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Calliergonella (Bryopsida) in the Iberian Peninsula.

INTRODUCTION

The genus Calliergonella was described by Loeske (1911) to place the single species C. cuspidata (Hedw.) Loeske and has been mainly treated in modern floras and monographs as a distinct and monotypic genus belonging to the Amblystegiaceae G. Roth (e.g. Nyholm 1965, Kanda 1976, Tuomikosky & Koponen 1979, Crum & Anderson 1981), although some exceptions exist. Thus, Boros (1968), Karczmarz (1971) and Smith (1978) included Calliergonella cuspidata in the genus Calliergon (Sull.) Kindb., following the old treatments by Kindberg (1896), Loeske (1910) and Monkemeyer (1927), while Richards & Wallace (1950) and Scott & Stone (1976) included it in Acrocladium Mitt., in accordance with Lindberg (1879), Brotherus (1923) and Dixon (1927).

Tuomikoski & Koponen (1979) and later Hedenas (1989, 1992) definitively established Calliergonella as a genus distinct from Calliergon and Acrocladium. Their results were based on morphological and anatomical characters that had been neglected before by most taxonomists of pleurocarpous mosses. Tuomikoski & Koponen (1979) suggested that the three genera could even belong to different families, retaining only Calliergon in the Amblystegiaceae, although they did not place Calliergonella and Acrocladium in any other family. Nishimura et al. (1984) and Hedenas (1989) placed Calliergonella in the Hypnaceae Schimp., and the latter also moved Acrocladium to the Plagiotheciaceae (Broth.) M. Fleisch., a result that was recently confirmed by phylogenetic studies based on molecular and morphological data (Pedersen & Hedenas 2002). Hedenas (1992, 1995, 1998) was unable to place Calliergonella with certainty either in the Amblystegiaceae or in the temperate Hypnaceae, since the limits between both families were badly defined at that point. However, recent phylogenetic analyses based on molecular data (Vanderpoorten et al. 2001) and on molecular and morphological data (Vanderpoorten et al. 2002a, 2003; Hedenas et al. 2005) have better circumscribed the Amblystegiaceae, and it is now established that there are at least two families within it, the Amblystegiaceae s.str. and the Calliergonaceae (Kanda) Vanderpoorten et al. (see new classification in Vanderpoorten et al. 2002b). It seems now clear that Calliergonella belongs neither to the re-circumscribed family Amblystegiaceae s.str. nor to the Calliergonaceae, and its closest relatives seem to be among the temperate Hypnaceae. Ignatov & Ignatova (2004) have just resurrected the family Pylaisiaceae Schimp. to place Calliergonella together with Breidleria, Pseudohygrohypnum (Hygrohypnum eugyrium and H. subeugyrium), Callicladium, Stereodon, Ptilium, Homomallium and Pylaisia.

Calliergonella had been traditionally considered a monotypic genus (C. cuspidata) until Hedenas (1992) transferred Hypnum lindbergii Mitt. to it. A few species were earlier described under Calliergonella or transferred to it. Grout (1931) included Pleurozium schreberi (Willd. ex Brid.) Mitt. in the genus Calliergonella, nowadays generally accepted to belong to the Hylocomiaceae (Rohrer 1985a, 1985b; Hadenas 2004). Lawton (1971) described a new species in the genus, Calliergonella conardii E. Lawton, which was later synonymized with Fontinalis hypnoides Hartm. (Tuomikoski & Koponen 1979).

Calliergonella lindbergii (Mitt.) Hadenas was first clearly defined as a species separate from Hypnum pratense Koch ex Spruce (currently Breidleria pratensis (Koch ex Spruce) Loeske) by Lindberg (Angstrom 1861). Hitherto, both these species had been confused in the descriptions, or the first was treated merely as variety <<beta>> (Wilson 1855; Schimper 1860). However, Lindberg made an unfortunate choice of an already used name, Hypnum arcuatum. Therefore, the name was an illegitimate homonym, and Mitten (1864) proposed a new name for this species, Hypnum lindbergii Mitt., honouring Lindberg. Calliergonella lindbergii had traditionally been considered to belong to Hypnum (Nyholm 1965, Ando 1973, Smith 1978) or to Breidleria (Loeske 1910) and had always been placed in the Hypnaceae, until Hadenas (1992) circumscribed the genera Calliergonella and Breileria based on many morphological and anatomical characters never used before for this group.

