et Ikongo, sur les
h
rameaux grOlesdes arbres,1892, Besson,holotype PC#
Orthostichopsis subimbricata (Hampe) Broth. (Argent, 1973; Crosby et a1.,1983)
Orthostichopsis subimbricata var. flageyi (Renauld et Cardot) Cardot (Crosby er
al., 1983) End, Type
Zone infdrieure des forOts, Di6go-Suarez,
Chenagon;Zone sup6rieure, Andrangoloaka, Sikora, herb (N.S.).
Pterobryopsis longissima Dixon (Crosby et a1.,1983)End, Type - unknown.
i
Renauldia hildebrandtielloides Mi.ill.Hal. (Argent,1973; Crosby et a1.,1983)r{-tr14_,'
Renauldia patentissima (Hampe) Broth. (Argent , 1973; Crosby et al., 1983) End,
Type -1892, Besson,holotype PC.
Renauldia patentissima var. mutica Th6r. (Crosby et al., 1983) End, Type Montagne d'Ambre, alt. 1000 m,911926, 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,
isolectotypesH, NY
Trachyphyllum inflexum (Harv.) A. Gepp (Buck, I9l9; Crosby et a1.,1983)
Ptychomitriaceae
Ptychomitrium crispatum (Hedw.) A. Jaeger (Crosby et a\.,1983)
Ptychomitrium deltorii (Ther.) Broth. (Crosby et a1.,1983)
244
L.Marline et al.
Pylaisiadelphaceae
Isopterygium ambreanum Renauld et Cardot (Crosby et al., 1983) Bnd, Type Di6go-Suarez, sur les 6corces,1890, Ch4nagon,holotype PC.
Isopterygium antsihqnaken^se Cardot (Crosby et al., 1983) Bnd, Type - Zone
moyenne des forOts,pays d'Antsihanaka, Perrot, herb (N.S.).
Isopterygium argillicola (Renauld et Cardot) Broth. (Crosby et al., 1983) End,
Type - Entre Tananarive et Betafo, CaussAque,holotype PC.
Isopterygium argyroleucum Besch. (Crosby et a1.,1983)
Isopterygium austrodenticulatum (Renauld et Cardot) Broth. (Crosby et a1.,1983)
End, Type - Andriba,189J, Dorr, holotype PC.
Isopterygium combaeBesch. (Crosby et a1.,1983)End, Type -Zone infdrieure des
forOts,Nosy Be et Nosy Komba, Marie, herb (N.S.).
Isopterygium gracile Renauld et Cardot (Crosby et a1.,1983)
Isopterygium intortum (P. Beauv.) A. Jaeger (Crosby et a|.,1983)
Isopterygium intortum var. chenagonii Renauld et Cardot (Crosby et al., 1983)
End, Type - Zone inf6rieure des forOts, Di6go-Suarez, Chdnagon, herb
(N.s.).
Isopterygium leptoblastum(Mtill.Hal.) A. Jaeger(Crosbyet al., 1983)
Isopterygiumluteonitens(Paris)Paris(Crosbyet a1.,1983)End, Type - Zone dl
Plateau central, bords gramineuxdu lac d'Ambohipo, Cambou6,herb
(N.s.).
Isopterygium meylanii Cardot (Crosby et a1.,1983) End, Type - Zone du Plateau
central, Imerina, Villaume, holotype G.
Isopterygium podomhizum Renauld et Cardot (Crosby et a1.,,1983) Bnd, Type Zone du Plateau central, Ambatomanga, dans un gazon d'Ectropothecium regulare, alt.1200-1300m, 1894, Talazac, herb (N.S.).
Isopterygium subleptoblasturm Miill.Hal. (Crosby et al., 1983)
Taxithelium argyrophyllum Renauld et Cardot (Crosby et a1.,1983) End, Type Zone inf6rieure des forOts, sur les troncs pourrissants h Di6go-Suarez,
Ch€nagon,holotype PC.
Taxithelium confusum Cardot (Crosby et a1.,1983) End, Type - Ile Sainte Marie,
Boivin, herb (N.S.).
Taxithelium glaucophyllum Besch. (Crosby et al., 1983) End, Type - Zone
inf6rieure des for0ts, Nosy Be, forOt de Lokobe, Marie, herb (N.S.).
