Journal of Bryology (2008) 30: 159–177
Bryological Notes
Cheilolejeunea cedercreutzii (H.Buch & Perss.) Grolle on Madeira
Island
Grolle & Long (2000) list 23 species of Lejeuneaceae for
Europe and Macaronesia (24 species if L. holtii Spruce is
considered distinct from L. eckloniana Lindenb.). However,
most of these species have a very restricted distribution in
the oceanic parts of Western Europe and in Macaronesia
(Azores, Madeira and Canary Islands). The genus
Cheilolejeunea has been reported within these territories
only from the Azores, where a single species C. cedercreutzii
(H.Buch & Perss.) Grolle is known (Buch & Persson, 1941;
Allorge & Allorge, 1950; Sjögren, 1978; Schumacker, 2001;
Sjögren, 2001; Gabriel et al., 2005). The genus has not been
reported from any of the other Macaronesian archipelagos
(Söderström, Urmi & Váňa, 2002; 2007).
In this paper, Cheilolejeunea is reported for the first time
from Madeira. The collection agrees well with material from
the Azores and it is here referred to C. cedercreutzii. Some
features of the plant, including oil-bodies, are illustrated.
Cheilolejeunea cedercreutzii (Fig. 1) is easily distinguished
from all other Lejeuneaceae of Europe and Macaronesia,
and is characterized as follows:
underleaves broadly oval to suborbicular, 1.4–2 6 width
of stem, bilobed to 1/4, with acute sinus and subacute lobes
lobules strongly inflated, forming distinct angle with the
keel, subtubular at the apex, the free margin incurved
apical tooth of lobule single, to 30 mm long, with distal
hyaline papilla
leaf cells with thickened walls and large, sometimes
bulging trigones
oil-bodies large, compound, solitary, rarely 2, per cell
perianths 5-carinate
subfloral innovations pycnolejeuneoid (the first leaf at
the base of the innovation is an underleaf).
The shoots are small, to ca 0.7 mm wide (wider at the
gynoecia), opaque and dark green in color. The leaf lobes
are obtuse or broadly pointed and the apex is decurved. The
cells of the lobe are prominent and rounded on the dorsal
surface, more rarely submamillose, giving the keel a more
or less crenulate appearance in profile (Fig. 1f). The oilbodies persist for a few weeks in dried specimens but are
soon lost; they are almost invariably solitary (exceptionally
2 per cell) and fill more than half of the cell lumen (Fig. 1d).
After they have disintegrated, the cell contents appear
darkly opaque and granular.
N
N
N
N
N
N
N
# British Bryological Society 2008
DOI: 10.1179/174328208X282472
Most of the gynoecia in the specimen from Madeira lack
mature perianths, and the few perianths observed were not
in good condition. However, they are clearly 5-carinate. No
androecia were detected, and this suggests that the species is
dioicous, as indicated by Buch & Persson (1941) in the
original description.
The Madeiran collection has been compared with plants
from Terceira (Azores) collected by Dr J.W. Bates in 1994,
and no significant differences could be found.
Pycnolejeuneoid subfloral innovations were noted in the
Azores material. However, oil-bodies had disintegrated in
the Azores material and could not be compared.
In the inflated lobules forming a distinct angle with the
keel, C. cedercreutzii has a superficial resemblance to
Harpalejeunea molleri (Steph.) Grolle and Lejeunea patens
Lindb. H. molleri is a slightly smaller plant, with
conspicuous ocelli near the base of the leaf lobes; it has
2–6 (occasionally more) oil-bodies per cell, and the underleaves are very different, being retuse or shallowly bilobed
with widely spreading broadly rounded lobes. Lejeunea
patens is pale to whitish green, commonly has 2–8 oilbodies per cell, and the underleaves are bilobed to half of
their length.
The taxonomic status of C. cedercreutzii as an independent species is outside the scope of this note and remains
uncertain. A full assessment is hardly possible without a
wide-ranging revision of both neotropical and palaeotropical species. Grolle (1983) noted that it is closely related to
the African C. usambarana (Steph.) Grolle, with which it
shares pycnolejeuneoid subfloral innovations. However C.
cedercreutzii differs in its apparently dioicous sexual
condition. According to Wigginton (2004), C. usambarana
is autoicous, although this is not always readily apparent as
the androecia are often remote from the gynoecia. C.
cedercreutzii also has many points in common with C.
oncophylla (Ångstr.) Grolle & Reiner from the Neotropics.
However, that species too is autoicous, and as described by
Grolle & Reiner-Drehwald (1997), it has more deeply lobed
underleaves (to 40% of the length) and more strongly
mamillose lobe cells.
The locality for C. cedercreutzii is to the east of the
Encumeada Pass on the north side of Pico Topeiro at an
altitude of ca 1150 m. The sample was collected from a
Received 29 January 2008. Revision accepted 17 March 2008
160
BRYOLOGICAL NOTES
Figure 1. Cheilolejeunea cedercreutzii from Madeira. (a) Vegetative shoot, ventral view; (b) underleaf; (c) single leaf, ventral view; (d) median
cells of leaf lobe, with oil-bodies; (e) detail of lobule apex, flattened, showing apical tooth and distal papilla; (f) detail of leaf keel. Drawn by
M.C. Watling from Blockeel 36/102.
loose piece of bark on a moribund Erica bush in tall heath
vegetation. It was closely associated with Harpalejeunea
molleri and sparsely with Microlejeunea ulicina (Taylor)
A.Evans, Lejeunea patens Lindb. and Frullania teneriffae
(F.Weber) Nees. The northern slopes of the mountains of
Madeira at this altitude are very humid with frequent
accumulations of cloud. Myurium hochstetteri (Schimp.)
Kindb. occurs on the crags near the Cheilolejeunea site, and
the endemic Tylimanthus madeirensis Grolle & Perss. on
humus-covered ledges.
Sjögren (1978) indicates that, in the Azores, C. cedercreutzii is confined to habitats where air humidity almost
permanently exceeds 80%. He notes its occurrence on moist
old pieces of bark of Juniperus brevifolia (Seub.) Antoine, a
habitat very similar to that of the new site on Madeira.
Although in a later paper Sjögren (2003) comments that it
is ‘almost only epiphyllous’ in the Azores, it is certainly
not confined to this habitat, occurring also as an epiphyte
and in rock crevices. Schumacker (2001) states that it is
‘not rare at all’ on the Azorean islands on which it is
known, being overlooked because of its similarity to other
small Lejeuneaceae. It is likely that it will be found at
additional localities on Madeira, and probably on different
substrates.
BRYOLOGICAL NOTES
Cheilolejeunea cedercreutzii (H.Buch & Perss.) Grolle,
Feddes Repertorium 87: 188. 1976 [Euosmolejeunea cedercreutzii H.Buch & Perss., Commentationes Biologicae 8 (7):
9. 1941].
MADEIRA: on bark of moribund Erica, north side of Pico
Topeiro, to east of Encumeada, ca 1150 m a.s.l., 8 April
2007, leg. T.L. Blockeel no. 36/102 (LISU, duplicate in Hb.
Blockeel).
AZORES: Terceira: in damp crevice of lava field, Misterio
Negro, 10 km N.W. of Angra do Heroismo, UTM 75–87-,
ca 650 m a.s.l., 15 July 1994, leg. J.W. Bates & R. Gabriel,
no. 3498 (Hb. Bates); Terceira: on Laurus bark in steep
forest, Terra Brava, about 1 km N.E. of Algar do Carvão,
UTM 82–87-, ca 640 m a.s.l, 21 July 1994, leg. J.W. Bates &
R. Gabriel, no. 3650 (Hb. Bates); Terceira: epiphyte on
Juniperus in upper caldeira forest, Juncal, about 11 km N.
of Angra do Heroismo, 22 July 1994, leg. J.W. Bates
no. 3682 (Hb. Bates).
I am very grateful to Malcolm Watling for preparation of the
plate, and to Des Callaghan for photographic assistance.
Thanks are also due to Dr J.W. Bates for the loan of his
collections of C. cedercreutzii, and to Dr D.G. Long and Dr
R. Schumacker for assistance with the literature.
TAXONOMIC ADDITIONS AND CHANGES: Nil.
REFERENCES
Allorge V, Allorge P. 1950. Hépatiques récoltées aux Îles Açores. Revue
Bryologique Lichénoligique 19: 90–118.
Buch H, Persson. 1941. Bryophyten von den Azoren und Madeira.
Societas Scientarum Fennica, Commentationes Biologicae 8: 1–15.
161
Gabriel R, Sjögren E, Schumacker R, Sérgio C, Frahm J-P, Sousa E.
2005. Lista dos Briófitos (Bryophyta)/List of Bryophytes
(Bryophyta). In: Borges PAV, Cunha R, Gabriel R, Martins AF,
Silva L, Vieira V, eds. Listagem da Fauna (Mollusca e Arthropoda)
e Flora (Bryophyta, Pteridophyta e Spermatophyta) Terrestres dos
Açores/A list of terrestrial fauna (Mollusca and Arthropoda) and
flora (Bryophyta, Pteridophyta and Spermatophyta) from the
Azores. Horta, Angra do Heroı́smo and Ponta Delgada:
Direcção Regional do Ambiente and Universidade dos Açores,
pp. 117–129.
Grolle R. 1983. Hepatics of Europe including the Azores: an annotated
list of species, with synonyms from the recent literature. Journal of
Bryology 12: 403–459.
Grolle R, Long DG. 2000. An annotated check-list of the Hepaticae and
Anthocerotae of Europe and Macaronesia. Journal of Bryology 22:
103–140.
Grolle R, Reiner-Drehwald ME. 1997. Cheilolejeunea oncophylla
(Ångstr.) Grolle & Reiner comb. nov. (Lejeuneaceae), from the
Neotropics. Journal of Bryology 19: 781–785.
Schumacker R. 2001. The hepatic flora of the Azores: brief historical
outline, present knowledge, endemics and phytogeographical
aspects. Belgian Journal of Botany 134: 51–63.
Sjögren E. 1978. Bryophyte vegetation in the Azores Islands. Memórias
da Sociedade Broteriana 26: 5–283.
Sjögren E. 2001. Distribution of Azorean bryophytes up to 1999, their
island distribution and information on their presence elsewhere,
including Madeira and the Canary Islands. Boletim do Museu
Municipal do Funchal 7: 1–89.
Sjögren E. 2003. Azorean bryophyte communities – a revision of
differential species. Arquipélago, Life and Marine Sciences 20A: 1–
29.
Söderström L, Urmi E, Váňa J. 2002. The distribution of Hepaticae and
Anthocerotae in Europe and Macaronesia. Lindbergia 27: 3–47.
Söderström L, Urmi E, Váňa J. 2007. The distribution of Hepaticae and
Anthocerotae in Europe and Macaronesia – Update 1–427.
Cryptogamie, Bryologie 28: 299–350.
Wigginton MJ. 2004. E.W. Jones’s liverwort and hornwort flora of West
Africa. Meise: National Botanic Garden of Belgium.
TOM L BLOCKEEL, 9 Ashfurlong Close, Dore, Sheffield S17 3NN, UK. E-mail: Tblockeel@aol.com
Journal of Bryology (2008) 30: 161–167
# British Bryological Society 2008
Received 24 February 2008. Revision accepted 29 February 2008
DOI: 10.1179/174328208X282463
New national and regional bryophyte records, 18
Intending contributors to this column should consult the
Instructions for Authors in part 1 of this volume, and
should address their contributions to the column editor.