Nowadays it is usually accepted that Calliergonella includes two species, C. cuspidata and C. lindbergii, especially since the circumscription of the genus suggested by Hadenas (1992) has also been supported by phylogenetic analyses based on molecular (Tsubota et al. 2002) or molecular and morphological data (Vanderpoorten et al. 2002a), which showed both species belonging to a very well supported clade (100% bootstrap support).

History of Calliergonella in the Iberian Peninsula

Calliergonella cuspidata was reported from the Iberian Peninsula since the middle of the nineteenth century (Bory de Saint-Vicent (1820), Spruce (1849), Colmeiro (1889) and Roll (1897) in Spain, and Henriques (1889) and Luisier (1907, 1910) in Portugal). At the beginning of the twentieth century C. cuspidata was already well-known as a widely distributed moss in Spain and Portugal (Casares 1915; Machado 1918, 1931). Calliergonella lindbergii was first cited in the Iberian Peninsula (in the Pyrenees) by Jeanbernat & Renauld (1885) as Hypnum arcuatum Lindb. Only a few records of this species have been reported since then, most of them in the Pyrenees (Casas 1986, Ruiz & Brugues 1997) and only one in the Cantabrian Range (Fuertes & Martinez-Conde 1989).

Recent floras and checklists of the Iberian Peninsula still place Calliergonella lindbergii under Hypnum (Casas 1991, Ruiz & Brugues 1997, Casas et al. 2001) and keep Calliergonella as a monotypic genus containing only C. cuspidata. In view of the old-fashioned taxonomic treatment of the genus Calliergonella in the Iberian Peninsula, and since there is not a complete Iberian bryophyte flora, the present work was considered to be needed in order to: (1) assess the taxonomic status of the Iberian specimens of Calliergonella in the light of modern taxonomic works, (2) determine the current distribution and ecology of the species of Calliergonella in the studied area, (3) provide descriptions and illustrations specific for the Iberian specimens of Calliergonella. This study is part of a larger one containing all the genera traditionally placed under Amblystegiaceae s.l.

MATERIALS AND METHODS

This revision is based mainly on herbarium material from PC and the main Iberian herbaria (BCB, FCO, GDAC, LISU, MA, MACB, MAF, MUB, PAMP, SALA, VAB, VIT). Many specimens were also collected by the authors in the studied area; these are now deposited in MACB. Due to the large number of Spanish herbarium specimens belonging to Calliergonella cuspidata, only a selection of these were studied (see Appendix 1), while all the available Spanish specimens belonging to Calliergonella lindbergii have been revised. About 300 specimens were checked. 110 morphological and anatomical characters have been studied (Appendix 2). The descriptions of the gametophytic characters and sexual branches, as well as the habitat descriptions and illustrations are based on Iberian specimens. Nevertheless, since there were no sporophytes in any of the studied Iberian specimens, the sporophytic characters were described from non-Iberian specimens, kept at S and BM, during stays of one of the authors in these institutions. The type specimens kept at S and BM was checked. Nomenclature follows Crosby et al. (1999) and Hadenas (1992). Names of authors are abbreviated according to Brummit & Powell (1992).

RESULTS

Calliergonella Loeske, Hedwigia 50: 248, 1911.

Calliergon sect. Calliergonella (Loeske) Monk., Laubm. Eur. 744, 1927. Type: Calliergonella cuspidata (Hedw.) Loeske, Hedwigia 50: 248, 1911. -- Hypnum cuspidatum Hedw., Sp. Musc. Frond. 254, 1801 (basionym).

Calliergon subgen. Pseudacrocladium Kindb., Eur. N. Amer. Bryin. 1: 80, 1897. -- Hypnum sect. Pseudacrocladium (Kindb.) Paris, Index Bryol. Suppl. 285, 1900. -- Acrocladium sect. Pseuda crocladium (Kindb.) Broth. Nat. Pflanzenfam. I(3): 1038, 1908. Type: Calliergon cuspidatum (Hedw.) Kindb., Canad. Rec. Sci. 6(2): 72, 1894. -- Hypnum cuspidatum Hedw., Sp. Musc. Frond. 254, 1801 (basionym).

