S. Rob Gradstein

Institute of Plant Sciences, University of Göttingen, 37073 Göttingen, Germany



Barbara M. Thiers,

New York Botanical Garden, Bronx, NY 10458-5126, U.S.A.


The hepatic flora of tropical America is still very inadequately studied. There is no Flora treating the species of any single region in the Neotropics. Our goal is to write a concise Flora of the Hepaticae and Anthocerotae of Central French Guiana. The present catalogue is the first step in that direction.

Central French Guiana is probably one of the richest lowland rain forest areas in the world for hepatics (liverworts, hornworts). An inventory of 28 trees in a few hectares of rain forest near Saül (Gradstein et al. 1990, Montfoort & Ek 1990) yielded 88 different species, being the highest number of hepatics recorded of any piece of lowland rain forest worldwide. The collections were especially representative because the inventory included detailed analysis of the forest canopy, where at least 50% of the species of hepatics occur.

The present catalogue is based on identified material kept in the herbarium of the University of Utrecht (on long-term loan in Göttingen) gathered by G. Cremers, ORSTOM, around Saül and on Mt. Galbao (Onraedt & Cremers 1980) and by Gradstein and students during May-October 1986 and in 1987 in the same area. Further collections from the area in NY (leg. Buck, Thiers, a.o.) are still in the process of identification and have not yet been included here.

In all, 172 species of hepatics (15 families, 59 genera) and 2 species of hornworts (2 genera, 2 families) are reported from Central French Guiana. Together they constitute more than half of the lowland hepatic flora of northeastern South America (French Guiana, Suriname, Guyana, below 1000 m; see Gradstein & Hekking 1989, Gradstein 1997). The most important hepatic family is the Lejeuneaceae, which include 123 species (in 40 genera) or about 75% of the entire hepatic flora of Central French Guiana. Of next importance are the Plagiochilaceae (9 spp.), Jubulaceae (8 spp.), Lepidoziaceae (6 spp.) and Radulaceae (6 spp.). The remaining twelve families have less than five species; seven families, including the two hornwort families Dendrocerotaceae and Notothyladaceae, are represented by only 1 species each.

Since the available collections are based on limited sampling around Saül and on Mt. Galbao, further exploration in Central French Guiana is necessary to assess the biodiversity of the liverworts and hornworts of the area. Fieldwork should include careful analysis of the forest canopy and of epiphylls, which were incompletely studied and may yield many unique and possibly novel taxa. We estimate that a minimum of 3-4 expeditions of about one month will be necessary to adequately explore the hepatic flora of the region.

The Flora of the Hepaticae and Anthocerotae of Central French Guiana shall be written by S.R. Gradstein & B.M. Thiers as a joint effort. Thiers will treat Lepidoziaceae, Radulaceae and Lejeuneaceae subfam. Cololejeuneoideae, Gradstein the remaining groups. Illustrations will be made at NY under supervision of Thiers and shall include a plate for one species per genus with selected diagnostic details of additional species.


Alphabetical list of families, genera and species of Hepaticae and Anthocerotae

recorded from Central French Guiana

Taxa followed by an * are species found only on Mount Galbao (ca. 650 m, near Sal)





Aneura Dumortier

Aneura sp.

Riccardia S.F. Gray

R. amazonica (Spruce) Schiffner


Calypogeia Raddi

C. lechleri (Stephani) Stephani *
C. miquelii Montagne


Odontoschisma (Dumortier) Dumortier

O. falcifolium Stephani (det. Vána)


Cylindrocolea Schuster
C. planifolia (Stephani) Schuster


Leptoscyphus Mitten

L. porphyrius (Nees) Grolle

Lophocolea (Dumortier) Dumortier

L. bidentata (Linnaeus) Dumortier
L. liebmanniana Gottsche
L. martiana Nees


Herbertus S.F. Gray

H. divergens (Stephani) Herzog (Onraedt 1980 as H. juniperoidea (Swartz) Grolle) *

JUBULACEAE (=Frullaniaceae)

Frullania Raddi

F. apiculata (Reinwardt et al.) Dumortier (= F. exilis Taylor)

F. arecae (Sprengel) Gottsche or F. ecklonii (Spreng.) Spreng. (check Yuzawa, J. Hattori Bot. Lab. 70: 259-271. 1991)

F. caulisequa (Nees) (= F. gymnotis Nees & Montagne = F. obcordata (Lehmann & Lindenberg) Lehmann & Lindenberg)

F. ericoides (Nees) Montagne
F. gibbosa Nees
F. kunzei (Lehmann & Lindenberg) Lehmann & Lindenberg
F. nodulosa (Reinwardt et al.) Nees
F. riojaneirensis (Raddi) Aongstroem