1. Aloina brevirostris (Hook. & Grev.) Kindb.
Contributors: H.W. Matcham and J.G. Duckett
Chile: REGION XII MAGALLANES: PROV. DE MAGALLANES:
roadside cutting along Route 5, 58 km north of Punta
Arenas, 52u21988.10S, 71u05917.10E, ca 20 m a.s.l., leg.
H.W. Matcham & J.G. Duckett, 6 September 2006
(UMAG, Priv. Herb. J.G. Duckett, Priv. Herb. H.W.
Matcham).
Predominately a northern hemisphere species, Aloina
brevirostris was previously known from the southern
hemisphere only from South Africa (Hodgetts, Matcham
& Duckett, 1999) and the Alexander Island and James Ross
Island group in the Antarctic (Ochyra, Bednarek-Ochyra &
Lewis Smith, 1998). At the site in Chile, A. brevirostris was
associated with Pterygoneurum ovatum (Hedw.) Dixon. The
Chilean specimens are synoicous, whereas the related A.
catillum (Müll.Hal.) Broth. is described (Delgadillo, 1975)
as being dioicous, cladautoicous or paroicous and has so
far been recorded only from Argentina. According to
Delgadillo (1975) its taxonomic position requires further
study.
2. Bryum archangelicum Schimp.
Contributors: R. Ochyra and R. D. Seppelt
Heard Island: no locality or habitat details given,
53u050S, 73u300E, 19 March 1980, leg. J. Jenkin No. 80/
115 (Seppelt no. 10743) (ADT).
162
BRYOLOGICAL NOTES
Bryum archangelicum is here broadly conceived to
include the species long known as B. inclinatum (Brid.)
Turton (Holyoak, 2004). Because the latter name is a later
homonym, this common Holarctic moss has been named B.
amblyodon Müll.Hal. in recent decades. This reservation is
necessary since some authors (e.g. Nyholm, 1993; Zolotov,
2006) still argue for the specific distinctness of these two
species. The broadly interpreted B. archangelicum is a
bipolar species with some intermediate stations in the
tropical and subtropical regions of South America (Ochi,
1980 as Bryum amblyodon). In the temperate and polar
regions of the Southern Hemisphere B. archangelicum is
scattered in western Patagonia, the Falkland Islands and
the Tierra del Fuego archipelago (Ochi, 1982). Moreover, it
deeply penetrates the maritime Antarctic, extending along
the western coast of the Antarctic Peninsula to central
Alexander Island (Ochyra & Ochi, 1986; Ochyra, Lewis
Smith & Bednarek-Ochyra, in press) and recently it was
also discovered in the Schirmacher Oasis in Dronning
Maud Land on the continent (Ochyra & Singh, in press). In
the Subantarctic, the species has so far been known from a
highly isolated station on Îles Kerguelen (Ochi, 1972) and
the present record represents an extension of the species in
the Kerguelen Province. The material consists of a single
mature capsule which allows an accurate determination.
3. Bryum orbiculatifolium Cardot & Broth.
Contributors: R. Ochyra and R. D. Seppelt
Heard Island: no locality or habitat details given,
53u050S, 73u300E, 19 March 1980, leg. J. Jenkin No. 80/
097 (Seppelt no. 10997) (ADT).
Bryum orbiculatifolium is a very characteristic and readily
recognised species with deeply concave, orbicular leaves. It
has been very occasionally collected in western Patagonia
(Ochi, 1982), subantarctic South Georgia (Ochyra,
Bednarek-Ochyra & Lewis Smith, 2002) and Tristan da
Cunha (Dixon, 1960 as Bryum cymbifoliellum). It extends to
the northern maritime Antarctic where is known from the
volcanic Leskov Island and Candlemas Island in the South
Sandwich Islands archipelago (Convey et al., 2001) and
Deception Island in the South Shetland Islands (Lewis
Smith, 2005a, b, c) and recently, it was also encountered on
the continent in the Schirmacher Oasis in Dronning Maud
Land (Ochyra & Singh, in press). The report of the species
from King George Island in the latter archipelago (Ochyra
& Ochi, 1986; Ochyra, 1998) is based on a misdetermination. The discovery of B. orbiculatifolium on Heard Island
in the Kerguelen Province represents a major extension of
its range and confirms its phytogeographical status as an
amphiatlantic south-temperate species.
4. Bucklandiella didyma (Mont.) Bednarek-Ochyra &
Ochyra
Contributors: H. Bednarek-Ochyra and R. Ochyra
Australia: NEW SOUTH WALES: Diggers Creek, 24 km
N.E. of Mt Kosciusko, 36u210S, 148u290E, 1500 m a.s.l., in
open Eucalyptus forest with introduced Salix along stream,
on exposed rock besides stream, 9 February 1978, leg. H.
Streimann no. 5435 (ALTA, H, KRAM). AUSTRALIAN
CAPITAL TERRITORY: Namadgi National Park, Mt
Gudgenby, 54.5 km SW of Capital Hill, Canberra,
35u46.50S, 148u54.50E, ca 1720 m a.s.l., rocky pinnacle of
Mt Gudgenby, on exposed boulders, 16 January 1985, leg.
J. A. Curnow no. 268 & H. Lepp (DUKE).
New Zealand: OTAGO: Clinton Valley, wet rocks, leg. W.
Petrie no. 619 (in Herb. T. W. Naylor Beckett) and sine
numero (in herb. W. Bell) (H, two specimens).
Bucklandiella didyma is one of the few austral species of
the broadly conceived genus Racomitrium which had not
been considered conspecific with Racomitrium crispulum
(Hook.f. & Wilson) Hook.f. & Wilson by Clifford (1955).
The name has long remained in obsolescence but Deguchi
(1984) examined the original material of this species and
confirmed its distinctiveness. Bednarek-Ochyra, Ochyra &
Buck (1999) outlined the morphological differences
between Bucklandiella didyma and B. crispula (Hook.f. &
Wilson) Bednarek-Ochyra & Ochyra and extended its
geographical range to south-eastern Brazil. Subsequent
revisionary studies of the austral collections revealed that
the species is widely distributed on subantarctic islands
from South Georgia to the Kerguelen Province, and in
Tristan da Cunha, South Africa, and in the northern
maritime Antarctic (Ochyra, Lewis Smith & BednarekOchyra, 2008). Its range is now extended to south-eastern
Australia and the South Island of New Zealand.
Accordingly, B. didyma is defined as a pan-south-temperate
species and the monographic study of Bucklandiella in the
Southern Hemisphere (Bednarek-Ochyra & Ochyra, in
progress), should precisely establish its geographical range
in Australasia.
5. Bucklandiella pachydictyon (Cardot) BednarekOchyra & Ochyra
Contributors: H. Bednarek-Ochyra and R. Ochyra
Prince Edward Islands: MARION ISLAND: no locality or
habitat details given, 53u050S, 73u300E, 14 March 1952, leg.
R.W. Rand no. 3642 (GRO).
Îles Crozet: ÎLE DE LA POSSESSION: summit of Mont
Branca, west of Port Alfred, 380 m a.s.l., 46u250S, 51u500E,
on exposed rock surface, 23 December 1978, leg. Bell 1457
(AAS, KRAM).
Bucklandiella pachydictyon is an amphiatlantic southtemperate species which is widely distributed but scattered
in the Fuegian region and in the Nothofagus zone at
the western fringes of southern South America where it
extends northward to the Cautin Province of Chile in
the Valdivian region (Herzog, 1938), with an isolated
location in Central Andes of Bolivia at an elevation of
3600 m (Churchill, Griffin & Muñoz, 2000). In addition, it
occurs frequently on subantarctic South Georgia (Bell,
1974) and infrequently on Îles Kerguelen (Cardot, 1916;
Ochyra, personal observations), and penetrates southwards
into the maritime Antarctic (Ochyra, Lewis Smith &
Bednarek-Ochyra, in press). The discovery of the species
BRYOLOGICAL NOTES
on Marion Island in the Prince Edward Islands archipelago
and on Île de la Possession in the Îles Crozet archipelago
completes its continuous range in the western part of the
Subantarctic.
6. Entodon concinnus (De Not.) Paris
Contributors: Tamás Pócs, B.O. van Zanten and Peter
Erzberger
Hungary: GÖDÖLLŐ HILLS: Comit. Pest, Vácrátót,
Botanical Garden, in the turf of the Conifer collection, 8
September 1983, leg. et det. T. Pócs & B.O. van Zanten s.n.,
conf. P. Erzberger, 30 April 2007 (EGR, dupl. in B, BP).
In the checklist of Hungarian bryophytes (Erzberger &
Papp, 2004), Entodon concinnus was excluded because
earlier reports had been considered doubtful by
Hungarian authorities (Boros, 1968; Orbán & Vajda,
1983) and no specimen could be located. Recently, T.P.
made available several specimens from EGR, which had
not been seen during the preparation of the checklist, and
one proved to be E. concinnus.
Perhaps this species has been overlooked in the past due
to a superficial similarity to common species like
Pseudoscleropodium
purum
(Hedw.)
M.Fleisch.,
Pleurozium schreberi (Willd. ex Brid.) Mitt., or
Calliergonella cuspidata (Hedw.) Loeske. These species
have in fact been confused with E. concinnus, as is evident
from several incorrectly named specimens in Hungarian
herbaria seen by P.E.
The apparent rarity of E. concinnus in Hungary is
somewhat enigmatic, since there should be sufficient
potential growth sites in the limestone districts.
163
This specimen was unnamed by the collector; however,
later ‘Grimmia apocarpa (L.) Hedw.’ was added in a
different handwriting (perhaps by Á. Boros or L. Vajda).
During revision of Schistidium specimens from BP by P.E.
and Wiebke Schröder (Erzberger & Schröder, 2008), the
specimen was recognised as Grimmia sp. and sent to E.M.
for revision, who established its true identity. The collection
site on the north side of Lake Balaton is famous for its
basalt rocks with the relict fern Notholaena marantae (L.)
Desv., and interesting bryophytes like Pterogonium gracile
(Hedw.) Sm., Saelania glaucescens (Hedw.) Broth. in
crevices, and Mannia fragrans (Balb.) Frye & L.Clark,
Oxymitra incrassata (Brot.) Sérgio & Sim-Sim, Pyramidula
tetragona (Brid.) Brid. and others in the open grasslands
near the top (414 m a.s.l.).
A second Hungarian locality of G. elatior was discovered
recently. In August 2007, P.E. and P.Sz. collected two
specimens of Grimmia, which were revised as G. elatior by
E.M.: Com. Pest, Mt Naszály near Vác, 47u499300N,
19u099500E, ca 550 m a.s.l., on sandstone boulder scree
near the summit, associated with Hedwigia ciliata var.
leucophaea Bruch & Schimp., leg. P. Erzberger nos. 12613,
12614 and P. Szűcs, 11 August 2007, det. E. Maier, 22
October 2007 (B).
Grimmia elatior is not included in the recent checklist of
Hungary (Erzberger & Papp 2004). It is new to the
bryophyte flora of Hungary.
