Bol. Soc. Esp. Briol. 46-47: 5-10 (2016) CINCLIDIUM STYGIUM SW. (MNIACEAE, BRYOPSIDA) IN EASTERN PYRENEES Vincent Hugonnot Le bourg, 43380 Blassac, France. E-mail: vincent.hugonnot@wanadoo.fr Abstract: Cinclidium stygium Sw. is recorded for the first time in the Pyrenees (PyrénéesOrientales, France) which is a significant south-western extension of its known range, approximately 400 km. The new locality is described, and recommendations are made for conservation of the species. Resumen: Se cita Cinclidium stygium Sw. por primera vez en los Pirineos (Pyrénées-Orientales, Francia), lo que supone una extensión significativa de su área de distribución hacia el suroeste de aproximadamente 400 km. Se describe la nueva localidad y se dan recomendaciones para la conservación de la especie. Keywords: Mosses, Cinclidium stygium, rich-fen relict, Pyrenees, ecology, Madres. INTRODUCTION In the course of bryological surveys in the eastern part of the Pyrenees, several bryophytes of phytogeographical interest were found. Among them, Cinclidium stygium Sw. deserves to be mentioned because it had not been previously recorded south of the Alps, which means a significant southwards extension of its known range. The worldwide distribution of Cinclidium stygium was reviewed by Mogensen (1973). This species has a circumpolar distribution in boreal and low-arctic regions in the northern hemisphere, and a remarkable disjunction in South America. In the Holarctic, its range extends farther south than the other species of Cinclidium, none of which has been recorded in southern Europe. Conversely, it is rare or absent in high-arctic regions. In North America, it is widespread in Canada (Koponen, 1974), from Yukon to Quebec. It is locally frequent in Alaska and Greenland. The species is widespread in Fennoscandia (Koponen, 1969), in Altaï Mountains (Ignatov, 1994), and Kamtschatka (Czernyadjeva & Ignatova, 2008), in Switzerland (http://www.nism.uzh.ch/map/map_en.php), of local occurrence in the north of the British Isles and Ireland (Blockeel et al., 2014), in Denmark and Germany (Meinunger & Schröder, 2007), as well as in Poland (Karczmarz, 1962). Its southwestern limit lies in French Alps, where it is very rare (Sarrassat, 1939). In this country the most important populations are in the Jura (Hétier, 1896; Hillier, 1954). Cinclidium stygium is experiencing a gradual decline, particularly in the southern part of its range. It has disappeared from several countries (Netherlands, Belgium), and is hence of conservation concern. It is for example VU in Ireland, CR in Slovakia, EN in Romania (Hodgetts, 5 Boletín de la Sociedad Española de Briología 46–47 (2016) 2015). In France it is protected under the law in the region of Franche-Comté (Arrêté of 22 June 1992, Journal Officiel of 4 August 1992). This paper intends to report the presence of an isolated population of Cinclidium stygium in the Pyrenees, as well as to describe its ecology and discuss conservation issues. STUDY AREA Madres-Coronat massif occupies a position between the Mediterranean Sea and the first high summits of the Pyrenean range. The Pic Madres culminates at 2469 m. It is settled less than 60 km from the Mediterranean Sea. The geologic bedrock is mostly composed of schists and granite, with veins of granodiorite and other magmatic rocks very rich in Ca, Mg, K, etc. The summit plateau has been carved by glaciers which formed typical U-shaped valleys, where horizontal sections and drops in elevation alternate. The plateau has been fractured by erosion, and the cracks are filled with detritic granitic sand that plays the role of an aquifer restituting slowly accumulated large quantities of water. This may account for the particular abundance of peaty habitats on the slopes of the massif. The climate is marked by oceanic influence but strongly modified by altitude. The annual mean precipitation is 1500-1900 mm at Madres summit. The summer may experience violent storms. The mean annual temperature is low (around 3°C at 2300 m). Snow is abundant in winter and may last from November to