Plants forming loose or dense tufts or mats, green or shiny yellowish, medium-sized to large. Stem erect or prostrate, usually branched in one plane, pinnately or irregularly branched; stem transverse section roundoval, with central strand, large and thin-walled medullar cells and 3-5 layers of yellowish-brown, thick-walled cortical cells enclosed by a well developed hyalodermis; axillary hairs abundant, medium-sized, with 1-5 hyaline or brownish apical cells and 1 quadrate to shortly rectangular and brownish basal cell; pseudoparaphyllia foliose, broad; paraphyllia absent. Rhizoids in bundles, below leaf costa insertion, on ventral surface of stem in prostrate plants, red-brown, scarcely branched, smooth. Stem leaves [+ or -] concave, either straight, oblong-ovate with abruptly acute or obtuse and apiculate or rounded apex, or falcate, ovate to oblong-ovate with gradually and shortly to longly acuminate apex; erect-spreading to imbricate, usually complanate; margin plane, entire or slightly denticulate near apex; costa short and double, occasionally absent; median lamina cells linear-flexuose, becoming shorter and wider towards apex and slightly shorter, wider and porose towards base; marginal cells similar or slightly shorter and wider than adjacent median lamina cells; alar cells numerous, hyaline, inflated, forming a well delimited group, slightly or longly decurrent. Branch leaves narrower and smaller than stem leaves or only slightly smaller. Dioicous. Perigonia lateral on stem, perigonial leaves ovate, abruptly acuminate, margin entire, ecostate. Perichaetia lateral on stem; inner perichaetial leaves straight, plicate, lanceolate, erect, margin entire, apex acuminate, costa absent or sometimes short and single, cells fusiform to linear, thin-walled and smooth; vaginula with paraphyses. Seta long (3,5-5,0 cm), reddish, slightly twisted, smooth. Capsule inclined to horizontal, curved, broadly cylindrical, inflated, brownish, furrowed when dry. Exothecial cells irregular, rounded, hexagonal or quadrate in ventral part, longer and rectangular in dorsal part; stomata at base of capsule, phaneropore. Separating annulus present, consisting of 3-4 rows of cells, cells of marginal rows rectangular to shortly rectangular and cells of central rows quadrate to rounded. Peristome well developed; exostome yellowish, brownish towards base, lower outside ornamentation cross-striolate, upper outside ornamentation papillose, border broad, widened where ornamentation changes from cross-striolate to papillose; endostome hyaline to yellowish, basal membrane high, about 35-40% of endostome height, segments perforated, papillose, cilia 2-3, long, appendiculate at least in upper part of cilia, otherwise nodose. Lid conical, obtuse and apiculate. Calyptra cucullate, naked. Spores 12,5-20,0 [micro]m, brownish, finely papillose.
The genus Calliergonella includes two species, both present in the
Iberian Peninsula.

1.- Leaves straight, oblong-ovate; apex abruptly acute or obtuse
and apiculate or rounded. Apices of stems and branches usually
straight, acute and stiff                             1. C. cuspidata

1.- Leaves falcate, ovate to oblong-ovate; apex gradually and
shortly to longly acuminate. Apices of stems and branches usually
hooked                                               2. C. lindbergii


[FIGURE 1 OMITTED]

[FIGURE 2 OMITTED]

1. Calliergonella cuspidata (Hedw.) Loeske, Hedwigia 50: 248, 1911. (Fig. 1).

Hypnum cuspidatum Hedw., Sp. Musc. Frond. 254. 1801 (basionym). -- Stereodon cuspidatus (Hedw.) Brid., Bryol. Univ. 2: 824, 1827. -- Amblystegium cuspidatum (Hedw.) Mitt., Nat. Hist. Azores 312, 1870. -- Acrocladium cuspidatum (Hedw.) Lindb., Musci Scand. 39, 1879. -- Calliergon cuspidatum (Hedw.) Kindb., Canad. Rec. Sci. 6(2): 72, 1894. Lectotype: <<Hypnum cuspidatum. Lipsiae lectus. Hedwig>>, G, selected by Hadenas (1992).