7. Grimmia decipiens (Schultz) Lindb.
Contributors: Peter Erzberger and Eva Maier
Hungary: BALATON UPLANDS: Comit. Veszprém, on the
north face of the hill ‘Halápihegy’ near Zalahaláp, north of
Tapolca, 46u55950.40N, 17u27915.80E, ca 250 m a.s.l., on
basalt scree (waste from quarry), leg. P. Erzberger
no. 10833, 30 March 2005, det. E. Maier, 19 September
2005 (B).
Grimmia decipiens was considered doubtful in the recent
checklist of the bryophytes of Hungary (Erzberger & Papp,
2004) because earlier reports had been rejected by Boros
(1968) and no specimen had been located. Therefore, the
above-mentioned collection is the first confirmed record for
Hungary.
9. Grimmia lisae De Not.
Contributors: Peter Erzberger and Eva Maier
Hungary: BÖRZSÖNY MTS: Comit. Nógrád, on andesite
boulders in the valley of the stream ‘Kemence-patak’ near
the village of Diósjenő, leg. L. Vajda, 10 April 1955 (sub
Grimmia apocarpa (L.) Hedw.), det. E. Maier, March 2007.
Specimen in BP (46335). The original label on the specimen
reads: Herbarium Musei Hist. Nat. Hungar. Budapest,
Flora Hungarica, Grimmia apocarpa (L.) Hedw. Comit.
Nógrád. In rupibus andesiticis vallis rivi Kemencepatak,
prope Diósjenő, montes Börzsöny. 10/IV 1955 Leg.: Det.:
L. Vajda.
This specimen was sent to E.M. for determination during
revision of Schistidium in Hungarian herbaria by P.E. and
Wiebke Schröder (Erzberger & Schröder, 2008).
Grimmia lisae is not included in the recent checklist of
Hungary (Erzberger & Papp, 2004), it is a new species in the
bryophyte flora of Hungary.
8. Grimmia elatior Bruch ex Bals.-Criv. & De Not.
Contributors: Peter Erzberger, Eva Maier and Péter Szűcs
Hungary: BALATON UPLANDS: Comit. Veszprém, on the
hill ‘Szentgyörgyhegy’ near Tapolca, on basalt, leg. J.
Szepesfalvi, 8 April 1927, det. E. Maier, July 2007 (BP,
159733). The original label on the specimen reads: Herbar.
Musei Nat. Hungar. Budapest, Flora Hungarica, Grimmia
apocarpa (L.) Hedw. Comit. Zala. In monte
‘Szentgyörgyhegy’ ad opp. Tapolca. Solo basaltico.
1927.IV.8. leg. J. Szepesfalvi.
10. Orthotrichum pulchellum Brunt.
Contributors: V. Plášek and I. Marková
Czech Republic: BOHEMIA: České Švýcarsko National
Park, 2.5 km W.N.W. of Chřibská village, valley of
Doubický potok stream, loc. ‘hájenka Saula’, bark of
Fraxinus excelsior, GPS coordinates (WGS 84) 50u529210N,
14u269480E, 335 m a.s.l., 3 August 2006. leg. I. Marková
no. 78/2006, det. V. Plášek (Priv. Herb. Marková).
The specimen cited above is the first collection of
this epiphytic moss from the Czech Republic. In
164
BRYOLOGICAL NOTES
Europe, it is reported from many countries situated mainly
along the western seacoasts (from southern Scandinavia to
northern Spain). The number of its localities has increased
recently and the moss appears to be spreading to the east
(Frahm, 2002; Lüth, 2004). In the Czech Republic, it was
recorded growing vertically on the bark of Fraxinus
excelsior at a height of 205 cm above the ground, with an
E.N.E. aspect. The size of the population was 2 cm2, and
associated species were: Brachythecium salebrosum (Hoffm.
ex F.Weber & D.Mohr) Schimp., Hypnum cupressiforme
Hedw. and Orthotrichum pumilum Sw. ex Anon.
11. Philonotis polymorpha (Müll.Hal.) Broth.
Contributors: R. Ochyra, H. Bednarek-Ochyra and R. D.
Seppelt
Macquarie Island: (1) Sawyer Creek Gorge, 54u309S,
158u53930.00E, on rock on steep seepage slope opposite
main waterfall gorge, 5 December 1981, leg. R. D. Seppelt
no. 12109 (ADT, KRAM); (2) Flat Creek Waterfall, S.E. of
Cormorant Point, 54u34928.00S, 158u529E, 25 m a.s.l., on
peat in hanging moss carpet in splash zone at edge of
watercourse, with Montia fontana, Brachythecium, 5
November 1981, leg. R. D. Seppelt no. 11600 (ADT,
KRAM).
Philonotis polymorpha is a poorly and still imperfectly
known hydrophytic species, growing mostly on stream
banks, as well as shores of ponds and lakes, occasionally
forming swards of several square metres. The plants can
sometimes reach a considerable length, to 30 cm. It is
widely distributed on the Southern Ocean islands, including
South Georgia (Clarke, 1973 as P. acicularis), Prince
Edward Islands (van Zanten, 1971 as P. cf. angustifolia;
Ochyra, personal observations), Tristan da Cunha (Dixon,
1937) and Îles Kerguelen from whence it was originally
described (Müller, 1883, 1889), and reaches the northern
maritime Antarctic (Ochyra, Bednarek-Ochyra & Lewis
Smith, 1998). With the present record, its geographical
range is markedly extended to subantarctic Macquarie
Island in the Australasian sector of the Subantarctic.
Accordingly, the phytogeographical status of P. polymorpha has to be redefined as circumsubantarctic. This species
is omitted from the recently published moss flora of
Macquarie Island and it was misnamed Aulacomnium
palustre (Hedw.) Schwägr. (Seppelt 2004), although the
latter species also occurs on the island. The discovery of P.
polymorpha and the recent addition of Bucklandiella
lamprocarpa (Müll.Hal.) Bednarek-Ochyra & Ochyra
(Bednarek-Ochyra & Ochyra, 2007) increases the number
of moss species known from this subantarctic island to 86,
with a further four taxa which have not yet been definitely
named to species.
12. Physcomitrium spathulatum var. spathulatum
(Hornsch.) Müll.Hal.
Contributors: H.W. Matcham and J.G. Duckett
Botswana: Tuli Game Reserve, ca 200 km south east of
Francistown, sandy soil, north bank of the River Limpopo,
22u12936.50S, 29u06934.10E, ca 519 m a.s.l., leg. H.W.
Matcham & J.G. Duckett, 24 November 2005 (Priv.
Herb. J.G. Duckett, Priv. Herb. H.W. Matcham).
13. Pterygoneurum ovatum (Hedw.) Dixon
Contributors: H.W. Matcham and J.G. Duckett
Chile: REGION XII MAGALLANES: PROV. DE MAGALLANES:
roadside cutting along Route 5, 58 km north of Punta
Arenas, 52u21988.10S, 71u05917.10E, ca 20 m a.s.l., leg.
H.W. Matcham & J.G. Duckett, 6 September 2006
(UMAG, Priv. Herb. J.G. Duckett, Priv. Herb. H.W.
Matcham).
Pterygoneurum ovatum is new to Latin America.
Elsewhere in the southern hemisphere, it is recorded from
Antarctica (Lewis Smith, 1985; Ochyra, Lewis Smith &
Bednarek-Ochyra, in press) and Australia (Scott & Stone,
1976). At the site in Chile, it was associated with Aloina
brevirostris (Hook. & Grev.) Kindb.
14. Schistidium frigidum H.H.Blom var. frigidum
Contributor: Louis Thouvenot
Andorra (Pyrenees): ORDINO: Coma del Forat (Arcalis),
UTM 31T CH7519, at base of sheer slope of mica schist
crag, 2456 m a.s.l., with Andreaea rupestris var. rupestris
Hedw., leg. L. Thouvenot no. 3281, 15 July 2007, conf. H.
H. Blom (PC, Priv. Herb. Thouvenot).
Schistidium frigidum has a circumpolar arctic-alpine
distribution and in Europe, it is known from the Nordic
countries and the Alps (Blom, 1996). This species is
unrecorded in the Pyrenees, the Iberian Peninsula (Casas,
2000) and France. The present discovery in Andorra
(Eastern Pyrenees) extends its area towards the southwestern European high mountains. According to Blom, the
species is very variable and the features of the Andorran S.
frigidum include small cupulate urns, somewhat longer than
wide, distinct marginal basal leaf cells (in 1–3 rows), mostly
hyaline, with thickened cross-walls, and basal cells near
costa wider and thin-walled, forming a conspicuous hyaline
group. The vegetative leaves are mostly epilose and the
median cells variously smooth or sinuose.
15. Scorpiurium sendtneri (Schimp.) M.Fleisch.
Contributor: V. Hugonnot
France: ARDÈCHE: Les Vans, Paı̈olive, 4.25988uE,
44.4428uN, leg. V. Hugonnot, 28 March 2005, and
additional sites in the Paı̈olive district (PC, Priv. Herb.
Hugonnot).
Mastracci (2001) reported Scorpiurium sendtneri as new
to France, from an old herbarium sheet labelled ‘Thamnium
alopecurum forme Draguignan’ in ‘Provence, Draguignan,
du pont de clorques a Encapis 8 janvier 1912 Girod ver:
Theriot’. The specimen is apparently stored in G but seems
to be lost. Cortini Pedrotti (2006, p. 862) writes that after
examination of the type specimen of Thamnobryum
cossyrense (Bott.) A.J.E.Sm. stored in FI, she came to the
conclusion that it is identical with Scorpiurium circinatum
(Brid.) M.Fleisch. & Loeske and not with S. sendtneri. This
165
BRYOLOGICAL NOTES
is in contradiction with the views of Mastracci (2001), who
placed it in synonymy with S. sendtneri. I (V.H.) have been
able to study an isolectotype of Thamnium cossyrense Bott.
(under PC0080128) and it is equally referable to
Scorpiurium circinatum. This casts some doubt about the
identity of the French specimen of ‘Thamnium alopecurum
forme Draguignan’ and consequently the latter should be
re-evaluated if the specimen can be traced. All the specimens of Scorpiurium sendtneri stored in PC have been
examined and no additional specimen originating from
France could be found. Consequently, S. sendtneri is here
confirmed for France. It is basically a northern
Mediterranean species with radiations in to Macaronesia
and south-western Asia. It has recently been added to the
African bryoflora, in Morocco at Jbel Bouhalla (Jiménez
et al., 2002). Walther (1975) uses the informal combination
‘monstr. Microphylla’ to describe the epiphytic monstrous
form bearing numerous caducous branchlets bearing
reduced leaves. This form has been observed very frequently in south Ardèche, including over calcareous rocks.
The basic structure of that form is generally not evident in
the field because it is greatly obscured by the profusion of
caducuous branchlets (tertiary axes). All the new localities
in Ardèche are situated in the Paı̈olive karstic forest, on
small vertical and dry calcareous walls, well shaded by the
trees in the immediate vicinity. Scorpiurium sendtneri has
been observed too on the bark of Quercus humilis Miller,
but this habitat seems to be more exceptional locally.
16. Vittia elimbata Hedenäs, Vanderpoorten & Goffinet
Contributors: H.W. Matcham, J.G. Duckett and L.
Hedenäs
Chile: REGION XII MAGALLANES: PROV. DE ULTIMA
ESPERANZA: Laguna Sofia, ca 40 km north-west of Puerto
Natales, 51u549S, 72u599E, ca 111 m a.s.l., leg. H.W. Matcham
& J.G. Duckett, 7 September 2006, det. L. Hedenäs (BM,
KRAM, S, UMAG, Priv. Herb. H.W. Matcham).