Taxithelium hirtellumParis e/ Renauld (Crosby et a1.,1983)End, Type - Zone des
savanesorientales, Maintirano, Paris 66,holotype PC.
*Taxithelium nepalense (Schwiigr.) Broth. - For0t d'Ankara, Sambirano, sur sol,
sur calcaire, alt. 200 m,2110811977,Tixier 10746,TAN.
Taxithelium nossi&num Besch. (Crosby et a1.,1983)
Taxithelium planulum Besch. (Crosby et al., 1983)
Wijkia bessonii (Renauld et Cardot) H.A. Crum (Crosby et a1.,1983)Bnd, Type Zone moyenne des forets, plateau d'Ikongo, Besson,herb (N.S.).
Wijkia madagassc (Th6r.) H.A. Crum (Crosby et a1.,1983)End, Type -Val16e de
I'Anony, sous-bois,0510811921,
Decary, holotyp PC.
(Miill.Hal.)
rutenbergtt
H.A.
Wijkia
Crum (Crosby et al., 1983) End, Type Massif d'Ankaizina, herb (N.S.).
Wijkia rutenbergii var. eflagellaris (Cardot) H.A. Crum (Crosby et a1.,1983)
Checklist of the bryophytes of Madagascar
245
Racopilaceae
Powellia elliptica (Renauld) Zanten (Crosby et a1.,1983; Zanten,2003) End, Type
- Entre Fianarantsoa et Mananjary, 1880,Besson,holotype PC.
P o w eI I i a p o csii Zanten (Zanten, 2003)
Racopilum africanum MitL (Crosby et al., 1983)
Racopilum cardotii Renauld (Crosby et a1.,1983)End, Type -Zone infdrieure des
for0ts, Di6go-Suarez, 1890, Ch4nagon, PC; Andevoranto, 1,896,Perrot,
PC; holotype (N.S.).
Racopilum madag&ssumRenauld (Crosby et a1.,1983)
Racopilum madagassum var. grandistipulaceum Th6r. e/ Renauld (Crosby et al.,
1983) End, Type - Zone du Plateau central, pays Betsileo, Villaume,
holotype BM#
Racopilum marginatum Dixon (Crosby et a1.,1983)
Racopilum mauritianumMill.Hal. ex Besch. (Crosby et a1.,1983)
Racopilum microdictyon Besch. (Crosby et a1.,1983)
Racopilum penieri Thdr. (Crosby et a1.,1983)End, Type - Entre la Sahambana et
I'Ihosy, ForOt d'Analamanara, alt. 1300 m,0611919, Perrier de la Bathie,
holotype PC.
Racopilum plicatum Renauld et Cardot (Crosby et a1.,1983)
Racopilum tomentosum (Hedw.) Brid. (Crosby et a1.,1983)
Regmatodontaceae
Regmatodon orthostegius Mont. (Crosby et a1.,1983;Eakin, 1998)
Rhabdoweisiaceae
Oreoweisia erosa (Miill.Hal.) Kindb. (Crosby et a1.,1983)
Rhachitheciaceae
Rhachithecium perpusillum (Thwaites et Mitt.) Broth. (Crosby et a1.,1983)
Rhacocarpaceae
Rhacocarpus purpurascens (Brid.)
Delgadillo, 1995)
Paris (Crosby et al., 1983; Frahm, 1996;
Rhizogoniaceae
Pynhobryum spinifurme (Hedw.) Mitt. (Crosby et aI., 1983; Frahm et al., 2003;
Delgadillo, L995)
Rigodiaceae
Rigodium toxarion (Schwaegr.) A. Jaeger var. toxarion (Crosby et al., 1983;
Zomlefer, 1993)
Rutenbergiaceae
Rutenbergiq borbonica Besch. (Crosby et a1.,1983)
Rutenbergia cirrata Renauld et Cardot (Crosby et al., 1983) End, Type - ForOt
d'Analam azoatra, Camb ou6 295, holotype PC.
Rutenbergis limbata (Hampe) Besch. (Crosby et al., 1983) End, Type - ForOt
d'Analam azaotra,,B o r chgr evink, herb (N. S.).