Plants forming loose or dense tufts, green or shiny greenish-yellow to brownish, 5-10(12) cm. Stem often erect, sometimes prostrate, occasionally julaceous, [+ or -] pinnately branched, branches 1-2 cm; stem transverse section round-oval, with central strand, large and thin-walled medullar cells and 4-5 layers of yellowish-brown, thick-walled cortical cells enclosed by a well developed hyalodermis; axillary hairs abundant (4-7 per leaf axil), with 3-5 hyaline or brownish apical cells; pseudoparaphyllia foliose, broad, [+ or -] quadrate with irregular mar gin. Rhizoids scarce in erect plants, more abundant on ventral surface of stem in prostrate plants. Stem leaves straight, slightly concave, oblong-ovate, 1,2-1,3 mm wide x 1,8-2,2 mm long; erect-spreading to imbricate, usually strongly appressed and inrolled towards apices of stem and branches, forming straight, acute and stiff tips, usually complanate; leaf margin plane, entire, sometimes slightly denticulate near apex; apex abruptly acute or obtuse and apiculate or rounded; costa short and double (up to 25% of length leaf); median lamina cells linear-flexuose, 3,7-5,0(6.2) [micro]m wide x 60-120(130) long, becoming shorter and wider towards apex and slightly shorter, wider and porose towards base; marginal cells similar to adjacent median lamina cells; alar cells rectangular to shortly rectangular, hyaline, strongly inflated, thin-walled, forming a very well delimited group, triangular or [+ or -] quadrate, usually excavate and decurrent. Branch leaves smaller and narrower than stem leaves, lanceolate, 0,4-0,8 mm wide x 1,3-2,1 mm long. Dioicous. Perichaetia lateral on stem; inner perichaetial leaves straight, plicate, lanceolate, erect, margin entire, apex acuminate, costa absent or sometimes short and single, cells fusiform to linear, thin-walled and smooth; vaginula with paraphyses. Sporophyte typical for genus. Sporophytes not known from Iberian specimens and rare in non-Iberian specimens.

Habitat: All kinds of wet and nutrient rich habitats. Rich fens, swampy meadows, moist soil in woods and bushy areas with Erica sp., beside eutrophic pools, brooks, etc., on acid or basic substrates. It has been found occasionally submerged in ponds (Penuelas & Comelles 1984).

General distribution: Eurasia, North and South America, Macaronesia, northern and north-eastern Africa, New Zealand and Australia (Hadenas 2003a, 2003b).

Distribution in the Iberian Peninsula (Fig. 2): Calliergonella cuspidata is most abundant in the northern half of the Iberian Peninsula, where it is only absent in the dry areas of the Northern Sub-plateau, while in the southern half this species is restricted to mountainous areas above 1200 m a.s.l. In the Eurosiberian Region C. cuspidata grows from the low montane to alpine belts (30-2000 m a.s.l.), in the north-western coasts of Portugal (Douro Litoral and Beira Litoral) and Spain (Coruna, Lugo), from the Galician-Portuguese Massif (Orense, Minho, Tras-Os-Montes and Alto Douro) to the Cantabrian Range (Lugo, Asturias, Leon, Zamora, Palencia, Cantabria and Basque Mountains (Guipuzcoa, Vizcaya)) and the Pyrenees (Navarra, Huesca, Andorra, Lerida and Gerona). In the Mediterranean Region, it grows in higher altitudes, in the supra-oromediterranean belts of the high mountains (700-2200 m a.s.l.) in the Central Range (Guadalajara, Madrid, Avila, Segovia, Salamanca and Beira Alta), Mountains of Toledo (Caceres) and the Iberian Range (Burgos, La Rioja, Soria, Guadalajara, Cuenca and Teruel). In contrast, in the southern half of the peninsula, which is much dryer, C. cuspidata is restricted to the Penibetic Ranges (Sierra de los Filabres (Garcia-Zamora et al. 1999) and Sierra Nevada in Granada) and Sub-Betic Ranges (Sierra de Cazorla, Segura y Las Villas).

Selected specimens examined: Appendix 1.

Specimens erroneously identified as Calliergonella cuspidata: Appendix 3.

2. Calliergonella lindbergii (Mitt.) Hadenas, Lindbergia 16: 167, 1992 (<<1990>>). (Fig. 3).

Hypnum lindbergii Mitt., J. Bot. 2: 123, 1864 (basionym).. -Hypnum curvifolium subsp. lindbergii (Mitt.) Kindb., Canad. Rec.

Sci. 6: 74, 1894. - Stereodon lindbergii (Mitt.) Braithw., Brit. Moss Fl. 3: 157, 1902.. - Drepanium lindbergii (Mitt.) G. Roth, Eur. Laubm. 2: 628, 55 f. 5, 1904.. - Breidleria lindbergii (Mitt.) W. Schultze-Motel, Nova Hedwigia 5: 88, 1963. Lectotype: <<270. Hypnum arcuatum Lindb. In terra argillaceae circa Holmian. Leg. S. O. Lindberg>>, S!, selected by Hadenas (1992). Isolectotype: BM!: BM000851575.