Acanthocoleus Schuster (see Gradstein 1994)

A. aberrans (Lindenberg & Gottsche) Kruijt

Acrolejeunea (Spruce) Schiffner (see Gradstein 1994)

A. torulosa (Lehmann & Lindenberg) Schiffner

Anoplolejeunea (Spruce) Schiffner

A. conferta (Meissner) Schiffner *

Amphilejeunea Schuster

A. reflexistipula (Lehmann & Lindenberg) Gradstein, Mem. N.Y. Bot. Garden 86: in press (= L. reflexistipula (Lehmann & Lindenberg) Gottsche et al.)

Aphanolejeunea Evans (see Pócs & Bernecker-Lücking in prep.)

A. exigua Evans (det. Tixier) *
A. gracilis Jovet-Ast
A. sicaefolia (Gottsche ex Stephani) Evans

Archilejeunea (Spruce) Schiffner

A. auberiana (Montagne) Evans
A. fuscescens (Hampe ex Lehmann) Fulford
A. parviflora (Nees) Schiffner

Brachiolejeunea (Spruce) Schiffner (see Gradstein 1994)

B. conduplicata (Stephani) Gradstein

Bryopteris Nees (see Gradstein 1994)

B. filicina (Swartz) Nees

Ceratolejeunea (Spruce) Schiffner (Dauphin in prep.)

C. cubensis (Montagne) Schiffner
C. confusa Schuster
C. cornuta (Lindenberg) Schiffner (=C. maritima (Spruce) Stephani, fide G. Dauphin in prep.)

(C. grandibracteata Fulford, det. Onraedt = ? see Dauphin in prep)

C. guianensis (Montagne & Nees) Stephani
C. megalophysa (Spruce) Stephani
C. patentissima (Hampe & Gottsche) Evans (identification probably erroneous)
C. plumula (Spruce) Stephani

(C. valida Evans, det. Tixier, = ? see Dauphin in prep.) *

Cheilolejeunea (Spruce) Schiffner

C. adnata (Kunze) Grolle

C. clausa (Nees & Montagne) Schuster (good species, not a synonym of C. trifaria, see Reiner-Drehwald, Tropical Bryology 14: 55. 1998)

C. inflexa (Hampe ex Lehmann) Grolle *
C. rigidula (Montagne & Nees) Schuster
C. surinamensis (Stephani) X.-L. He, Ann. Bot. Fenn. 34: 1997 (= Trachylejeunea?, lobule teeth paired!)
C. trifaria (Reinwardt et al.) Mizutani

Cololejeunea (Spruce) Schiffner (see Tixier & Pócs in prep.)

C. apressa (Evans) Benedix
C. bekkeri Tixier
C. cardiocarpa Montagne

?C. linopteroides H. Robinson (identified by Tixier as C. sigmoidea Jovet-Ast & Tixier, which is Asiatic and is replaced in the Neotropics by C. linopteroides, see Bernecker-Lücking & Morales, The Bryologist 102: 437-441. 1999]

C. obliqua (Nees & Montagne) Schiffner (= C. scabriflora Gottsche ex Stephani)
C. platyneura (Spruce) S. Arnell
C. subcardiocarpa Tixier
C. subscariosa (Spruce) Stephani

Colura (Dumortier) Dumortier

C. clavigera Gottsche ex Jovet-Ast
C. cylindrica Herzog
C. tortifolia (Nees & Montagne) Trevisan

(Crossotolejeunea = Lejeunea subgen. Crossotolejeunea)

Cyclolejeunea Evans (see Bernecker- Lücking 1998)

C. angulata Herzog (Onraedt 1998, check identification)
C. convexistipa (Lehmann & Lindenberg) Evans
C. luteola (Spruce) Grolle

C. peruviana (Lehmann & Lindenberg) Evans

Cyrtolejeunea Evans

C. holostipa (Spruce) Evans

Diplasiolejeunea (Spruce) Schiffner

D. brunnea Stephani (det. Tixier) *
D. cavifolia Stephani (= D. brachyclada Evans)
D. pellucida (Meissner ex Sprengel) Schiffner
D. rudolphiana Stephani