Vittia elimbata was collected from the edge of the lake at
this site, where it would certainly have been inundated
periodically; it was possibly washed up by wave action from
deeper water. Reassessing the taxonomic position of the
Vittiaceae, Vanderpoorten et al. (2003) described V.
elimbata from a single collection in Bolivia at Laguna
Ventanani. Gathered from running water on the face of a
waterfall at 3810 m a. s. l., it differed from the other two
species described in the genus, viz. V. pachyloma (Mont.)
Ochyra and V. salina Hedenäs & J. Muñoz, in lacking a bior multistratose leaf border, but had a partly bistratose
lamina in the upper part of the leaf. Vanderpoorten et al.
(2003)
synonymized
the
Vittiaceae
into
the
Amblystegiaceae.
Our collection from Laguna Sofia in the Patagonian
steppe represents a markedly southerly extension of range
of this aquatic moss, although it is not an entirely
unexpected extension phytogeographically, considering
the number of other disjunct species found in the Andes
and in southern South America.
ACKNOWLEDGEMENTS
H. Bednarek-Ochyra and R. Ochyra are grateful to
Curators at ALTA, DUKE, GRO and H for arranging
specimens on loan; their studies have gained financial
support from the Polish Ministry of Science and
Higher Education through grants no. N 303 063 32/2264
for H. Bednarek-Ochyra and no. 2 P04G 043 29 for R.
Ochyra. P. Erzberger and E. Maier wish to thank B. Papp,
the curator of the herbarium at BP, for the loan of
specimens.
TAXONOMIC ADDITIONS AND CHANGES: Nil.
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T. L. BLOCKEEL1, 9 Ashfurlong Close, Dore, Sheffield S17 3NN, UK. E-mail: Tblockeel@aol.com
H. BEDNAREK-OCHYRA & R. OCHYRA, Laboratory of Bryology, Institute of Botany, Polish Academy of Sciences, ul. Lubicz 46, 31-512
Kraków, Poland. E-mails: Halina.Bednarek@ib-pan.krakow.pl and Ryszard.Ochyra@ib-pan.krakow.pl
JEFFREY G. DUCKETT, School of Biological Sciences, Queen Mary, University of London, Mile End Road, London, E1 4NS, UK.
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P. ERZBERGER, Belziger Str. 37, D-10823 Berlin, Germany. E-mail: erzberger@erzfisch.de
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I. MARKOVÁ, Bohemian Switzerland National park Administration, Pražská 52, CZ-407 46 Krásná Lı́pa, Czech Republic.
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HOWARD W. MATCHAM, 21, Temple Bar, Strettington, Chichester, West Sussex, PO18 0LB, UK. E-mail: hwlgmatch@yahoo.co.uk
V. PLÁŠEK, Department of Biology and Ecology, University of Ostrava, Chittussiho 10, CZ-710 00 Ostrava 10, Czech Republic.
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T. PÓCS, Research Group for Bryology of the Hungarian Academy of Sciences, Department of Botany, Eszterházy College, Eger, P.B. 222,
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P. SZŰCS, Verő J.u. 1, H-2931 Almásfüzitő, Hungary.
BRYOLOGICAL NOTES
167
LOUIS THOUVENOT, 11 rue Saint Léon, F-66000 Perpignan, France. E-mail: thouloup@club-internet.fr
B. O. van ZANTEN, State University Groningen, Biological Centre, Department of Biology of Plants, PO Box 14, NL-9750 AA Haren, The
Netherlands. E-mail: bovzanten@home.nl
1
Column editor, to whom contributions should be sent.
Journal of Bryology (2008) 30: 167–170
# British Bryological Society 2008
Received 26 February 2008. Revision accepted 26 March 2008
DOI: 10.1179/174328208X300570
Notes on some newly recorded bryophytes from Montenegro
The bryophyte flora of Montenegro is still poorly explored,
although considerable progress has been made recently.
This is reflected in the publication of a checklist based to a
large extent on literature reports (Dragićević & Veljić, 2006)
and of several comprehensive contributions (Milikić et al.,
2001; Veljić et al., 2001; Dragićević et al., 2001, 2003;
Martinčič, 2006; Erzberger & Papp, 2007; Papp &
Erzberger, 2007).
The present paper reports observations made during a
collection trip to the Orjen Mts and the Bjelasica Mts in
July 2007. A complete account of the floristic results will be
published elsewhere, but here we wish to draw attention to
25 taxa, eight liverworts and 17 mosses, which, to our
knowledge, have not yet been documented for Montenegro.
The Bjelasica Mts are one of the few Montenegran
mountain ranges composed of siliceous bedrock, while the
Orjen Mts are carboniferous limestone. Owing to their
situation close to the sea, the Orjen Mts receive an
exceptionally high amount of precipitation (y8000 mm/y;
Komar, 1995), the runoff from which vanishes almost
immediately in the karstic underground. This can lead to
very special microsites where erosion exposes patches of
subneutral soil from which the bases have been washed out,
and where some subneutrophytic and even acidophytic
bryophytes that would not be expected in a calcareous
mountain region can be observed: Trematodon ambiguus,
Pleuridium acuminatum, Pohlia andalusica, Pogonatum
nanum, Polytrichum commune var. perigoniale and
Lophozia sudetica (Table 1, site 6).
Another site that deserves special mention is the region
around Zekova glava peak (2020–2080 m a.s.l.) in the
Bjelasica Mts (Table 1, sites 14 and 15). Here, in an area
with obvious late snow beds, several species new to
Montenegro have been found: Anthelia juratzkana,
Diplophyllum taxifolium, Gymnomitrion concinnatum,
Marsupella
sphacelata,
Encalypta
microstoma,
Pseudotaxiphyllum elegans and Schistidium umbrosum.
Out of 15 sites visited, novelties are reported from nine
sites (Table 1). Nomenclature follows Hill et al. (2006) and
Grolle & Long (2000). Voucher specimens are held in the
Botanical Museum at Berlin–Dahlem (B: collected by P.E.)
and the Hungarian Natural History Museum at Budapest
(BP: collected by B.P.). In evaluating additions to the
bryoflora of Montenegro, we consulted, in addition to the
papers mentioned above, the regional checklists by Düll
(1995), Düll et al. (1999), Sabovljević & Stevanović (1999),
Sabovljević (2000), Ganeva & Natcheva (2003), Martinčič
(2003), Natcheva & Ganeva (2005), Colacino & Sabovljević
(2006), Sabovljević & Natcheva (2006), and a preliminary
version of the check-list of the mosses of south-east Europe
(in prep.).
Anthelia juratzkana (arctic-alpine)
15a, on soil, associated with Gymnomitrion concinnatum,
Marsupella sphacelata, Saelania glaucescens, Timmia austriaca, leg. P.E. 12565, 12569, conf. J. Váňa (B); 15b leg.
B.P. (BP 49229/H).
Calypogeia suecica (suboceanic-montane)
8, on decaying wood with Riccardia palmata, leg. B.P.
(BP 49230/H). This species, known from six countries in
south-east Europe, has perhaps been overlooked. In Serbia,
it is a recent addition (Papp & Sabovljević, 2002).
Diplophyllum taxifolium (boreal subarctic-alpine)
14b, in rock fissures, associated with Bartramia ithyphylla, Heterocladium dimorphum, leg. P.E. 12558, rev. J.
Váňa (B). This is the first record of the genus Diplophyllum
for Montenegro.
Gymnomitrion concinnatum (subarctic alpine)
14a, on soil in siliceous rock crevices associated with
Barbilophozia hatcheri, Marsupella sphacelata, Lophozia
sudetica, leg. B.P. (BP 49231/H); 15a, on soil, associated
with Anthelia juratzkana, Marsupella sphacelata, Saelania
glaucescens, Timmia austriaca, leg. P.E. 12561, 12564,
12565, 12573, conf. J. Váňa (B); 15b, leg. B.P. (BP 49232/
H). G. concinnatum, like A. juratzkana and M. sphacelata, a
typical late-snow bed hepatic, has been added to the
Serbian list recently (Erzberger & Papp, 2007).
Lophozia ascendens (boreal-montane)
8, on decaying wood with Lophozia ventricosa, Lepidozia
reptans, leg. B.P. (BP 49233/H); and leg. P.E. 12398 (B). In
Serbia, L. ascendens is a recent addition (Papp & Sabovljević,
2002). We have also found it in the Prokletije Mts in
Montenegro recently (Papp & Erzberger, unpublished).
Lophozia sudetica (boreal-montane)
6, on soil in limestone rock crevices associated with
Pohlia andalusica and P. cruda, leg. B.P. (BP 49234/H). 9,
on siliceous rock associated with Andreaea rupestris, leg.
B.P. (BP 49235/H); and 14a, on soil in siliceous rock
crevices with Barbilophozia hatcheri, Gymnomitrion concinnatum and Marsupella sphacelata, leg. B.P. (BP 49236/
H).
168
BRYOLOGICAL NOTES
3, on soil overlying limestone, near road, leg B.P. (BP
174415) and P.E. 12303 (B). The occurrence of C.
chloropus, previously known from seven countries in
south-east Europe, was to be expected in the mediterranean
part of Montenegro.
Encalypta microstoma
14a, on soil among siliceous rocks associated with
Heterocladium dimorphum, leg. B.P. (BP 174416) This rare
species of the higher mountains has been recorded in southeast Europe only in Bulgaria (Rila Mts) and recently in
Serbia (Stara Planina Mts; Erzberger & Papp, 2007).
Grimmia anomala (subarctic-subalpine)
9, on dry siliceous rock outcrops, associated with
Grimmia caespiticia, leg. P.E. 12441, 12443, 12445, conf.
E. Maier (B) and leg. B.P. (BP 174417). G. anomala, a rare
species, has been added to the Serbian and Albanian lists
recently (Papp, Erzberger & Sabovljević, 2004; Erzberger,
2008), but is absent from the other south-east European
countries except Romania. In a Red Data List of European
Like the previous species, a recent addition to Serbia
(Papp & Erzberger, 2005).
Marsupella sphacelata (northern suboceanic-montane)
14a, on soil in siliceous rock crevices associated with
Barbilophozia
hatcheri,
Gymnomitrion
concinnatum,
Lophozia sudetica, leg. B.P. (BP 49237/H); 15a, on soil,
associated with Gymnomitrium concinnatum, Anthelia juratzkana, leg. P.E. 12561, 12565, det. J. Váňa (B); 15b, leg.
B.P. (BP 49238/H). This appears to be the first record for
the genus Marsupella in Montenegro. M. sphacelata,
previously recorded in three countries only in south-east
Europe, appears to be rare in the region.
Tritomaria quinquedentata (circumboreal arctic-alpine)
8, on soil near path, associated with Plagiochila
porelloides, Eurhynchium angustirete, leg. P.E. 12426; 9,
on dry siliceous rock outcrops, associated with
Heterocladium dimorphum, leg. P.E. 12489, conf. J. Váňa
(B).
Cheilothela chloropus (oceanic-mediterranean)
Table 1.
Additions to the bryoflora of Montenegro (July 2007).