Hypnum arcuatum Lindb., Ofvers. Forh. Kongl. Svenska Vetensk.-Akad. 18: 371, 1861, illegitimate, later homonym. Original material: S!, BM!: BM000851575.

Hypnum pratense var. arcuatum Molendo, Ber. Naturhist. Vereins Augsburg 18: 184, 1865. - Breidleria arcuata (Molendo) Loeske, Stud. Morph. Syst. Laubm. 172, 1910. Type not seen (cf. Ignatov & Afonina, 1992). Full synonymy in Hadenas (1992).

Plants forming quite dense tufts or carpets, green, shiny yellowish or brownish, 3-8 cm. Stem usually erect, irregularly branched, branches 1,0-2,5 cm; stem transverse section round-oval, with central strand, large and thin-walled medullar cells and 3-4 rows of yellowish-brown thick-walled cortical cells enclosed by a well developed hyalodermis; axillary hairs abundant (4-6 per leaf axil), with 1-4 hyaline or brownish apical cells; pseudoparaphyllia foliose, very broad, transversally rectangular with irregular margin. Rhizoids (abundant on ventral surface of stem in prostrate plants. Stem leaves falcate, hooked at stem apices, concave, ovate to broadly ovate, 0,5-0,8 mm wide x 1,5-2,2 mm long; erect-spreading, usually complanate; leaf margin plane, entire or slightly denticulate near apex; apex gradually and shortly to longly acuminate; costa short and double (up to 35% of length leaf), occasionally absent, seldom single; median lamina cells linear, flexuose, 5-7 [micro]m wide x 50-110(125) [micro]m long, becoming shorter and wider towards apex and shorter, wider and porose towards base; marginal cells slightly shorter and wider than adjacent median lamina cells; alar cells rectangular to shortly rectangular, hyaline, inflated, thin-walled, forming a very well delimited group, triangular, sometimes excavate and not or slightly decurrent. Branch leaves slightly smaller than stem ones. Dioicous. Perichaetia lateral on stem; inner perichaetial leaves straight, plicate, lanceolate, erect, margin entire, apex acuminate, costa absent or sometimes short and single, cells fusiform to linear, thin-walled and smooth; vaginula with paraphyses. Sporophyte typical for genus. Sporophytes not known from Iberian specimens and rare in non-Iberian specimens.

Habitat: Wet and nutrient-rich habitats. Wet or moist meadows, pool or brook shores, moist and loamy soils in woods and beside paths, on basic or acid substrates.

[FIGURE 3 OMITTED]

General distribution: Eurasia, North America, and a few localities in Brazil (Hadenas, 2003a, 2003b).

Distribution in the Iberian Peninsula (Fig. 4): Calliergonella lindbergii is restricted to the Eurosiberian Region, along the Pyrenees (Navarra, Huesca, Lerida and Gerona) and Cantabrian Range (Asturias and Cantabria). In the Pyrenees it grows in the high-montane to alpine regions (1200-2200 m a.s.l.), while in the Cantabrian Range it has been collected at lower altitudes, from 50 to 100 m a.s.l.

Specimens examined: Appendix 1.

DISCUSSION

Calliergonella cuspidata and C. lindbergii are very similar in almost all the characters except in the shape and symmetry of the stem leaves and sometimes in leaf orientation. It seems now surprising that they had never been thought to be closely related until Hadenas (1992) placed them together in the same genus. However, considering that for a long time the classifications of bryophytes were based on a few key characters, such as those mentioned above, maybe it is not so strange that the similarity in many other characters of Calliergonella cuspidata and C. lindbergii were disregarded for a long time. This stresses the importance of using the highest possible number of characters, either morphological or molecular, in order to get accurate taxonomies based on well-resolved phylogenies.