Drepanolejeunea (Spruce) Schiffner

D. bidens Stephani *
D. crucianella (Taylor) Evans
D. fragilis Bischler
D. inchoata (Meissner) Stephani *
D. lichenicola (Spruce) Stephani *
D. mosenii (Stephani) Bischler (Onraedt & Cremers 1980)
D. orthophylla (Nees & Montagne) Bischler
D. pinnatiloba Schiffner (det. Onraedt)
D. spinosa Herzog *

Echinocolea Schuster

E. asperrima (Spruce) Schuster

Frullanoides Raddi (see Gradstein 1994)

F. tristis (Stephani) van Slageren

Harpalejeunea (Spruce) Schiffner

H. oxyphylla (Nees & Montagne) Stephani (see Grolle & Reiner-Drehwald, J. Bryology 21: 38. 1999) *
H. stricta (Lindenberg & Gottsche) Stephani (check identification!)

H. tridens Bescherelle & Spruce (= H. heterodonta Evans, see Grolle & Reiner-Drehwald, J. Bryology 21: 40. 1999) *

H. uncinata Stephani (check identification!)

Lejeunea Lib.

L. boryana Montagne (= Crossotolejeunea boryana) (see Reiner-Drehwald, Bryophyt. Biblioth. 54: 22. 1999)
L. brittoniae (Evans) Grolle, Bryophyt.. Biblioth. 48: 81. 1995 (= Rectolejeunea brittoniae Evans)
L. caespitosa Lindenberg
L. flava (Swartz) Nees
L. laetevirens Montagne & Nees

Lejeunea phyllobola Nees & Montagne (= Rectolejeunea phyllobola (Nees & Montagne) Evans) - species with true flagellae! (see Grolle, Bryophyt. Biblioth. 48: 16. 1995)

(L. reflexistipula (Lehmann & Lindenberg) Gottsche et al. = Amphilejeunea reflexistipula (Lehmann & Lindenberg) Gradstein)

Lepidolejeunea Schuster

L. involuta (Gottsche) Grolle

Leptolejeunea (Spruce) Schiffner

L. elliptica (Lehmann & Lindenberg) Schiffner
L. exocella (Spruce) Evans
L. foraminulosa Stephani (det. Tixier) *
L. moniliata Stephani
L. cf. serratifolia Schiffner

Leucolejeunea Evans

L. unciloba (Lindenberg) Evans

Lopholejeunea (Spruce) Schiffner (see Gradstein 1994)

L. quelchii Stephani (= L. howei Evans) (det. Tixier) *
L. nigricans (Lindenberg) Schiffner (= L. muelleriana (Gottsche) Schiffner) *
L. subfusca (Nees) Schiffner

Marchesinia S.F. Gray (see Gradstein 1994)

M. brachiata (Swartz) Schiffner

Mastigolejeunea (Spruce) Schiffn. (see Gradstein 1994)

M. auriculata (Wilson) Schiffner
M. innovans (Spruce) Schiffner
M. plicatiflora (Spruce) Schiffner

Metalejeunea Grolle

M. cucullata (Reinwardt et al.) Grolle, Bryophyt. Biblioth. 48: 100. 1995 (=

Microlejeunea monoica Bischler)

Microlejeunea Stephani

M. acutifolia Stephani
M. aspera Stephani (det. Tixier) *
M. bullata (Taylor) Stephani *
M. crenulifolia (Gottsche) Stephani
M. epiphylla Bischler

(M. monoica Bischler = Metalejeunea cucullata (Reinwardt et al.) Grolle, Bryophyt. Biblioth. 48: 100. 1995)

Neurolejeunea (Spruce) Schiffner (see Gradstein 1994)

N. breutelii (Gottsche) Evans
N. seminervis (Spruce) Schiffner

Odontolejeunea (Spruce) Schiffner (see Gradstein 1994)

O. decemdentata (Spruce) Stephani
O. lunulata (Weber) Schiffner

O. rhomalea (Spruce) Stephani

Pictolejeunea Grolle

P. picta (Gottsche ex Stephani) Grolle

Prionolejeunea (Spruce) Schiffner

P. aemula (Gottsche) Evans *
P. macrocardia (Spruce) Stephani

Pycnolejeunea (Spruce) Schiffner (see X.-L. He 1999)

P. contigua (Nees) Grolle
P. macroloba (Nees & Montagne) Schiffner

(P. surinamensis Stephani = Cheilolejeunea surinamensis (Stephani) X.-L. He, Ann. Bot. Fenn. 34: 1997 = Trachylejeunea cf. pandurantha?)