Sites
1
Anthelia juratzkana
Calypogeia suecica
Diplophyllum taxifolium
Gymnomitrium concinnatum
Lophozia ascendens
Lophozia sudetica
Marsupella sphacelata
Tritomaria quinquedentata
Cheilothela chloropus
Encalypta microstoma
Grimmia anomala
Grimmia torquata
Orthotrichum acuminatum
Pogonatum nanum
Polytrichum commune var. perigoniale
Pseudotaxiphyllum elegans
Racomitrium lanuginosum
Schistidium dupretii
Schistidium papillosum
Schistidium umbrosum
Syntrichia virescens
Taxiphyllum wissgrillii
Trematodon ambiguus
Ulota bruchii
Weissia controversa var. crispata
Distribution in SE-Europe
2
3
5
6
8
9
14
15
Previously known
N
BG, HR, MK, RO, SI
BA; BG, HR, RO, RS, SI
BG, RO, SI
BG, RO, RS, SI
BG, HR, RO, RS
BG, MK, RO, RS, SI
BG, GR, RO
AL, BG, HR, MK, RO, RS, SI
AL, BA, BG, GR, HR, RS, TR
BG, RS
AL, RO, RS
BG, GR, RO, RS
BA, GR, HR
All except AL, ME
HR, GR, MK, RS, SI
BA, BG, GR, HR, RO, RS, SI
BA, BG, GR, RO, RS, SI
GR, MK, RO
BG, GR, RO, RS, (SI)
BA
All except AL, ME, MK
All except AL, ME, MK
BG, RO, SI
BA, HR, RO, RS, SI
BA, HR, MK, RS, SI
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
List of localities: 1–6: Orjen Mts; 8–15: Bjelasica Mts.
1: around Vrbanje, 42u33929.60N, 18u30930.70E, 1005 m a.s.l., 13 and 16 July 2007; 2a: near Orjen sedlo, in Pinetum heldreichii, 42u33927.30N,
18u33906.60E, ca 1600 m a.s.l., 14 July 2007; 2b: near the summit of Zubački kabao (Orjen peak), 42u34909.20N, 18u32938.60E, 1900 m a.s.l., 14 July
2007; 3: around Kruševice, 42u32904.80N, 18u29940.00E, 665m a.s.l., 15 July 2007; 5: between Orjen sedlo and Crkvice, around Reovci north of Velji
Kabao, 42u33946.40N, 18u35938.40E, 1240 m a.s.l., 16 July 2007; 6: between Orjen sedlo and Crkvice, Reovci, grassland with dolinas, 42u34909.20N,
18u35914.90E, 1215 m a.s.l., 16 July 2007; 8: Kolašin town, Jezerine, at Svinjača stream, 42u49949.50N, 9u37906.70E, 1360 m a.s.l., 18 July 2007; 9:
Katun Rupe at Kolašin town, 42u49909.90N, 19u36959.70E, 1660 m a.s.l., 19 July 2007; 14a: between Troglava and Zekova glava peaks at Kolašin
town 42u51905.70N, 19u40908.80E, 2022 m a.s.l., 20 July 2007; 14b: between Troglava and Zekova glava peaks at Kolašin town, 42u51902.50N,
19u40915.70E, ca 1990 m a.s.l., 20 July 2007; 15a: Zekova glava peak at Kolašin town, siliceous rock outcrops at slope above lake Pešica Jezero,
42u51900.10N, 19u41905.10E, ca 2020 m a.s.l., 20 July 2007; 15b: Zekova glava peak at Kolašin town, siliceous rocks, 42u51904.20N, 19u40954.60E,
2080 m a.s.l., 20 July 2007.
Abbreviations (ISO 3166-1): AL5Albania; BA5Bosnia and Hercegovina; BG5Bulgaria; GR5Greece; HR5Croatia; ME5Montenegro;
MK5FYR Macedonia; RO5Romania; RS5Serbia; SI5Slovenia.
BRYOLOGICAL NOTES
Grimmia species proposed by Greven (1995), G. anomala is
classified as an endangered species. The populations in the
Bjelasica Mts are very large.
Grimmia torquata (boreo-arctic montane)
9, under overhanging siliceous rocks, leg. P.E. 12486 (B)
and leg. B.P. (BP 174418).
G. torquata has been added to the Serbian list recently
(Erzberger & Papp, 2007).
Orthotrichum acuminatum (submediterranean-montane)
3, on the bark of Tilia with O. affine, O. pallens, O.
striatum, leg. B. P. (BP 174539) conf. R. Garilleti.
Pogonatum nanum (suboceanic-submediterranean)
6, on freshly exposed soil in an erosion runnel, leg. P.E.
12363 (B).
Polytrichum commune var. perigoniale (sub-boreal
holarctic)
6, on soil among limestone rocks, leg. B.P. (BP 174420).
For the distribution of this taxon in south-east Europe,
only Düll et al. (1999) could be consulted, since other
sources do not differentiate it from P. commune. Var.
perigoniale is considered as a full species by some
bryologists (Schoepe, 2000; Meinunger & Schröder, 2007).
In Serbia and Greece, it is a recent addition (Papp &
Erzberger, 2005; Tsakiri, Papp & Szurdoki, 2006).
Pseudotaxiphyllum elegans (suboceanic)
14a, on soil in siliceous rock crevices associated with
Plagiothecium
denticulatum,
Lophozia
ventricosa,
Barbilophozia hatcheri, leg. B.P. (BP 174421).
Racomitrium
lanuginosum
(boreo-arctic
montane
circumpolar)
14b, on siliceous rock, associated with Dicranum
scoparium, Dicranoweisia crispula, leg. P.E. 12544, 12545
(B). This species is a recent addition to Serbia (Erzberger &
Papp, 2007).
Schistidium dupretii (circumboreal-montane)
2a, on limestone rock, leg. P.E. 12266 conf. W. Schröder
(B); 8, on concrete of a broken-down bridge, associated with
S. crassipilum, Tortula muralis, Syntrichia ruralis, Encalypta
streptocarpa, leg. P.E. 12397 conf. W. Schröder (B). This
species has been reported from Greece, Macedonia and
Romania (Blom, 1996); it is new to Montenegro. Its
distribution is imperfectly known, since it has only recently
been brought to the attention of bryologists. The same can
be said for the other species of the genus.
Schistidium
papillosum
(circumpolar
boreo-arctic
montane)
9, on dry siliceous rock outcrops, leg. P.E. 12483 conf. W.
Schröder (B).
This species is in south-east Europe at present known
from Bulgaria, Greece (Lüth, 2003), Romania and Serbia,
where it is a recent addition (Erzberger & Papp, 2007), and
is doubtfully recorded from Slovenia (Martinčič, 2003).
Schistidium umbrosum
15a, on soil, associated with Dicranoweisia crispula,
Anthelia juratzkana, Saelania glaucescens, Timmia austriaca, leg. P.E. 12567, 12568 det. H. Köckinger (B). This
appears to be the second record of the species for the
169
Balkan peninsula. Blom (1996) mentions a single collection
from Bosnia-Hercegovina. S. umbrosum appears to be a
rare alpine species.
Syntrichia virescens (european temperate)
1, on the bark of Robinia near the well at the monument,
intermingled with cushions of Syntrichia laevipila, but
differentiated by smaller size, the denticulate hair-point
and the costa papillose on the abaxial side, leg. P.E. 12388
(B). 9, on siliceous rocks associated with Pseudoleskea
patens, Grimmia muehlenbeckii, Grimmia anomala, leg. B.P.
(BP 174423) and leg. P.E. 12461 (B); and on the bark
of Fagus associated with Pseudoleskeella nervosa,
Orthotrichum stramineum, O. pumilum, Pterigynandrum
filiforme leg. B.P. (BP 174424), and leg. P.E. 12456 (B).
Taxiphyllum wissgrillii (suboceanic)
5, on shaded limestone rock associated with
Gymnostomum aeruginosum, leg. B.P. (BP 174425).
Trematodon ambiguus
6, on soil, leg. B.P. (BP 174426), dupl. (B Erzberger
12385). This rare species with a scattered distribution in
Europe was known from Bulgaria (only found in the first
half of the twentieth century), Romania and Slovenia.
During, Verduyn & van Tooren (2007) studied the
ecological requirements of this species in Belgium and the
Netherlands, where it has been observed more frequently in
the recent past, probably due to the increased use of sod
cutting as a conservation management tool. They conclude
that the species requires a rather high pH in an otherwise
acid environment, and they state that they ‘‘do not know of
any locality for this species with large-scale base-rich
conditions’’ (During et al., 2007). Our finding, on the other
hand, seems to be just this: the underlying limestone
bedrock surely provides overall base-rich conditions, and
under these conditions neutral or mildly acid soil is
available in restricted patches only. In an area without tree
cover, the often copious amounts of rain must lead to
strong soil erosion. Obviously, these factors combine to
wash out bases from the soil superficially and create
patches of bare soil, where short-lived species like T.
ambiguus can thrive without competitors.
Ulota bruchii (northern suboceanic european)
8, on a thin branch of Picea abies near the water, leg. P.E.
12406 (B).
Weissia controversa var. crispata (Weissia fallax Sehlm.)
(submediterranean-montane holarctic)
1, on soil between partly shaded calcareous rocks,
associated with Homalothecium sericeum and Tortella
tortuosa leg. P.E. 12245 (B); 2b, on soil among limestone
rocks in Pinetum heldreichii, associated with Ditrichum
flexicaule, leg. P.E. 12278 (B). This taxon is considered a
full species by some bryologists (Meinunger & Schröder,
2007; Schlüsslmayr, 2005).
We are very grateful to H. Köckinger and W. Schröder
for the examination of Schistidium specimens, to J. Váňa
for revision of hepatics, and to E. Maier for the
examination of Grimmia specimens, to R. Garilleti for the
170
BRYOLOGICAL NOTES
confirmation of Orthotrichum acuminatum, and to M.
Sabovljević for making available a preliminary version of
the checklist of the mosses of south-east Europe. The
warm-hearted hospitality of D. Vincek during our stay at
Kolašin and his enthusiasm for our work deserve our
special thanks.
TAXONOMIC ADDITIONS AND CHANGES: Nil.
REFERENCES
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preliminary check-list. Cryptogamie Bryologie 27: 471–498.
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the flora of Montenegro. Flora Mediterranea 11: 109–113.
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the knowledge of the moss flora of the Tara river canyon. Natura
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Trematodon ambiguus in lowland Belgium and the Netherlands.
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data on their distribution. I. Hepaticae and Anthocerotae.
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Anthocerotae of Europe and Macaronesia. Journal of Bryology 22:
103–140.
Hill MO, Bell N, Bruggeman-Nannenga MA, Brugués M, Cano MJ,
Enroth J, Flatberg KI, Frahm J-P, Gallego MT, Garilleti R, Guerra
J, Hedenäs L, Holyoak DT, Hyvönen J, Ignatov MS, Lara F,
Mazimpaka V, Muñoz J, Söderström L. 2006. An annotated
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Bryology 28: 198–267.
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25.
Martinčič A. 2003. Seznam listnatih mahov (Bryopsida) Slovenije.
Hacquetia 2: 91–166.
Martinčič A. 2006. Moss flora of the Prokletije Mountains (Serbia,
Montenegro). Hacquetia 5: 113–130.
Meinunger L, Schröder W. 2007. Verbreitungsatlas der Moose
Deutschlands. Volume 2. Regensburg: Regensburgische
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of the Morača River Basin. Archives of Biological Sciences,
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II. Musci. Cryptogamie Bryologie 26: 209–232.