[FIGURE 4 OMITTED]

Calliergonella cuspidata is easily distinguished from other related or similar taxa by the stem with hyalodermis, oblong-ovate leaves, usually obtuse-rounded and apiculate apex, costa short and double, and alar cells inflated and hyaline, forming a well delimited group. These diagnostic characters are constant in all the plants belonging to this taxon. Nevertheless, some other features can vary considerably, which has resulted in the description of a high number of varieties (Karczmarz 1971). It seems that the variability depends on habitat conditions, and many aquatic and terrestrial forms and varieties have been distinguished. Iberian specimens that had been referred to two varieties of Calliergonella cuspidata were found in the collections that were checked for the present work, C. cuspidata var. pungens (Schimp.) Latzel and C. cuspidata var. fluitans (H. Klinggr.) Riehm. All of them were re-identified as Calliergonella cuspidata, since we consider that they fit perfectly within the variability of the species, and their differences are only due to habitat conditions. Karczmarz (1971) reported Calliergonella cuspidata var. caespitosum (H. Whitehouse) Jansen & Wacht. for Spain, but we have seen no Spanish specimens under this name in the revised herbaria.

During this study we found that plants of Calliergonella cuspidata growing in dry and exposed habitats, forming carpets and presenting julaceous stems with acute and stiff apices and yellow-brownish colour, had sometimes been confused with Pleurozium schreberi (Willd. ex Brid.) Mitt. (see Appendix 3). Their appearance in the field may be very similar, but they can be easily differentiated when observed with a microscope. P. schreberi has very different alar cells, rectangular and thick-walled, never hyaline and inflated, and a stem without hyalodermis. C. cuspidata has also occasionally been confused with Scorpidium revolvens (Sw. ex Anonymo) Rubers (Fuertes et al. 2005), probably due to the presence of a hyalodermis on the stem in both species. However, they can easily be separated because S. revolvens has falcate stem leaves and a single costa reaching midleaf or beyond.

Calliergonella lindbergii is easily distinguished from other related or similar taxa by the stem with hyalodermis, ovate to oblong-ovate leaves, gradually and shortly to longly acuminate apex, costa short and double and alar cells inflated and hyaline, forming a well delimited group.

Habitat and distribution in the Iberian Peninsula

In the Iberian Peninsula Calliergonella cuspidata is basically present in all the wetlands with not too poor soils. Since wetlands are more abundant in the northern half of the Iberian Peninsula, this is where C. cuspidata is most abundant. It seems that C. cuspidata's distribution is only determined by the availability of water, and neither any other climatic factor, nor altitude, nor the kind of substrate, seem to affect it. The presence of Calliergonella cuspidata in Sintra (Casares 1915), Mafra (Machado 1918), Algarve (Solms-Laubach 1868) and Valle de la Miel (Allorge & Allorge 1946) could not been confirmed since no material from those localities was found in the revised herbaria (see Fig. 3).

Although the first record for Calliergonella lindbergii in Spain was given by Jeanbernat & Renauld (1885), Spruce had gathered it in the Pyrenees before, although it was distributed by Schleicher as Hypnum circinatum (Mitten 1864). Spruce (1849) reported Hypnum pratense in Bagneres de Bigorre, in the French Pyrenees. Also Wilson (1855) affirmed that Hypnum pratense <<var. a>> was common in the Pyrenees. However, Breidleria pratensis has not been reported from the Spanish Pyrenees, while Calliergonella lindbergii is relatively common in that area (Ruiz & Brugues 1997, Casas 1986). Taking into account that both species were not well defined until Lindberg (in Angstrom 1861) described Hypnum arcuatum, it is possible that the preceding references to Hypnum pratense correspond either to Calliergonella lindbergii or to Breidleria pratensis.

Ruiz & Brugues (1997), in a revision of the genus Hypnum for peninsular Spain, affirmed that Calliergonella lindbergii (Hypnum lindbergii in their treatment) is restricted to the Pyrenees. However, they ignored that a record of this taxon for the Cantabrian range had already been published (Fuertes & Martinez-Conde 1989). We confirm the distribution of this species along the Pyrenees and Cantabrian Range. Calliergonella lindbergii grows in the Pyrenees from 1200 to 2200 m a.s.l., whereas in the Cantabrian Range it has been collected at low altitudes (50 m a.s.l.). This difference in altitude preferences could be due to the humidity conditions. Since the south slope of the Pyrenees is quite continental, mosses with high requirements of humidity have to grow at high altitudes, where the dryness of the summer cannot affect them. The Cantabrian Range has a more Atlantic influence, which provides higher humidity conditions, allowing C. lindbergii and many other mosses to grow at lower altitudes than they do in the Pyrenees.

Although the habitat preferences of Calliergonella cuspidata and C. lindbergii seem to be very similar, the latter is restricted to the Eurosiberian region, while C. cuspidata has a wider distribution. It is likely that C. lindbergii is more dependent on the environmental humidity than C. cuspidata, since it usually grows in more exposed habitats, and consequently it is more sensitive to dry periods. However, it is also possible that the distribution of C. lindbergii is affected by other climatic factors that restrict its growth to the Eurosiberian Region.