Rectolejeunea Evans

R. berteroana (Gottsche) Evans
R. emarginuliflora (Gottsche ex Schiffner) Evans
R. flagelliformis Evans

Schiffneriolejeunea Verdoorn (see Gradstein 1994)

S. amazonica Gradstein

Stictolejeunea (Spruce) Schiffner (see Gradstein 1994)

S. balfourii (Mitten) E.W. Jones var. bekkeri Gradstein
S. squamata (Willdenow) Schiffner

Symbiezidium Trevisan (see Gradstein 1994)

S. barbiflorum (Lindenberg & Gottsche) Evans
S. transversale (Swartz) Trevisan

Taxilejeunea (Spruce) Schiffner

T. debilis (Lehmann & Lindenberg) Stephani
T. fissistipula Stephani (det. Tixier) (probably synonym) *
T. aff. sordida (Nees) Eifrig (Asiatic species!)
T. sulphurea (Lehmann & Lindenberg) Schiffner
T. surinamensis (Lindenberg & Gottsche) Stephani (Onraedt & Cremers 1980)

Thysananthus Lindenberg (see Gradstein 1994)

T. amazonicus (Spruce) Schiffner

Trachylejeunea (Spruce) Schiffner

T. crenata (Nees & Montagne) Schuster. J. Hattori Bot. Lab. 72: 253. 1992 (det. Onraedt) (= Crossotolejeunea crenata)

T. pandurantha (Spruce) Stephani
T. spec. (? see Cheilolejeunea surinamensis)


Arachniopsis Spruce

A. diacantha (Montagne) Howe

Bazzania S.F. Gray

B. bidens (Nees) Trevisan
B. breuteliana (Lindenberg & Gottsche) Trevisan
B. hookeri (Lindenberg) Trevisan *

Micropterygium Lindenberg

M. trachyphyllum Reimers

Telaranea Spruce ex Schiffner

T. nematodes (Gottsche ex Austin) Howe


Metzgeria Raddi (see Pinheiro da Costa 1999)

M. albinea Spruce *
M. leptoneura Spruce *
M. thomeensis Stephani (?, check identification) *


Symphyogyna Nees & Montagne

S. brogniartii Montagne *


Plagiochila (Dumortier) Dumortier

P. contigua (Nees) Grolle

P. disticha Lehmann & Lindenberg (=P. orbigniana Nees & Montagne, fide Heinrichs & Gradstein, Nova Hedwigia, in press)

P. laetevirens Lindenberg (= P. hondurensis Herzog)
P. macrifolia Taylor (probably synonym)

P. montagnei Nees (= P. hypnoides (Willdenow) Lindenberg, fide Heinrichs & Gradstein, Nova Hedwigia, in press)

P. perrotettiana Montagne (probably synonym)
P. pittieri Stephani (probably synonym)
P. subplana Lindenb. (= P. amazonica Spruce, fide Heinrichs et al., Haussknechtia Beiheft 9; in press)
P. tenuis Lindenberg


Radula Dumortier

R. cf. antilleana Castle (det. Yamada)

R. macrostachya Lindenberg & Gottsche (= R. caldana Ångström = R. longifolia Stephani = R. surinamensis Stephani, see Uribe & Gradstein 1998)

R. flaccida Lindenberg & Gottsche
R. husnotii Castle (det. Yamada)
R. kegelii Gottsche ex Stephani
R. stenocalyx Montagne (Onraedt & Cremers 1980)


Riccia Linnaeus

R. subplana Stephani (det. Jovet-Ast)


Trichocolea Dumortier

T. tomentosa (Swartz) Gottsche *
T. sprucei Steph. (Onraedt & Cremers 1980; check identification) *




Dendroceros Nees

D. crispus (Swartz) Nees *


Notothylas Sullivant

N. breutelii (Gottsche) Gottsche


For help with identifications we thank R. Grolle (Pycnolejeunea), S. Jovet-Ast (Colura, Riccia), M. Onraedt (selected Lejeuneoideae), P. Tixier (miscellaneous species), J. Vána (Odontoschisma) and K.Yamada (Radula).



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Gradstein, S. R. 1994. Lejeuneaceae: Ptychantheae, Brachiolejeuneae. Flora Neotropica Monograph 62: 1-216.

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