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Biosphere Reserve and some adjacent sites (SW Serbia, SerbiaMontenegro). Studia Botanica Hungarica 36: 101–116.
Papp B, Erzberger P. 2007. Contribution to the bryophyte flora of
Montenegro. Studia Botanica Hungarica 38: 79–94.
Papp B, Erzberger P, Sabovljević M. 2004. Contributions to the
bryophyte flora of Kopaonik Mts (Serbia, Serbia–Montenegro).
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Papp B, Sabovljević M. 2002. The bryopyhte flora of Tara National
Park (W Serbia, Yugoslavia). Studia BotanicaHungarica 33: 25–
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hornworts of Southeast Europe. Phytologia Balcanica 12: 169–180.
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Republic of Yugoslavia. Flora Mediterranea 9: 65–95.
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P. ERZBERGER, Belziger Str. 37, D-10823 Berlin, Germany. E-mail: erzberger@erzfisch.de
B. PAPP, Department of Botany, Hungarian Natural History Museum, H-1476 Budapest, Pf. 222, Hungary.
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Journal of Bryology (2008) 30: 170–173
# British Bryological Society 2008
Received 23 November 2007. Revision accepted 6 January 2008
DOI: 10.1179/174328208X281969
A contribution to the understanding of Heterocladium heteropterum
and its relationship with H. wulfsbergii
Nomenclatural issues and lectotypifications
The authorship of Heterocladium heteropterum is confused
due to the early history of its epithet. Despite several
deviating treatments (e.g. Crosby et al., 1999; Crum &
Anderson, 1981; Frey et al., 2006; Hill et al., 2006; Nyholm,
171
BRYOLOGICAL NOTES
1960; Smith, 2004), this species can only be understood as
having been originally described as new by Schimper (in
Bruch, Schimper & Gümbel, 1851–1855). The name is thus
not based on Pterigynandrum heteropterum Brid. (BridelBrideri, 1827), even if the interpretation of early treatments
is partly confusing. The name is also not based on
‘Pterogonium heteropterum Bruch ex Schwaegr.’ (cf.
Nyholm, 1960; Smith, 2004), since Schwaegrichen (1827)
referred to Pterigynandrum heteropterum and page 176 in
part 2 of Bridel-Brideri (1827) and it is therefore clear that
Pterogonium heteropterum is a new combination by
Schwaegrichen based on the Pterigynandrum name.
Schimper (in Bruch et al., 1851–1855) first treated
Pterigynandrum heteropterum as a variety of P. filiforme
Hedw. in fascicles 46–47 of Bryologia Europaea (1851).
Later, in the historical part of the generic protologue of
Heterocladium (fasc. 49–51, 1852), he expressly stated that
his H. heteropterum is one of the two new (‘neu’,
‘nouvelles’) species of the genus. At the same time, he
made it clear that Hypnum heteropterum (Brid.) Hampe,
based on Pterigynandrum heteropterum Brid., is not what he
then described as Heterocladium heteropterum but represents a variety of Pterigynandrum filiforme – ‘Hypnum
heteropterum der Auctoren ist durchaus nicht unser H.
heteropterum, sondern eine Varietät von Pterigynandrum
filiforme’. No doubt, the latter statement must refer to his
1851 treatment and thus proves definite exclusion of
Bridel’s taxon from his own Heterocladium heteropterum.
On the other hand, in connection with the description of
his Heterocladium heteropterum (fasc. 49–51, 1852), Schimper
(in Bruch et al., 1851–1855) cited ‘Hypnum heteropterum
(Schwaegr.) Rob. Spruce Ms.’. The Spruce material included
among the syntypes, no. 56 in Musci Pyrenaici (Spruce, 1847)
which label bears the name ‘Hypnum (heteropterum,
Schwaegr.)’ and ‘Pterogonium heteropterum Schwaegr.’
and no indication that Spruce considered this to be a new
species, thus proving that Spruce apparently believed that his
plant was the same as the one of Schwaegrichen, and
therefore in the end referring back to Bridel’s Pterigynandrum
heteropterum or Hypnum heteropterum (Brid.) Hampe
(Hampe, 1837). Nevertheless, as presented by Schimper (in
Bruch et al., 1851–1855), the Spruce reference can only mean
Hypnum heteropterum sensu Spruce (in sched., 1847), a mere
misapplication lacking any nomenclatural status, yet forming
quite essential element of Schimper’s new species (see the
lectotypification below). Accordingly, his cryptic Spruce
citation under Heterocladium heteropterum by no means
Table 1.
indicates indirect inclusion of Bridel’s Pterigynandrum
heteropterum.
Neither Schimper’s Heterocladium heteropterum nor
Bridel’s Pterigynandrum heteropterum appear to have been
lectotypified, and lectotypes are therefore selected here to
stabilise the use of these names.
Heterocladium
heteropterum
Schimp.,
Bryologia
Europaea 5: 154 (fasc. 49–51. Monogr. 2). 1852. Type:
[France] 56. Hypnum (heteropterum Schwaegr.). HAB. ad
saxa in sylvis Pyrenaeorum centralium; sat frequens sed
rarissime fructificans. Prope B.-de-Bigorre capsulis onustum legi 17 Octobris [no year indicated on label]. [Spruce,
Musci Pyrenaici] (BM-herb. Schimper, lectotype, selected
here by Hedenäs; BM isolectotype).
In herbarium Schimper in BM, only one of the syntypes
that are mentioned in the protologue of Heterocladium
heteropterum Schimp. exists (Len Ellis, pers. comment), the
specimen with sporophytes collected by Spruce in 1845
(according to Spruce, 1849) at Bagnères-de-Bigorre in the
Pyrenees as Musci Pyrenaici 56. This syntype represents
the particular gathering from which Schimper drew the
sporophyte description and illustrations of H. heteropterum
and is here selected as lectotype of the name; the BM
isolectotype is another Musci Pyrenaici 56 specimen. A
possible isosyntype, also with sporophytes, but collected in
January 1846, exists in BM-herb. W. Wilson.
Pterigynandrum filiforme Hedw., Species Muscorum
Frondosorum 81. 1801. Type: Specimen ‘a’ on a sheet with
reference to ‘Descr. IV, p. 18 t. VII [leg. Timm,...]’ (G-herb.
Hedwig-Schwaegrichen, lectotype; selected by Hedenäs &
Geissler, 1999).
Synonym: Pterigynandrum heteropterum Brid., Bryologia
Universa 2: 176. 1827. Hypnum heteropterum (Brid.)
Hampe, Flora 20: 274. 1837. P. filiforme var. heteropterum
(Brid.) Schimp., Bryologia Europaea 5: 122 (fasc. 46–47.
Monogr. 4). 1851. Type: [Switzerland] Pterigynandrum
curvatum/heteropterum. [Bryol. Univ.] In alpibus Valesia a
Sabadiaer Thomas 1820 (B-herb. Bridel, reg. no. B 31 0679
02, specimen ‘d’, lectotype, selected here by Hedenäs).
In Bridel’s herbarium in B syntypes of Pterigynandrum
heteropterum Brid. from all four localities mentioned in the
protologue are extant (Bridel-Brideri, 1827), and from Mt.
Hohack in the Vosges there are three specimens, thus in
total six syntypes. Five of these syntypes, including those
with spore capsules, belong to P. filiforme Hedw., in the
Italian specimen mixed with Heterocladium dimorphum
(Brid.) Schimp. One of the syntypes from the Vosges
Characters that differentiate the sporophytes of Heterocladium heteropterum from those of H. wulfsbergii.
Character
H. heteropterum
H. wulfsbergii
Capsule shape, when moist
Obloid or oblong, sometimes curved in
transition to seta
(2–)3–5
Of Brachythecium-type or oblong
2–3(–4)
White or pale whitish yellow
Red-brown to usually orange-brown, at least in basal part
Rows of differentiated exothecial cells
below capsule mouth
Exostome colour
172
BRYOLOGICAL NOTES
belongs to Heterocladium heteropterum Schimp. Among the
specimens belonging to P. filiforme Hedw. one with
sporophytes is selected as lectotype in order to choose a
specimen that is as complete as possible as regards the
presence of available characters.
At present, there appears to be no consensus regarding how
to treat the gametophytically relatively similar taxa
Heterocladium heteropterum Schimp. and H. wulfsbergii
I. Hagen. Even if most authors consider the two taxa as
distinct species (e.g. Casas et al., 2006; Crundwell & Smith,
2000; Hedenäs, 1992; Hill et al., 2006; Nyholm, 1960;
Smith, 2004), others treat H. wulfsbergii as a synonym of H.
heteropterum (e.g. Dirkse, Bouman & Losada-Lima, 1993),
or the two taxa are treated as varieties (e.g. Düll, 1992; Frey
et al., 2006) or subspecies (Jensen, 1939) of H. heteropterum. The history of these taxa was recently reviewed and
gametophytic characters that were considered useful for the
separation at the species level were described (Crundwell &
Smith, 2000). The various treatments of the taxa at levels
below the species are probably due to the rather wide
plasticity of the gametophyte combined with the fact that
sporophytes are rarely observed (cf. Braithwaite, 1896–
1905; Frey et al., 2006).
Most bryologists who saw sporophytes in either of these
species appear to have seen them in at most one or two
specimens, and often sporophyte descriptions found in the
literature are second-hand. Examples of descriptions of H.
heteropterum based on first-hand observations include the
following. (1) Based on Spruce material collected in the
Pyrenees, Schimper (in Bruch et al., 1851–1855) described
and illustrated the capsules as orthogonal (for terminology,
see Hedenäs, 2007) and described the peristome teeth as of
the kind described for the genus, that is, reddish orange
with paler distal parts. The description of the exostome
colour is a bit strange, since the lectotype that Schimper
could have seen has got a whitish exostome. (2) Braithwaite
(1896–1905) described capsules with pale peristomes and
stated that they are homotropous, but illustrated an almost
orthogonal capsule. (3) Jensen (1939) described the capsules
as orthogonal, and the exostome as pale. (4) Nyholm (1960)
described the spore capsules as homotropous to orthogonal
and illustrated homotropous to orthotropous ones. (5)
Smith (2004) described the capsule as homotropous to
orthogonal, and illustrated an orthotropous or almost
orthotropous capsule. The last author also described the
capsule of H. wulfsbergii, as homotropous to orthogonal.
Studies of Heterocladium heteropterum and H. wulfsbergii
in S, during collecting on Madeira, and of the original
material of H. heteropterum and a specimen of H.
wulfsbergii in herbarium Schimper in BM included four
H. heteropterum specimens with capsules and eight H.
wulfsbergii ones. Two of the latter were old and had no
peristome left, and could only be used to study capsule
orientation. This allowed a close comparison of the
sporophytes of the two taxa and revealed some differences
(Table 1, Fig. 1). The strikingly different exostome colours,
Figure 1. Sporophytes of (A) Heterocladium heteropterum (United
Kingdom: S, reg. no. B116465) and (B) H. wulfsbergii (Madeira,
Portugal): S, reg. no. B42888). Scale lines: 0.5 mm. Photo by
Christopher Reisborg.
whitish or pale whitish yellow in H. heteropterum and the at
least basally red- or orange-brown in H. wulfsbergii clearly
suggest that the two are different. On the other hand,
capsule orientation, and to some degree shape, is obviously
very variable within both species and cannot be used as a
reliable separating character. The sporophytic differences
found between these species provide important evidence
that supports their treatment as distinct species.