Rutenbergia madagassa Geh. et Hampe (Crosby et al., 1983) End, Type
Ambaton dr azaka,herb (N.S.).
246
L.Marline et al.
Rutenbergia madagqssayar.flagellifurmis Renauld et Cardot (Crosby et a|.,1983)
End, Type
Zone du Plateau central, Ambohimahatsara prds
d'Ambos itr a, 1892, B erthieu, holotype PC.
Sematophyllaceae
Acanthomhynchium papillatum
rees)
(Harv.) M. Fleisch. (Crosby et al., 1983; Tixier,
Acroporium depressumThdr. (Crosby et a1.,1983)Bnd, Type - Mont Tsaratanana,
for6t ir sous-bois herbac€,, aIt.1200-2400 m, Perrier de la Bathie, holotype
PC.
Acroporium laevifuliurn (Renauld) Broth. (Crosby et al., 1983) End, Type Ankeramadinika, zone supdrieure des forOts, lIlI897, Dorr, holotype PC.
Acroporium megasporum (Duby) M. Fleisch. (Crosby et a1.,1983)
Acroporium megasporum var. densum (Renauld et Cardot) Wijk et Margad.
(Crosby et a|.,1983) End, Type - ForOt entre Fianarantsoa et Mananjary,
1892,Benson, holotype PC.
Acroporium stellatum (Renauld et Cardot) Broth. (Crosby et a1.,1983)End, Type
Zone inf6rieure des forOts, forOt e l'Est de Mahambo, environ
d'Andevorante, au milieu de touffes de Schlotheimia perroti,1890-1896,
Perrot, holotype PC.
Clastobryophilum bogoricurn (Bosch e/ Sande Lac.) M. Fleisch. (Crosby et al.,
1e83)
Macrohymenium acidodon (Mont.) Dozy e/ Molk. (Eakin, 1976; Crosby et al.,
1983;Een, 1997)
Macrohymenium muelleriDozy er Molk. (Eakin, 1976)
M acro hy m enium rufum Mi.ill. Hal. (Eakin, I97 6)
Macrohymenium strictum Bosch. e/ 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 forOts,forOt d'Analamazaotra, Cambou6, herb
(N.s.).
Radulina borbonica (Bel.) W.R. Buck (Crosby et a1.,1983)
Rhaphidothynchium qngustissimum (Geh.) Broth. (Crosby et al., 1983) End,
Type - ForOt de Vondrozona, Rutenberg, herb (N.S.).
Rhaphidothynchium cambouef (Renauld et Cardot) Broth. (Crosby et a1.,1983)
End, Type - Zone du Plateau central, Ankadivavala, Campenon, herb
(N.s.).
Rhaphidomhynchium rubricaule (Besch.) Broth. (Crosby et a1.,1983)
Sematophyllum coppeyi (Cardot) Broth. (Crosby et a1.,1983) End, Type - ForOt
de Manongarivo, 1909,Drouhard, holotype PC.
Sematophyllum dixonii (Thdr.) O'Shea (Crosby et a1.,1983) End, Type - Vall6e
de l'Anony, Imerimandroso, Ambatondrazaka, Amparafaravola, 300 km
NE, alt. 600 m, 0510811921,
Decary & Waterlot,holotype PC.
(Renauld)
Broth. (Crosby et al., 1983) End, Type Sematophyllum dubium
Ankeramadinika, II|I897, Dorr, holotype PC.
Sematophyllum nanopyxis (Geh.) O'Shea (Crosby et a1.,1983) End, Type - Zone
des for0ts, forOt de Vondrozona, Rutenberg,herb (N.S.).
Checklist of the bryophytes of Madagascar
247
Sematophyllum obtusifulium (Renatld et Cardot) Broth (Crosby et a1.,1983)
Sematophyllum rigescens(Cardot) O'Shea (Crosby et a1.,1983)End, Type -Zone
inf6rieure des forOts,forOt de Fito, district de Tamatave, Perrot, holotype
FH#
Sematophyllum sinuosulurn (Besch.) Broth. (Crosby et a1.,1983)
Sematophyllum sinuosulum var. Iatifulium (Cardot) Th6r. (Crosby et al., 1983)
End, Type - Zone du Plateau central, province d'Imamo, Cambou,4,
holotype G#
Sematophyllum subpinnatum (Brid.) E. Britton (Crosby et a1.,1983)
Sematophyllum subrugifulium Dixon (Crosby et a1.,1983)End, Type - unknown.