April in alpine habitats. Cold and dry winds are very frequent. MATERIAL AND METHODS This study is based upon examination of fresh and dry collections of plants referable to Cinclidium stygium from France. The material collected in Pyrénées-Orientales was studied immediatly after collecting while still fresh. Dry material previously collected by the author in Southern Alps and Jura was used for morphological comparison. The new locality is as follows: FRANCE: Pyrénées Orientales, Mosset, Madrès range, La Balmette d’Avall, 2150 m, in alpine fens, 42º39’027'‘N; 2º12’013'‘W. Leg. V. Hugonnot, 21 October 2015, Herbarium PyrénéesOrientales 7598, without reproductive organs. The morphological characters have been assessed in the course of our study from traditional examination of both dry and fresh plants. The voucher specimens are deposited in the author’s private herbarium. 6 HUGONNOT: CINCLIDIUM STYGIUM SW. (MNIACEAE, BRYOPSIDA) IN EASTERN PYRENEES Nomenclature of liverworts and mosses follows, respectively, Ros et al. (2007) and Ros et al. (2013). RESULTS The studied material has been confirmed to belong to Cinclidium stygium. However, it has been found rather variable as far as gametophytes are concerned. Frequently, the plants appeared with the vegetative parts depauperate, with small leaves, from 1.5 to 2 mm, of ovate to elliptical outline and an acute apex. The largest leaves (from 3 to 4.5 mm) were oblong to obovate-elliptic, longer than wide and sharply apiculate. The best developed ones were distinctly petiol-like at base and had an elongated areolation. The margin was uniformly unistratose. Old portions of stem were densely tomentose. Macronemata were initiated as indistinct longitudinal lines at the apex of shoots. No gametangia were observed. At 2 cm deep in the colony, the leaves were mostly damaged and reduced to their thick nerves, the lamina cells being destroyed. The colony was composed of an extensive red-blackish mat of densely interwoven stems, packed into a dense tomentum. The mat was strikingly superficial, not deep, as it is often the case in floating fens. Here, it did not exceed 2 cm deep, and grew upon a dense and black peat where no fragments of plants were recognizable. Cinclidium stygium grew on a very gentle slope (less than 5 deg. of inclination) in a subalpine Trichophorum cespitosum (L.) Hartm. fen, attributable to Caricion davallianae Klika 1934. Associated bryophytes were not abundant, with Scorpidium cossonii (Schimp.) Hedenäs, Ptychostomum pseudotriquetrum (Hedw.) J.R.Spence & H.P.Ramsay (C. sp.), Imbribryum muehlenbeckii (Bruch & Schimp.) N. Pedersen, Campylium stellatum (Hedw.) Lange & C.E.O.Jensen, Palustriella falcata (Brid.) Hedenäs, Sphagnum teres (Schimp.) Ångstr., and Calliergon richardsonii (Mitt.) Kindb. Fifteen square meters were occupied by Cinclidium stygium. Three main colonies were observed in distinct small similar fens. Several hundred meters of unoccupied fens were also present in the surroundings. DISCUSION Determination of material The genus Cinclidium includes 4 species (Mogensen, 1973). The distinction between them is generally straighforward taking into consideration sexual condition, leaf shape and capsule characters. Then, difficulties may arise when only sterile material is at hand, which is the case here. In particular, the distinction of Cinclidium stygium and C. articum (Bruch & Schimp.) Schimp. may raise serious problems. The areolation of Cinclidium stygium is longer (lamina cells more than two times as long as wide) than that of C. arcticum (less than two times as long 7 Boletín de la Sociedad Española de Briología 46–47 (2016) as wide). The leaf apiculum is long squarrose-pointed in Cinclidium stygium while being shortly acuminate in C. arcticum, which is a northern species that has been reported from Germany in recent times (Meinunger & Schröder, 2007). Reproduction and Ecology Cinclidium stygium is a synoicous species whose sporophytes are occasional. They are frequent in Jura or Southern Alps (pers. obs.), where they were considered frequent in the past (Hétier, 1896; Hillier, 1954). Absence of sporophytes in monoicous taxa may relate to the absence of gametangia, inefficient fertilization or abortion of sporophytes at early stages of development. In the case of Cinclidium stygium in the Pyrenees, it is clearly linked to the total absence of gametangia. The harsh climatic conditions could be responsible for a depleted fertility, which is common at the margin of plants distribution areas. Cinclidium stygium is a fen indicator included in many types of “eutrophic” communities (see review in Koponen, 1969), mostly Caricion lasiocarpae (Braun, 1968; Paul & Lutz, 1941). There is a great deal of confusion about fen nomenclature, deriving from the phytosociological approach to this vegetation type. Calcareous fens are widely considered eutrophic systems even though detailed chemical characteristic of water or soil are mostly lacking. Though admittedly rich in Ca++ solutes, or other minerals, they are mostly poor in biogenous elements (mostly P and N compounds), and then clearly oligotrophic (Braun, 1968; Lamentowicz et al. 2013; Kooijman et al., 2015). This is likely the case in the Pyrenees where no natural source of nutrient enrichment is known. Cinclidium stygium may extend into thinly wooded stands, which is the case in Alps or Jura, where it is occasionally found in wooded carr. It is generally observed in habitats where water trickles, as in the Pyrenees, or by springs or along rivulets (Mogensen, 1973; Sauer, 1990; Blockeel et al., 2014). In Europe, the habitat requirements appear to be much narrower in the southern regions than in the northern ones. It is frequently associated with Calliergon giganteum (Schimp.) Kindb., Meesia triquetra (L. ex Jolycl.) Ångstr., Scorpidium cossonii, S. scorpioides (Hedw.) Limpr., Hamatocaulis vernicosus (Mitt.) Hedenäs, Drepanocladus trifarius (F.Weber & D.Mohr) Broth. ex Paris (Paul & Lutz, 1941; Braun, 1970). By contrast, the community in the Pyrenees appears rather speciespoor. Calliergon richardsonii is perhaps a rarer associate which is mentioned from small spring mounds at sites of calcium-rich ground water outflow (Antipin et al., 2003). Cinclidium stygium is more frequent and achieves dominance in intermediate floating fens (along with Hamatocaulis vernicosus, Meesia longiseta Hedw., Sphagnum teres) than in aquatic hollows (Pseudocalliergon trifarium, Scorpidium scorpioides) (Braun, 1968). In the Pyrenees no topographic heterogeneity was detectable. Conservation Fen bryophytes are very sensitive to overgrazing, which results in trampling and eutrophication (Stammel et al., 2003; McBride et al., 2011). Grazing by cattle may well not be 8 HUGONNOT: CINCLIDIUM STYGIUM SW. (MNIACEAE, BRYOPSIDA) IN EASTERN PYRENEES appropriate in the case of relictual fens, and the dynamics of such stands are far from well understood. They may be stable on the long run and may not need anthropogenic action. Monitoring of the populations should be undertaken to unravel the vegetation trajectories. Climate warming is certainly of concern because it could enhance development of tall vegetation and overturn water balance on the long run. Additionally, taking advantage of the wide vegetative regeneration of mosses, the capacity of Cinclidium stygium for regrowth should be tested in adequate fen stands in the field. Target stands could be habitats where competition has been lowered, deteriorated, but where water feeding remains intact both from the point of views of quality and amounts. Clonal growth of dispersed gametophores fragments is to be expected and could represent an innovating device of conserving asexually reproducing relict bryophytes in isolated situation, far from their main range. REFERENCES ANTIPIN, V., T. BRAZOVSKAYA., M. BOYCHUK, N. STOIKINA & E. TALBONEN (2003). Flora and vegetation of the mire ecosystems of National park Vodlozersky. Finnish Envir. 