Besides these considerations, Calliergonella cuspidata is a since long very well-known taxon and easy to identify in the field, while C. lindbergii has been known only later, in part because it was described later as a separate species from Hypnum pratense. In addition, it is not so easy to recognize this species in the field, and it could have been confused with Hypnum species. Consequently, the distribution area of C. lindbergii in the Iberian Peninsula could be larger than known at present.

Conservation

Calliergonella cuspidata can be considered as a common moss of wet and moist habitats in Andorra, Portugal and Spain. Machado (1931) already made similar observation about C. cuspidata in Portugal: <<Uno de los musgos mas vulgares y caracteristicos de terrenos encharcados>>.

In contrast, Calliergonella lindbergii is considered as a vulnerable species in Spain and the Iberian Peninsula (Sergio et al. 1994), but not throughout Europe (Schumacker & Martiny 1995). According to Hallingback et al. (1998) a species is Vulnerable (VU) when it is not Critically Endangered or Endangered but is facing a high risk of extinction in the wild in the medium-term future, as defined by any of the criteria (A to E) shown in that work. There is only information to assess the threat category of C. lindbergii in the Iberian Peninsula under criterion B (species recently recorded in twenty or fewer 10 km x 10 km squares and found in ten or fewer localities/severely fragmented and in decline). So far C. lindbergii has been reported in sixteen 10 km x 10 km squares and in 19 localities, agreeing only with the first part of criterion B. On the other hand, it occurs in eutrophic habitats, such as roadsides, a circumstance that seems to indicate that C. lindbergii bears quite well the anthropogenic pressure, which is probably the main threat for the wet habitats in the Iberian Peninsula. In our opinion this species, although rare in the Iberian Peninsula, is not menaced.

Appendix 1. Specimens examined.

Calliergonella cuspidata

ANDORRA. 31TCH71, Pal. Pla de Borras, Casas, 13-07-78, BCB 10671; 31TCH91, Vall dlncles, Pont de l'Orry, E. Fuertes & G. Olivan, 17-10-2001, MACB.

PORTUGAL. Beira Alta: 29TPE16, Serra da Estrela, Fuertes, Acon & Olivan, 2000. Beira Litoral: 29TNE45, Penedo da Meditacao. Coimbra, P. Allorge, 1928, PC. Douro Litoral: 29TNF26, BoaNova, V. Allorge, 1950, PC. Minho: 29TNG62, Caldas de Gerez, P. Allorge, 1931, PC. Tras-os-Montes e Alto Douro: 29TPG82, Serra Nogueira sobre Rabordaos, P. Allorge, 1931, PC.

