Cryptogamie, Bryologie, 2009, 30 (2): 265-269
© 2009 Adac. Tous droits réservés
Fossombronia caespitiformis De Not. ex Rabenh. subsp.
multispira (Schiffn.) J.R. Bray et Cargill in Belgium,
a remarkable extension of its European range
André SOTIAUXa,b, Herman STIEPERAEREb*,
Odette SOTIAUXc & Henri POHLd
aUniversité
bNational
de Liège, Institut de Botanique, B22 Sart Tilman, B-4000 Liège.
Botanic Garden of Belgium, Domein van Bouchout, B-1860 Meise
cChaussée
dPlace
de Bruxelles 676, B-1410 Waterloo
de Baileux 33, B-6464 Baileux
(Received 10 September 2008, accepted 8 October 2008)
Abstract – Fossombronia caespitiformis subsp. multispira has been found at Gozée
(province of Hainaut, Belgium), 330 km outside its known range. This subspecies is
characterised by elaters having 3-4 spirals. The plants from Gozée correspond well with the
original description of Fossombronia husnotii, a synonym of this subspecies. As the rhizoids
are purple, not colourless as those traditionally described for F. husnotii, rhizoid colour is
not a reliable character to distinguish F. caespitiformis subsp. multispira.
Résumé – Fossombronia caespitiformis subsp. multispira a été découvert à Gozée (province
de Hainaut, Belgique), 330 km en dehors de son aire connue. Cette sous-espèce est
caractérisée par des élatères à 3-4 spires. La morphologie du matériel belge correspond
bien à la description originale du F. husnotii décrit par Corbière en 1904, un synonyme de
cette sous-espèce. Comme les rhizoïdes sont pourpres, non hyalins comme ceux
habituellement décrits pour F. husnotii, la couleur des rhizoïdes n’est pas un caractère
fiable pour distinguer F. caespitiformis subsp. multispira.
Fossombroniaceae / Fossombronia caespitiformis / Fossombronia husnotii / rhizoid colour
/ Belgium
INTRODUCTION
In the recent checklist of Belgian bryophytes (Sotiaux et al., 2007) several
species have been added to the list. This was mainly due to intensive prospecting
in Southern Belgium. Two species considered extinct were rediscovered,
Philonotis marchica (Hedw.) Brid. and Pterygoneurum lamellatum (Lindb.) Jur.,
* Correspondence and reprints : Stieperaere@br.fgov.be
266
A. Sotiaux, H. Stieperaere, O. Sotiaux & H. Pohl
both not seen for nearly a century. One of the species that still remained on the
‘hit list’ was Fossombronia caespitiformis De Not. ex Rabenh., found once in 1903
between Lives and Bossimé (Prov. Namur) in the Belgian district of the Meuse
(Mansion, 1905). This find has been much debated. Düll (1983) listed this species
for Belgium and indicated that he had seen material. However, the only specimen
is (now) sterile and even lacks perianths. De Zuttere & Schumacker (1984) did
therefore not accept the presence of this submediterranean species in their
checklist and it remained absent from the subsequent checklists and in the flora
of Siebel & During (2006). Nevertheless Schumacker & Vá≈a (2005) follow Düll
(1983) and list F. caespitiformis for Be, the abbreviation for the geographical
territories of both Belgium and the Grand Duchy of Luxembourg.
When in June 2008 A.S. studied a Fossombronia specimen he just had
collected, with O.S. & H.P, on the side of a ditch between a wheat field and a
forest near Gozée (Prov. Hainaut, Belgium), he found the characteristic spores of
F. caespitiformis. However, spore diameter of the recent collection (32-35 µm) was
much lower than that of typical F. caespitiformis. We decided therefore to study
the specimen in more detail.
MORPHOLOGY AND ECOLOGY OF THE BELGIAN SPECIMEN
Gametophyte resembling F. pusilla (L.) Nees, rhizoids violet. Spores well
developed, (30.3) 32-35 (37.4) µm in diameter, distally densely covered with
4-6 µm long truncate, flattened spines, broadened at the base to form a star like
pattern, some spines confluent at their base but not forming short lamellae pattern
(Figs 1-3); spores proximally without a clear triradiate mark, covered with low,
irregular alveolae (Fig. 4). Elaters with 3 (4) spirals, exceptionally bispiralled.