485: 137-140. BLOCKEEL, T.L., S.D.S. BOSANQUET, M.O. HILL & C.D. PRESTON (2014). Atlas of British & Irish bryophytes. The distribution and habitat of mosses and liverworts in Britain and Ireland. Volume 1 & 2. British Bryological Society, Pisces Publications, Newbury, 555 pp & 652 pp. BRAUN, W. (1968). Die Kalkflachmoore und ihre wichtigsten Kontaktgesellschaften im bayerischen Alpenvorland. Diss. Bot. 1: 1-134. BRAUN, W. (1970). Bestimmungsübersicht für die Kalkflachmoore und deren wichtigsten Kontaktgesellschaften im Bayerischen Alpenvorland. Ber. Bayer. Bot. Ges. 42: 109-138. CZERNYADJEVA, I.V. & E.A. IGNATOVA (2008). Mosses of the Bystrinsky Nature Park (Kamchatka peninsula, Russian Far East). Arctoa 17: 49-62. HÉTIER, F. (1896). Note sur quelques plantes rares ou nouvelles de la flore Française récoltées dans le Jura. Bull. Soc. Bot. France 43: 66-70. HILLIER,, L. (1954). Catalogue des mousses du Jura. Ann. Sci. Univ. Besançon 2ème sér. 3: 1-221. HODGETTS, N.G (2015). Checklist and country status of European bryophytes - towards a new Red List for Europe. Irish Wildlife Manuals, 84. National Parks and Wildlife Service, Department of Arts, Heritage and the Gaeltacht, Ireland, 125 pp. IGNATOV, M. S. (1994). Bryophytes of Altai Mountains. I. Study area and history of its bryological exploration. Arctoa 3: 13-27. KARCZMARZ, K. (1962). Distribution of Cinclidium stygium Sw. in Poland. Ann. Univ. Mariae Curie Sklodowska Lublin, Polinia sect. C 17: 427-431. KOOIJMAN,, A., C. CUSELL, I.S. METTROP & L.P.M. LAMERS (2015). Recovery of target bryophytes in floating rich fens after 25 yr of inundation by base-rich surface water with lower nutrient contents. App. Veg. Sci 19: 53-65. KOPONEN,, T. (1969). The moss genus Cinclidium (Mniaceae) in Finland. Ann. Bot. Fenn. 6: 112-118. KOPONEN,, T. (1974). A guide to Mniaceae in Canada. Lindbergia 2: 160-184. LAMENTOWICZ, M., Ł. LAMENTOWICZ & R.J. PAYNE (2013). Towards quantitative reconstruction of peatland nutrient status from fens. The Holocene 12: 1661-1665. MCBRIDE, A., I. DIACK, N. DROY, B. HAMILL, P. JONES, J. SCHUTTEN, A. SKINNER & M. STREET. (2011). The Fen Management Handbook. Scottish Natural Heritage, Perth, 329 pp. 9 Boletín de la Sociedad Española de Briología 46–47 (2016) MEINUNGER,, L. & W. SCHRÖDER (2007). Verbreitungsatlas der Mosse Deutschlands. Band 2, 3. Herausgegeben von O. Dürhammer für die Regensburgische Botanische Gesellschaft, Regensburg, 699, 709 pp. MOGENSEN, G.S. (1973). A revision of the moss genus Cinclidium Sw. (Mniaceae Mitt.) Lindbergia 2: 49-80. PAUL,, H. & J. LUTZ (1941). Zur soziologisch-ökologischen Characterisierung von Zwischenmooren. Ver. Bayer. Bot. Gesellschaft 25: 5-32. ROS, R.M., V. MAZIMPAKA, U. ABOU-SALAMA, M. ALEFFI, T.L. BLOCKEEL, M. BRUGUÉS, M.J. CANO, R.M. CROS, M.G. DIA, G.M. DIRKSE, W. EL SAADAWI, A. ERDAĞ, A. GANEVA, J.M. GONZÁLEZ-MANCEBO, I. HERRNSTADT, K. KHALIL, H. KÜRSCHNER, E. LANFRANCO, A. LOSADA-LIMA, M.S. REFAI, S. RODRÍGUEZ-NUÑEZ, M. SABOVJLEVIĆ, C. SÉRGIO, H. SHABBARA, M. SIM-SIM & L. SÖDERSTRÖM (2007). Hepatics and Anthocerotes of the Mediterranean, an annotated checklist. Cryptogamie, Bryologie 28: 351-437. ROS, R.M., V. MAZIMPAKA, U. ABOU-SALAMA, M. ALEFFI, T.L. BLOCKEEL, M. BRUGUÉS, R.M. CROS, M.G. DIA, G.M. DIRKSE, I. DRAPER, W. EL-SAADAWI, A. ERDAĞ, A. GANEVA, R. GABRIEL, J.M. GONZÁLEZ-MANCEBO, C. GRANGER, I. HERRNSTADT, V. HUGONNOT, K. KHALIL, H. KÜRSCHNER, A. LOSADA-LIMA, L. LUÍS, S. MIFSUD, M. PRIVITERA, M. PUGLISI, M. SABOVLJEVIĆ, C. SÉRGIO, H.M. SHABBARA, M. SIM-SIM, A. SOTIAUX, R. TACCHI, A. VANDERPOORTEN & O. WERNER (2013). Mosses of the Mediterranean, an annotated checklist. Cryptogamie, Bryologie 34: 99-283. SARRASSAT, C. (1939). Muscinées récoltées en Haute-Savoie au cours de la session extraordinaire de 1937. Bull. Soc. Bot. France 86: 58-69. SAUER, M. (1990). Die Mniaceae (Sternmoose) Baden-Württembergs. Teil 2: Cinclidium Sw., Rhizomnium (Broth.) T. Kop., Plagiomnium T. Kop. unf Pseudobryum (Kindb.) T. Kop. (mit einem Nachtrag zum 1. Teil). Jh. Ges. Naturkde. Württ. 145: 183-220. STAMMEL, B., K. KIEHL & J. PFADENHAUER (2003). Alternative management on fens: response of vegetation to grazing and mowing. App. Veg. Sci. 6: 245-254. Recepción del manuscrito: 31-03-2016 Aceptación: 17-06-2016 10