Sematophyllum subscabrellum Renauld et Cardot (Crosby et a1.,1983) End, Type
- Andevoranto, au milieu d'autres mousses,1895,Perrot, holotype PC.
Trichosteleum debettei var. laevisetum Cardot (Crosby et a1.,1983) End, Type unknown.
Trichosteleum heterophyllumBizot et Onr. (Crosby et a1.,1983)End, Type - Nord
Maromandia, au Sud de Di6go-Suarez, sur sol caillouteux, au bord d'un
marigot permanent, sous forOt, alt. 50 m, I3l0BlI973, Cremers 2735,
holotype BR, isotypes BR, TAN, PC.
Trichosteleum leviusculumRenauld et Cardot (Crosby et a1.,1983)
Trichosteleum microdontum (Besch.) Renauld (Crosby et a1.,1983)
Trichosteleum microdontum var. megapterun Renauld et Cardot (Crosby et al.,
1983) End, Type - Zone moyenne des for0ts, forOt d'Analamazaotra,
Cambout & Campenon,herb (N.S.).
Trichosteleum permixtum Cardot (Crosby et a1.,1983)End, Type - Didgo-Surarez,
Ch4nagon, holotype PC.
Trichosteleum percotii (Renauld et Cardot) Renauld et Cardot (Crosby et al.,
1e83)
perrotii var. eurydictyonRenauldet Cardot(Crosbyet al., 1983)
Trichosteleum
End, Type - Zone inf6rieure des for6ts, ile de Sainte-Marie, Mahavelona
ou Foulpointe, Darbould, herb (N.S.).
Trichosteleum pervilleanum (Mill.Hal. ex Geh.) W.R. Buck (Crosby et a\.,1983)
Trichosteleum trachypyxis (Geh.) Renauld et Cardot (Crosby et al., 1983) End,
Type
Zone moyenne des forOts, forOt a I'est d'Ambatondrazaka,
Rutenberg, herb (N.S.).
Warburgiella leptothyncha (A. Jaeger) Broth. (Crosby et a\.,1983)
Serpotortellaceae
Serpotortella chenagonii (Renauld et Cardot) W.D. Reese et R.H. Zander
(C rosby et a| .,1983)
Serpotortella cyrtophylla (Besch.) W.D. Reese e/ R.H. Zander (Reese & Zander,
1987)
Sphagnaceae
Sphagnum c&penseHornsch. (Eddy, 1985)
Sphagnum ceylonicum Mitt. ex Warnst. (Eddy, 1985)
Sphagnum cuspidatulum MUll.Hal. (Eddy, 1985)
Sphagnum cuspidatum Hoffm. var. cuspidaturn (Eddy, 1985)
*Sphagnum dqvidii Warnst. - Massif de I'Ankaratra, S L9'20' E 47"20', alt.2700 m,
2411011972,
M.R. & C.A. Crosby 6636,TAN.
L.MarIine et al.
Sphagnum ericetorum Brid (Eddy, 1985)
Sphagnum magellanicum (Warnst.) A. Eddy subsp.grandirete (Bddy,1985) End,
Type - Andevoranto, Androrangobe, 1901,,holotype PC.
Sphagnum perichaetiale Hampe (Eddy, 1985)
Sphagnum peruieri Th6r. (Crosby et a1.,1983)End, Type - Massif de l'Andringitra,
alt. 1600-2500m,t92I, Perrier de la Bathie, holotype PC.