Belgium, province of Hainaut, Gozée, 50°19’37”N 4°21’11”E; IFBL/ H4.25.23, UTM/
ER9676, alt. 185 m, 26-06-2008, leg A. Sotiaux n° 36.552 (herb. Sotiaux)
Gozée is situated in the Mosan district, a loam plateau often overlying
calcareous deposits that are exposed in deep valleys. The locality near the
artificial lake of the Grand Vivier forms part of the pure loam plateau devoted
largely to agriculture. The north side of the drain where F.caespitiformis was
found had a varied flora. Next to the Fossombronia plants grew large colonies of
Phaeoceros carolinianus (Michx.) Prosk. with Riccia glauca L., Amblystegium
serpens (Hedw.) Schimp., Barbula unguiculata Hedw., Bryum violaceum Crundw.
et Nyholm, Dicranella schreberiana (Hedw.) Dixon, D. staphylina H.Whitehouse,
Phascum cuspidatum Hedw., Physcomitrium pyriforme (Hedw.) Bruch et Schimp.,
Pohlia melanodon (Brid.) A.J.Shaw and Tortula truncata (Hedw.) Mitt.
DISCUSSION
Fossombronia caespitiformis is a very variable taxon. Stotler et al. (2003)
distinguish two subspecies: F. caespitiformis subsp. caespitiformis and F.
caespitiformis subsp. multispira (Schiffn.) J.R.Bray et D.C.Cargill. The only
diagnostic character given for F. caespitiformis subsp. multispira is that the elaters
Fossombronia caespitiformis in Belgium
267
Figs 1-4. Fossombronia caespitiformis subsp. multispira, SEM-micrographs of spores (from
Sotiaux 36552). 1-3. Distal view. 4. Proximal view.
are consistently 3-5 spiralled, rarely bispiralled, whereas in subsp. caespitiformis
they are consistently bispiraled, rarely trispiraled centrally. They also reduce
F. husnotii Corb. to synonymy of F. caespitiformis subsp. multispira.
Corbière (1889) distinguished his F. husnotii from F. caespitiformis on
spore characters (spores with lamellae elevated at the intersection of the vague
and incomplete alveolae; creating a star like pattern on the surface) and on the
3-4 spiralled elaters. He did not mention rhizoid colour. He eventually reduced
F. husnotii to F. caespitiformis var. husnotii (Corb.) Corb. because of the many
overlaps in the spore characters of both taxa (Corbière, 1904).
Spore diameter is much lower than the figures for F. caespitiformis spores
given in Paton (1999) and Schumacker & Vá≈a (2005): 38-56(64) µm, rarely
30-40 µm, resp. 42-65 µm, especially since Paton (1999) states that the small
spored form has spines and tubercles largely reduced to conical papilla. The spore
diameter of F. husnotii given in Corbière (1889) is only ca. 40 µm compared to
45-50 µm for F. caespitiformis. Also, in his original description of F. caespitiformis
268
A. Sotiaux, H. Stieperaere, O. Sotiaux & H. Pohl
var. multispira Schiffner (1918†1) indicates that the spores are smaller than those
of the type variety “Differt a typo sporis minoribus, …”.
The 3(4)-spiralled elaters, and the smaller spore diameter of the Belgian
collection compared to typical F. caespitiformis var. caespitiformis makes us
confident that this is F. caespitiformis subsp. multispira. Also, spore ornamentation
of this collection matches very well the description and plate of its synonym
F. husnotii (Corbière, 1889).
Since Müller (1909), European authors (e.g. Paton, 1999) have stressed
the colourless or pale brown rhizoids and the variable spores of F. husnotii with
irregular truncate papillae, sometimes united to irregular lamellae or irregular
alveolae on the distal face. Thus, Schumacker & Vá≈a (2005) who follow the
nomenclature of Stotler et al. (2003) and Stotler & Crandall-Stotler (2007), still
key out F. caespitiformis subsp. multispira on the colourless to pale brown rhizoids
and variable spore ornamentation and do not mention the number of spirals in the
elaters.