Studied specimens with sporophytes. H. heteropterum:
France: lectotype and possible isosyntype. Sweden:
Bohuslän, Lycke, P. A. Larsson, S, reg. no. B116467
(one single capsule). United Kingdom: Wales, Barmouth,
J. Rogers, S, reg. no. B116465. H. wulfsbergii: Azores
(Portugal): Terceira, H. Persson, S, reg. no. B10010
(one capsule, lacking peristome). Madeira (Portugal):
Porto do Moniz, L. Hedenäs, S, reg. no. B42888; Between
Ribeiro Frio and Portelo, L. Hedenäs, S, reg. no. B9971;
Ribeiro Frio to Santo da Serra, L. Hedenäs, S, reg. no.
B9974 (only basal portions of capsules); Ribeiro Frio to
BRYOLOGICAL NOTES
Santo da Serra, L. Hedenäs, S, reg. no. B9974 (several
capsules, but only two with peristome remains); Sao
Vicente, L. Hedenäs, S, reg. no. B119021; Sao Vicente, L.
Hedenäs, S, reg. no. B119022. United Kingdom: In loco
‘the Jumbles’ dicto, prope Bolton, Lancastriae, Ad rupes
arenaceas humidas et c[..]brosus, 5 January 1874, J.B.W.,
BM-herb. Schimper.
We thank L. Ellis for the BM loan and information, and H.
Nowak-Krawietz for the B loan. Suzana Sa Fontinha
kindly provided the collecting permit for some of the
Madeiran material studied.
TAXONOMIC ADDITIONS AND CHANGES: Nil.
REFERENCES
Braithwaite R. 1896–1905. The British moss-flora. Vol. 3. Pleurocarpi.
London: L. Reeve & Co.
Bridel-Brideri SE. 1827. Bryologia universa seu systematica ad novam
methodum dispositio, historia et descriptio omnium muscorum
frondosorum hucusque cognitorum cum synonymia ex auctoribus
probatissimis. Vol. 2. Lipsiae: Sumtibus Joan. Ambros. Barth.
Bruch P, Schimper WP, Gümbel T. 1851–1855. Bryologia Europaea seu
genera muscorum Europaeorum monographice illustrata. Vol. 5.
Stuttgartiae: Sumptibus Librariae E. Schweizerbart.
Casas C, Brugués M, Cros RM, Sérgio C. 2006. Handbook of mosses of
the Iberian Peninsula and the Balearic Islands. Barcelona: Institut
d’Estudios Catalans.
Crosby MR, Magill RE, Allen BH, He S. 1999. A checklist of the
mosses. St. Louis: Missori Botanical Garden.
Crum H, Anderson LE. 1981. Mosses of eastern North America. New
York: Columbia University Press.
Crundwell AC, Smith AJE. 2000. Heterocladium wulfsbergii I. Hagen in
the British Isles. Journal of Bryology 22: 43–47.
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Dirkse GM, Bouman AC, Losada-Lima A. 1993. Bryophytes of the
Canary Islands, an annotated checklist. Cryptogamie, Bryologie
Lichenologie 14: 1–47.
Düll R. 1992. Distribution of the European and Macaronesian mosses
(Bryophytina). Annotations and progress. Bryologische Beiträge 8/
9: 1–223.
Frey W, Frahm J-P, Fischer E, Lobin W. 2006. The liverworts, mosses
and ferns of Europe. Essex: Harley Books.
Hampe E. 1837. Musci frondosi germaniae ad methodum naturalem
dispositi. Allgemeine botanische Zeitung 18: 273–287.
Hedenäs L. 1992. Flora of Madeiran pleurocarpous mosses (Isobryales,
Hypnobryales, Hookeriales). Bryophytorum Bibliotheca 44: 1–
165.
Hedenäs L. 2007. Morphological characters and their use in pleurocarpous moss systematics. Systematics Association Special Volume
71: 221–239.
Hedenäs L, Geissler P. 1999. Lectotypification of Hedwig names:
holarctic pleurocarpous mosses. Candollea 54: 417–432.
Hill MO, Bell N, Bruggeman-Nannenga MA, Brugués M, Cano MJ,
Enroth J, Flatberg KI, Frahm J-P, Gallego MT, Garilleti R, Guerra
J, Hedenäs L, Holyoak DT, Hyvönen J, Ignatov MS, Lara F,
Mazimpaka V, Muñoz J, Söderström L. 2006. An annotated
checklist of the mosses of Europe and Macaronesia. Journal of
Bryology 28: 198–267.
Jensen C. 1939. Skandinaviens bladmossflora. Köbenhavn: Ejnar
Munksgard.
Nyholm E. 1960. Illustrated moss flora of Fennoscandia. II, Musci. Fasc.
4. Lund: C. W. K. Gleerup.
Schwaegrichen F. 1827. Ioannis Hedwig Species Muscorum
Frondosorum descriptae et tabulis aenis coloratis illustratae.
Opus posthumum. Suppl. 3. Vol. 1. Lipsiae: Sumptus Ioannis
Ambrosii Barth.
Smith AJE. 2004. The moss flora of Britain and Ireland. 2nd edn.
Cambridge: Cambridge University Press.
Spruce R. 1847. Musci Pyrenaici, quos in Pyrenaeis centralibus
occidentalibusque, nec non in agro syrtico, A. D. 1845–6. Londini.
Spruce R. 1849. The Musci and Hepaticae of the Pyrenees. Transactions
of the Botanical Society 2: 103–128.
LARS HEDENÄS, Swedish Museum of Natural History, Department of Cryptogamic Botany, PO Box 50007, SE-104 05 Stockholm, Sweden.
E-mail: lars.hedenas@nrm.se
PEKKA ISOVIITA, Botanical Museum, PO Box 7, FI-00014 University of Helsinki, Finland. E-mail: pekka.isoviita@helsinki.fi
Journal of Bryology (2008) 30: 173–174
# British Bryological Society 2008
Received 4 February 2008. Revision accepted 28 March 2008
DOI: 10.1179/174328208X300624
Lejeunea laii nom. nov. for Lejeunea ramulosa (Herzog) R.M.Schust.
(Jungermanniopsida: Lejeuneaceae) from Taiwan
The Lejeuneaceae, with about 1000 species in about
90 genera, is the largest family of liverworts
(Marchantiophyta) (Wilson et al., 2007) and the most
dominant group of the epiphyllous flora (Gradstein, 1997;
Zhu & So, 2001). Lejeunea Lib. is a predominantly
pantropical genus. It is characterized by: (1) proximal
hyaline papilla of the leaf lobule, (2) lack of brownish-black
pigments in cell wall, (3) usually seven rows of larger
cortical cells of the stem, (4) lejeuneoid leaf sequence of
gynoecial innovations, (5) ocelli lacking, (6) 26 elaters per
capsule with either seven or six elaters in opposite valves, in
a ‘1z2z2z2’ or ‘2z2z2’ pattern in a valve, and (7) lack
of specialized cladia. While compilation of an updated
checklist of liverworts and hornworts of Taiwan, we found
that Lejeunea ramulosa (Herzog) R.M.Schust. (Schuster,
1963), a poorly known species known only from the type
from Taiwan (Herzog & Noguchi, 1955), is an illegitimate
homonym. A new name is, therefore, proposed here.
Lejeunea laii R.L.Zhu, nom. nov.
; Microlejeunea ramulosa Herzog in Herzog & Noguchi,
J. Hattori Bot. Lab. 14: 51. 1955. ; Lejeunea ramulosa
(Herzog) R.M.Schust, Beih. Nova Hedwigia 9: 133. 1963.
[non Lejeunea ramulosa Spruce, Trans. Proc. Bot. Soc.
Edinburgh 15: 274. 1884.].
TYPE: China. Taiwan, ‘Formosa: loco incerto, leg. G. H.
Schwabe, no. ?, 1947’ (holotype: JE).
174
BRYOLOGICAL NOTES
Illustrations: Herzog & Noguchi (1955, p.53, Fig. 15: e–h
as Microlejeunea ramulosa).
The species epithet is dedicated to the late Prof. Ming-Jou Lai
(194922007) who was a bryologist and lichenologist in Taiwan.
Owing to its minute size Lejeunea laii was placed
originally in the genus Microlejeunea by Herzog and
Noguchi (1955). Microlejeunea is a controversial genus
which was treated as a subgenus of Lejeunea (Schuster,
1963). Some authors admitted generic rank to Microlejeunea, which is characterized by dioecy, the presence of
basal ocelli in the leaf lobes, small size, stems with only
three rows of medullary cells, and female bracts with long
keels showing a tendency to develop wings (cf. Jones 1979;
Grolle, 1995). But as in the typical members of Lejeunea,
Microlejeunea ramulosa lacks ocelli in leaf lobes and wings
at female bracts, and it also has larger leaves and over three
rows of medullary cells of the stem. Microlejeunea
ramulosa, thus, is no doubt a member of typical Lejeunea
s. str. A new name, Lejeunea laii, is needed owing to
existence of an earlier taxon with this name, Lejeunea
ramulosa Spruce.
As shown in Herzog & Noguchi (1955), Lejeunea laii is
characterized by: (1) very small size (only 0.4 mm wide with
leaves), (2) densely branched stems, (3) small underleaves
with very narrow lobes, and (4) obtuse or rounded-obtuse
apex of the leaf lobes. Lejeunea ramulosa Spruce, known
from Brazil, Costa Rica and Ecuador (Dauphin 2005;
Reiner-Drehwald, 1999, 2007), is immediately distinguished
from Lejeunea laii by its dioecy, large underleaves 324
times as wide as stem, acute to acuminate apex of the leaf
lobes, and more or less dentate keels of the perianth. In the
autoicous sexuality, deeply bilobed underleaves, obovate
perianths with five keels, and usually reduced leaf lobules,
Lejeunea laii is very similar to depauperate plants of
Lejeunea anisophylla Mont. common in Taiwan and
tropical Asia as well as Oceania, but the latter differs in
the rounded apex of leaf lobes, large trigones of leaf cells,
and larger size of the plant. In its obtuse to obtuse-rounded
apex of leaf lobes, small size and deeply bilobed underleaves, Lejeunea laii may be confused with Lejeunea cocoes
Mitt., known from China, Indonesia (Java), Malaysia
(Sarawak), and Papua New Guinea (cf. Zhu & So, 1999).
The latter, however, is separated by its dioecy, strongly or
slightly caducous leaves, and the presence of regenerants at
the leaf margin.
This research was sponsored by the National Natural
Science Foundation of China (no. 30670155).
TAXONOMIC ADDITIONS AND CHANGES: Lejeunea laii
R.L.Zhu is proposed for Lejeunea ramulosa (Herzog)
R.M.Schust, nom. illeg.
REFERENCES
Dauphin G. 2005. Catalogue of Costa Rican Hepaticae. Tropical
Bryology 26: 141–218.
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Islands. An annotated catalogue. Bryophytorum Bibliotheca 48: 1–
178.
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von Formosa und den benachbarten Inseln Botel Tabago und
Kwashyoto. Journal of the Hattori Botanical Laboratory 14: 29–
70.
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(Hepaticae) of Latin America. Bryophytorum Bibliotheca 54: 1–101.
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Brazil. http:// drehwald.info/Lejeunea/Key_Lejeunea_Brazil_30_Apr.
pdf.