Sphagnum planifuliumvar. angustilimbatunr (Warnst.) A. Eddy (Eddy, 1985)
Sphagnum planifulium Mi.ill.Hal. var. planifulium (Eddy, 1985)
Sph agnum rut enbergii Mi.ill. Hal. (Eddy, 1985)
Sphagnum strictum (Miill.Hal.) A. Eddy subsp.psppeanum (Eddy, 1-985)
Sphagnum truncatum var. bordasii (Besch.) A. Eddy (Eddy, 1985)
Sphagnum tumidulum Besch. var. tumidulum (Eddy, 1985)
Sphagnum violascens Miill.Hal. (Eddy, 1985)
Splachnaceae
Brachymitrion moritzianum (Mlull.Hal.) A.K. Kop. (Koponen & Weber, 1972;
C rosby et a1.,1983)
Tayloria indica Mitt. (Crosby et a1.,1983)
Tayloria isleanq (Besch.) Broth. (Crosby et a1.,1983)
Tayloria orthodonta (P.Beauv.) Wijk et Margad. (Crosby et al., 1983; P6cs &
Geissler,2002)
Tayloria solitaria (Hedw.) T.J. Kop et W. Weber (Crosby et al., 1983; P6cs &
Geissler,2002)
Stereophyllaceae
Entodontopsis nitens (Mitt.) W.R. Buck etheland (Crosby et a\.,1983)
Entodontopsis tenuinervis (Broth.) W.R.Buck etheland (Crosby et a1.,1983)
Stereophyllum radiculosum (Hook.) Mitt. (Crosby et a|.,1983)
Symphyodontaceae
Symphyodon pygmaeus (Broth.) S. He er Snider (Crosby et al.,t9B3; He & Snider,
2000)
Thuidiaceae
Pelekium chenagon t (Mtill.Hal. ex Renauld et Cardot) Touw (Touw, I976; Crosby
eta|.,1983)
Pelekium gratum (P. Beauv.) Touw (Touw, 1976; Crosby et a1.,1983;Touw, 200I)
Pelekium intricatum (A. Jaeger) Touw (P6cs & Geissler,2002)
Pelekium investe (Mitt.) Touw (Touw, 1976;Crosby et a1.,1983;Touw, 2001.;P6cs
& Geissler,Z}12)
Pelekium ramusculosum (Mitt.) Touw (Touw, 1976;Crosby et a1.,1983)
Pelekium varians (Welw. er Duby) Touw (Touw, 1976;Crosby et a1.,1983)
Pelekium versicolor (MUll.Hal.) Touw (Touw, 1976; Crosby et al., 1983; Touw,
20or)
Ruuiella praelonga (Schimp.exBesch.)Wijk etMargad.(Touw,1976;Crosbyer
a1.,1983)
et Cardot (Touw, 1976;Crosbyet a1.,1983)
Thuidium aculeoserratumRenauld
Thuidium ussimile(Mitt.) A. Jaeger(Touw, 1976;Crosbyet a1.,1983)
(Mtill.Hal.) A.Jaeger (Touw, I976; Crosbyet a|.,1983)
Thuidiumpseudoinvolvens
249
Checklist of the bryophytes of Madagascar
Note on Riccardia:
According to GrolIe 1995, three species of the genus Aneura: A. caespitans Steph.,
A corbierl 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 1,144 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 R6union Island, a landmass both a fraction of the age and size of Madagascar
(Ah-Peng et a1.,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(Schwiigr.) 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 I8.97o/" 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
Taxa
751
7744
Genera
186
275
91,
Families
I
Hornworts
Endemics
Endemicity (%)
254
74
328
33.82"/"
18.97"/"
28.67"/"
250
L.Marline et al.
families with the largest number of endemic species are the Dicranaceae (41183),
Orthotrichaceae (38/66) and Sematophyllaceae (22140). Of the 74 endemic
liverwort speciesrecorded 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 (P6cs, 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 L800'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
Foundationfor Researchin Biodiversity),FEDER POE 3.21,MOVECLIM (ANR Net
Biome) for financialsupport.The last auihor thanksthe ClaudeLeon Harris Fellowshipfor
supportingher research.
This checklist would not have been possiblewithout the work and support of
Brian O'Sheaand Martin Wigginton,particularlyfor their checklistsof bryophytesfor subSaharanAfrica.
We are grateful to the MalagasyNational Park for the issuingof collecting and
export permits (No 253/09/MEF/SG/DGF/DCBSAP/SLRSE)
and also to StevenGoodman
and Roger Edmond for their support and encouragementto work on the bryophyte flora
of Madagascar.We thank Ida Bruggeman-Nannengafor the identification of two new
recordsof Fissidensfor Madaqascar.
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