Stotler et al. (2003) and Stotler & Crandall-Stotler (2007), however, do
not mention rhizoid colour, only the number of spirals in the elaters. As our find
shows, rhizoid colour is not a reliable character to distinguish F. caespitiformis var.
multispira. In fact, the rhizoids of two Dutch Fossombronia wondraczekii
collections were consistently brown to colourless (During in Gradstein & van
Melick, 1996). Fossombronia taxa with variable rhizoid pigmentation between
populations and even on individual plants have been described from Australia
(e.g. F. scrobiculata G.A.M.Scott & D.C.Pike and F. vermiculata G.A.M.Scott &
D.C.Pike; Scott & Pike, 1984) and from South Africa (F. cederbergensis Perold;
Perold, 1998).
The habitat at Gozée is rather nondescript and typical of intensive arable
fields. Mansion (1905) found F. caespitiformis in a calcareous field in Bossime,
between Live and Erpent, 4 km SE of Namur, Prov of Namur, in the Mosan
district, in the company of Fossombronia pusilla, Riccia glauca, Anthoceros
agrestis Paton and Phaeoceros laevis (L.) Prosk. It is therefore difficult to
understand why F. caespitiformis s.l. is so rare. Hill et al. (1991) also can not easily
explain the scarcity of this species in England in terms of its habitat requirements,
but think this may reflect that it is at its climatic limit.
Mansion (1905) wrote that his specimens were abundantly fertile; the
specimen at BR is sterile. Until another fertile specimen is found, this recent find
definitely documents F. caespitiformis s.l. for Belgium but makes the find of
Mansion more plausible.
Both subspecies of F. caespitiformis are present in nearly all the
Mediterranean countries (Ros et al., 2007) and in Madeira and the Canaries.
Along the Atlantic coast it is rare from Portugal to France up to the English
Channel and in Southern England (Hill et al., 1991). Fossombronia caespitiformis
subsp. caespitiformis is also present in Australia (McCarthy, 2006). The nearest
localities to Gozée are in Normandy (Villons-les-Buissons, at 365 km; Lecointe,
1978) and in Southern England (at ca. 330 km; Hill et al., 1991). Because of the
large overlap in distribution of the two subspecies of F. caespitiformis these could
also be treated as varieties, F. caespitiformis var. husnotii then has priority.
† The publication date of Fossombronia caespitiformis var. multispira is not 1917 as indicated by Stotler et al.
1.
(2003) but 1918. See page 113 of Oesterreichische Botanische Zeitschrift 67.
Fossombronia caespitiformis in Belgium
269
Acknowledgements. We thank Marcel Verhaeghen and Frank Van
Caekenberghe (Meise) for help with SEM micrographs and Quintin Groom (Meise), David
Long (Edinburgh) and an anonymous referee for constructive comments. The Walloon
Region DGRNE, convention C81, supported fieldwork.
REFERENCES
CORBIÈRE L., 1889 — Muscinées du Département de la Manche. Mémoires de la société nationale
des sciences naturelles et mathématiques de Cherbourg 26: 195-368.
CORBIÈRE L., 1904 — Contribution à la flore bryologique de l’Algérie. Revue bryologique 31: 31-42.
DE ZUTTERE PH. & SCHUMACKER R., 1984 — Bryophytes nouvelles, méconnues, rares,
menacées ou disparues de Belgique. Service de la conservation de la nature, Travaux 13:
1-160 + 9 fig., 40 distributionmaps.
DÜLL R., 1983 — Distribution of European and Macaronesian liverworts (Hepaticophytina).
Bryologische Beitraege 2: 1-115.
GRADSTEIN S.R. & VAN MELICK H.M.H., 1996 — De Nederlandse levermossen en hauwmossen.
Flora en verspreidingsatlas van de Nederlande hepaticae en Anthocerotae. Utrecht, Stichting
Uitgeverij van de Koninklijke Nederlandse Natuurhistorische Vereniging, 366 p.