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of Lejeuneaceae. I. Introduction; annotated keys to subfamilies
and genera. Beihefte Zur Nova Hedwigia 9: 1–203.
Wilson R., Gradstein SR, Schneider H, Heinrichs J. 2007. Unravelling
the phylogeny of Lejeuneaceae (Jungermanniopsida): evidence for
four main lineages. Molecular Phylogenetics and Evolution 43: 270–
282.
Zhu R-L, So ML. 1999. Additions of Lejeuneaceae taxa to the hepatic
flora of Yunnan, China. Annales Botanici Fennici 36: 219–229.
Zhu R-L, So ML. 2001. Epiphyllous liverworts of China. Nova
Hedwigia Beiheft 121: 1–418.
JIAN WANG and RUI-LIANG ZHU*, Department of Biology, School of Life Science, East China Normal University, 3663 Zhong Shan North
Road, Shanghai 200062, China. Email: lejeunea@163.com
*Corresponding author
Journal of Bryology (2008) 30: 174–177
# British Bryological Society 2008
Received 17 October 2007. Revision accepted 7 April 2008
DOI: 10.1179/174328208X300642
Lejeuneaceae (Marchantiophyta) share an indel in chloroplast rps4
gene
The Lejeuneaceae, with the estimated more than 1000
species and 91 genera, are the most diversified family of
liverworts (Gradstein, Reiner-Drehwald ME & Schneider,
2003). Recent molecular studies using three (ITS2, trnL-F
and rbcL, Ahonen, Muona & Piippo, 2003), two (rbcL and
nad4, Wilson et al., 2004) and four (rbcL, psbA, trnL-F and
ITS, Wilson et al., 2007a) loci, respectively, have demonstrated that the family is monophyletic.
The chloroplast rps4 gene has been used widely
for molecular phylogenetic studies of bryophytes. When
BRYOLOGICAL NOTES
analyzing an alignment of rps4 sequences for liverworts,
we noticed an indel (insertion/deletion) shared by all taxa
of Lejeuneaceae. A search of all liverworts rps4 sequences
on GenBank, did not reveal this indel in any taxa outside
the Lejeuneaceae. In order to test if the indel is a marker
for the Lejeuneaceae, a data set containing 93 rps4 sequences
from 91 liverwort genera was generated. All sequences
were obtained from GenBank (accession numbers are listed
in the Appendix); they mainly come from the following
studies: Davis (2004), He-Nygrén et al. (2004), Forrest &
Crandall-Stotler (2004), Forrest et al. (2005), de Roo et al.
(2007), Hendry et al. (2007) and Liu et al. (in press). In our
data set, 13 species and 11 genera of Lejeuneaceae were
included.
The sequence alignment was made using CLUSTAL X
1.83 (Thompson et al., 1997) and adjusted manually in
BioEdit 5.0.9 (Hall, 1999). In order to detect whether this
indel affects selection on the gene, nonsynonymous-synonymous substitution rate ratios (dN/dS) were calculated
using codeml program of PAML v3.15 (Yang, 1997) for
Lejeuneaceae, and also for other liverwort taxa.
The rps4 data matrix included 618 sites. The indel is a 9base-pair (bp) deletion occurring in all members of
Lejeuneaceae (Fig. 1). It is noteworthy that the deletion
does not occur in Nipponolejeunea nor in Jubulaceae but
was found in Bryopteris. Traditionally, Nipponolejeunea has
been assigned to the Lejeuneaceae (Mizutani, 1961;
Schuster, 1963; Gradstein, 1994, 2001; Crandall-Stotler &
Stotler, 2000) based mainly on the basis of leaf morphology: the lobule attaches to the lobe along a long keel. Weis
(2001) and Gradstein et al. (2003) described cladistic
analyses based on morphological characters, and both
authors recognized Nipponolejeunea as an early-diverging
lineage of Lejeuneaceae. However recent molecular analyses (e.g. Ahonen et al., 2003; Davis, 2004; Ahonen, 2006;
He-Nygrén et al., 2004, 2006; Wilson et al., 2004, 2007a;
Heinrichs et al., 2005; Forrest et al., 2006) have all
suggested this genus is more closely related to the
Jubulaceae. In fact, more than 50 years ago, Mizutani
(1961) placed Nipponolejeunea close to Jubula based on
morphological characters such as red alar cells in the capsule
wall and one or two archegonia per perianth. CrandallStotler and Guerke (1980) also indicated that
Nipponolejeunea appears to have a typical Lejeunea-type
lobule as in the Lejeuneaceae , but is only attached to the stem
narrowly. Bryopteris (Nees) Lindenb. was placed in its own
family (Bryopteridaceae) by Stotler & Crandall-Stotler (1974)
and Crandall-Stotler & Stotler (2000) because of its unique
branching type. However molecular (Ahonen et al., 2003;
Davis, 2004; He-Nygrén et al., 2004, 2006; Wilson et al., 2004,
2007a; Heinrichs et al., 2005; Forrest et al., 2006) and
morphological cladistic studies (Weis, 2001; Gradstein et al.,
2003) both resolve Bryopteris as a member of Lejeuneaceae.
The distribution of the 9 bp indel provides support for the
view that Bryopteris is a member of the Lejeuneaceae, but
Nipponolejeunea shares a more recent common ancestor with
genera in the Jubulaceae rather than the Lejeuneaceae.
175
Figure 1. The portion of the aligned matrix of rps4 showing the
unique indel shared by Lejeuneaceae. Dots indicate nucleotides identical to the top sequence, and dashes indicate gaps. The codon
grouping shown is the correct reading frame.
rps4 is a chloroplast gene located in the large single copy
region of the chloroplast genome. It encodes the chloroplast
ribosomal protein S4, which is one of the primary rRNA
binding proteins (Harris et al., 1994). As the deletion in
Lejeuneaceae rps4 gene dose not cause a frame shift of
176
BRYOLOGICAL NOTES
codon reading frames, no functional influence is expected.
The analyses of synonymous and non-synonmous substitutions in the rps4 gene suggests that there is no obvious
difference in substitution patterns between Lejeuneaceae
(dN/dS50.07) and other liverwort taxa (dN/dS50.08). It
appears that evolutionary constraints on the rps4 gene are
not relazed in the Lejeuneaceae.
Molecular approaches known as ‘DNA barcoding’
(Tautz et al., 2002; Pennisi, 2003; Hajibabaei et al., 2006)
are helping to identify species and classify taxa. The
exclusive indel found in the Lejeuneaceae maybe can be
used to help identifying members of this large family. The
lineages of this species-rich family appear to have accumulated more or less constantly through the Tertiary (Wilson
et al., 2007b). Whether all members of this monophyletic
group possess the indel, or if the indel occurs in other
liverworts remains unknown. More rps4 sequences are
needed for taxa in the Lejeaneaceae and other families.
APPENDIX
rps4 sequences used in this study and their Genbank
accession numbers.
Acromastigum exile, AY608041; Adelanthus lindenbergianus, AM398292; Anastrepta orcadensis, AM398339;
Anastrophyllum tubulosum, AM398237; Anthelia julacea,
AY608044; Aphanolejeunea gracilis, AY462338; Ascidiota
blepharophylla, AY608046; Austrolejeunea nudipes,
AY462339;
Balantiopsis
diplophylla,
AY608047;
Barbilophozia quadriloba, AM398375; Bazzania sp.,
AY608048; Blepharidophyllum densifolium, AM398306;
Bryopteris filicina, AY608051; Calypogeia integristipula,
AM398361;
Cephalozia
lunulifolia,
AM398315;
Cephaloziella varians, AM398310; Ceratolejeunea coarina,
AY608055; Chiloscyphus appalachianus, AY608056;
Clasmatocolea vermicularis, AM398246; Cryptochila paludosa, AM398275; Cyclolejeunea convexistipa, AY608058;
Delavayella serrata, AM398305; Diplophyllum taxifolium,
AM398354; Douinia ovata, AM398353; Fossombronia
pusilla, AY608062; Frullania tamarisci, AY462349;
Gackstroemia weindorferi, AY608063; Geocalyx graveolens, AM398367; Goebeliella cornigera, AY608064;
Gongylanthus reniformis, AM398261; Gymnocolea inflata,
AM398321; Gymnocoleopsis multiflora, AM398251;
Gymnomitrion corallioides, AM398302; Harpanthus flotovianus, AM398357; Herbertus aduncus, AM398291;
Herzogobryum teres, AY608072; Isotachis lyallii,
AY608073; Jamesoniella colorata, AM398314; Jensenia
decipiens, AY734695; Jubula pennsylvanica, AY608075;
Jubulopsis novae-zelandiae, AY608076; Jungermannia
exsertifolia, AM398369; Leiocolea rutheana, AM398345;
Leiomylia anomala, AM398269; Lejeunea catanduana,
AY462363; Lejeunea cavifolia, DQ787470; Lejeunea cladogyna, AY608079; Lepidogyna hodgsoniae, AY462366;
Lepidolaena sp., AY608082; Lepidozia cupressina,
AM398265; Leptoscyphus ovatus, AM398248; Lethocolea
congesta, AM398267; Lophocolea difformis, AM398352;
Lophozia perssonii, AM398374; Marsupella aquatica,
AM398281;
Marsupidium
latifolium,
AY608088;
Mastigolejeunea auriculata, AY462371; Mastigophora
woodsii, AM398298; Mesoptychia sahlbergii, AM398328;
Metzgeria decipiens, AM398259; Mnioloma fuscum,
AM398263; Mylia taylorii, AM398270; Nardia scalaris,
AM398273;
Neesioscyphus
argillaceus,
AM398235;
Neotrichocolea bissetii, AY608093; Nipponolejeunea pilifera, AY462376; Nowellia curvifolia, AM398293;
Odontolejeunea lunulata, AY608095; Odontoschisma
denudatum, AM398325; Pachyschistochila carnosa,
AM398296; Pallavicinia xiphoides, AY734700; Pellia epiphylla, AY507457; Plagiochila acanthocaulis, AM180486;
Plicanthus hirtellus, AM398304; Porella pinnata,
AY608101; Pseudolepicolea quadrilaciniata, AM398299;
Ptilidium ciliare, AY608103; Ptychanthus striatus,
AY462389; Radula complanata, AM398295; Riccardia
capensis, AM398264; Scapania compacta, AM398312;
Schistochila lehmanniana, AY608109; Southbya gollanii,
AM398294;
Sphenolobopsis
pearsonii,
AM398316;
Stenorrhipis madagascariensis, AY608111; Symphyogyna
podophylla, AM398258; Syzygiella setulosa, AM398256;
Telaranea pulcherrima, AY608114; Temnoma pulchellum,
AY608115;
Tetralophozia
setiformis,
AM398370;
Triandrophyllum subtrifidum, AY608117; Trichocolea
tomentosa,
AY608118;
Trichocoleopsis
sacculata,
EF460703; Tritomaria polita, AM398376; Trocholejeunea
sandvicensis, AY462401.
We thank R. Wilson for her valuable communion. Thanks
are also given to W. Wang and R.Q. Li for their suggestions. We specially thank A.J. Shaw for his help of
improving the language of the manuscript.
TAXONOMIC ADDITIONS AND CHANGES: Nil.
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BRYOLOGICAL NOTES
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YANG LIU and YU JIA, Institute of Botany, the Chinese Academy of Sciences, Beijing, 100093, China. Email: yjia@ibcas.ac.cn