HILL M.O., PRESTON C.D. & SMITH A.J.E., 1991 — Atlas of the Bryophytes of Britain and Ireland.
Vol. 1. Liverworts (Hepaticae and Anthocerotae). Colchester, Harley Books, 351 p.
LECOINTE A., 1978 — Les associations bryologiques des éteules en Normandie (France). Documents
phytosociologiques, nouvelle série, 2: 283-300.
MANSION A., 1905 — Note sur le Fossombronia caespitiformis De Not., espèce nouvelle pour la flore
belge. Bulletin de la société royale de botanique de Belgique 42(2): 65.
MCCARTHY P.M., 2006 — Checklist of Australian Liverworts and Hornworts. Australian Biological
Resources Study, Canberra. Version 6 April 2006. http://www.anbg.gov.au/abrs/liverwortlist/
liverworts_intro.html. Assessed August 2008.
MÜLLER K., 1909 — Die Lebermoose Deutschlands, Oesterreichs u. d. Schweiz. In Rabenhorst’s
Kryptogamenflora von Deutschland, Oestereich und der Schweiz, 2 edit., Leipzig, 6(7): 385448.
PATON J.A., 1999 — The liverwort flora of the British Isles. Colchester, Harley Books, 626 p.
PEROLD S.M., 1998 — Studies in the liverwort genus Fossombronia (Metzgeriales) from Southern
Africa. 5. A new species from Northern and Western Cape. Bothalia 28: 1-5.
ROS R.M., MAZIMPAKA V., ABOU-SALAMA U., ALEFFI M., BLOCKEEL T.L., BRUGUÉS M.,
CANO M.J., CROS R.M., DIA M.G., DIRKSE G.M., EL SAADAWI W., ERDA∆ A.,
GANEVA A., GONZÁLEZ-MANCEBO J.M., HERRNSTADT I., KHALIL K.,
KÜRSCHNER H., LANFRANCO E., LOSADA-LIMA A., REFAI M.S., RODRÍGUEZNUÑEZ S., SABOVLJEVI± M., SÉRGIO C., SHABBARA H., SIM-SIM M. &
SÖDERSTRÖM L., 2007 — Hepatics and Anthocerotes of the Mediterranean, an
annotated checklist. Cryptogamie, Bryologie 28: 351-437.
SCHIFFNER V., 1918 — Hepaticae Baumgartnerianae dalmaticae. III. Das Zaratiner Gebiet.
Oesterreichische Botanische Zeitschrift 27: 147-156.
SCHUMACKER R. & VÁ~A J., 2005 — Identification keys to the liverworts and hornworts of Europe
and macaronesia, second and fully revised edition. Pozna¬, Sorus, 209 p.
SCOTT G.A.M. & PIKE D.C., 1984 — New Species of Fossombronia from Australia. Journal of the
Hattori botanical laboratory 56: 339-349.
SIEBEL H. & DURING H., 2006 — Beknopte mosflora van Nederland en België. Utrecht, KNNV
Uitgeverij, 559 p.
SÖDERSTRÖM L., URMI E. & VÁ~A J., 2007 — The distribution of Hepaticae and Anthocerotae
in Europe and Macaronesia - Update 1-427. Cryptogamie, Bryologie 28:299-350.
SOTIAUX A., STIEPERAERE H. & VANDERPOORTEN A., 2007 — Bryophyte checklist and
European red list of the Brussels-Capital Region, Flanders and Wallonia (Belgium).
Belgian journal of botany 140 : 174-196.
STOTLER R.E. & CRANDALL-STOTLER B.J., 2007 — Worldwide Monograph of the Suborder
Fossombroniineae, http://bryophytes.plant.siu.edu/fossombronia.html, website assessed
6 Aug 2008.
STOTLER R.E., BRAY J.R. Jr, CARGILL D.C., KRAYESKI D. & CRANDAL-STOTLER B.J.,
2003 — Typifications in the genus Fossombronia (Marchantiophyta). The bryologist 106:
130-142.