SPAIN. Albacete: 30SWH46, Campamento de San Juan, Riopar, Ros, 1984, MACB 62114. Asturias: 29TPJ52, Ribadeo, P. Allorge, 1956, PC; 30TUP30, Entre Cangas de Onis y Covadonga, P. Allorge, 1933, PC; 30TTP80, Siero, El Plano, H. S. Nava, 1986, MA-Musci 15988; 29TQH27, Parque Natural de Somiedo, E. Fuertes & A. R. Burgaz, 1994, MACB s/n; 29TQJ3219, Pravia, Canedo, J. Munoz, 1988, MA-Musci 15986; 30TUN7499, Penamellera Baja; Narganes, J. Munoz, 1987, MA-Musci 15990; 30TTN6067, Lena, Ubina, vega del Meicin, C. Aedo, 1990, MA-Musci 17327; 30TUN39, Covadonga, camino a las minas, Simo, 1970, MACB 62316. Avila: 30TUK07, Sierra de Gredos, Puerto de la Pena Negra, Brugues, Cros & Lloret, 1993, BCB 41895; Pena Negra, Herguijuela, Costa, MACB 62111; 30TUK291656, Puerto del Pico, Cuevas del Valle, Soria & Ron, 1984, MACB 13841; 30TUK0568, Sierra de Gredos, Laguna del Cervunal, Granzow, Ortiz & Ron, 1983, MACB 11261; 30TUK26, Sierra de Gredos, Puerto del Pico, prope Cuevas del valle, E. Fuertes, 1985, MACB. Barcelona: 31TDF07, Garraf Olivella, Can Suniol, F. Lloret, 1993, BCB 50376. Burgos: 30TWM05, Neila. Refugio de Villavelayo, Casas, 1988, BCB 25353; 30TVM94, Quintanar de la Sierra, Casas, 1988, BCB 33419; 30TVN43, Huidrobo, Fuertes, 1988, MACB 29915. Caceres: 30STJ9784, Navalvillar de Ibor. Garganta Salobriga, Casas, 1995, BCB 49309; 30STJ9875, Rio Ibor, M. C. Viera, 1980, MA-Musci 5093. Cantabria: 30TUN87, Pena Sagra, Olivan & Fuertes, 1997, MACB 75705; Ibidem, Fuertes, 1985, MACB 20652; 30TUN57, Fuente De, MA-Musci 6978, Fuertes, 1981, MACB 28692; 30TUN6166, Vega de Liebana; Puertos de Rio Frio, J. Munoz, 1988, MA-Musci 15989; Ibidem, Fuertes & Acon, 1996, MACB 64277; 30TVN06, Hermandad de Campoo de Suso, Serna, J. Munoz, 1995, MA-Musci 19744; 30TUN9364, Hermandad de Campoo de Suso, Abiada, J. Munoz, 1987, MA-Musci 15987; 30TVP3307, Camargo; Maliano, <<El Carmen>>, J. Munoz, 1988, MA-Musci 15991; 30TUN7474, Cabezon de Liebana, La Tenada de Montecillo, pr. Perrozo, C. Aedo, 1991, MA-Musci 17328, 12220; 30TUN7580, Cabezon de Liebana; supra Cahecho, J. Munoz, 1996, MA-Musci 21307; 30TUN5975, Cosgaya, Viera & Ron, 1979, MACB 31390; 30TUN69, Urdon, Fuertes & Martinez-Conde, 1981, MACB 22187. Coruna: 29TNJ71, Caaveiro, Puentedeume, J. Reinoso, 1979, BCB 7417; 29TNJ83, Sierra de Capelada, Fuertes, Olivan & Sallent, 2000, MACB. Cuenca: 30TWK79, Hoz de Beteta, E. Fuertes & A. Silva, 1982, MA-Musci 276; Ibidem, E. Fuertes & M. Alonso, 1979, MA-Musci 7572; Ibidem, Fuertes & Alonso-Silva, 1980, MACB 10599. Gerona: 31TDG49, Vall Llobre, Lloret, 1985, BCB 25286; 31TDG6166, Carretera de Olot a Sta Pau, Km 6, Sole, 1984, BCB 32192. Granada: 30SVG61, Barranco del Genil, Varo, 1974, MACB 62315. Guadalajara: 30TVL9969, Sierra de Bulejo, Alvarez & Ayala, 1985, MACB 20046; 30TVL74, Prados de Campillejo, Ladero & Demetrio, 1970, MACB 62314; 30TVL95, Sierra del Alto Rey, arroyo Pelagallinas, Gomez, Pajaron & Ron, 1976, BCB 24530; 30TVL86, Hayedo de Cantalojas, rio Lillas, Riestra, 1985, MACB 14541. Guipuzcoa: 30TWN89, Pasajes, P. & V. Allorge, 1932, PC; 30TWN69, Entre Zarauz y Guetaria, P. Allorge, 1933, PC; 30TWP90, Irun, P. Allorge, 1927, PC; Monte Jaizquibel, cerca de Irun hacia el oceano, P. Allorge, 1930, PC; 30TWN69, Zarauz, P. & V. Allorge, 1933, PC; 30TWN88, Cuevas de Landarbaso, V. Allorge, 1968, PC; 30TWN35, Puerto de Arlaban, Gandoger, 1910, PC. Huesca: 31TBH91, Hospital de Benasque, subiendo al Pla dels Estanys, Casas, 1962, MACB 2707; 31TCH02, Benasque, Plan d'Estan, M. Infante & P. Heras, 2002, VIT 29322; 30TYN23, Panticosa, El Bozuelo, Casas, 1965, MACB 3219. La Rioja: 30TWM27, El Rasillo de Cameros, Zubia, MA-Musci 3676; Orillas del Ebro, Zubia, MA-Musci 3648; Orillas del Ebro, Zubia, MA-Musci 6090-3; 30TWM18, Sierra de la Demanda. Penas en la carretera forestal a San Millan de la Cogolla, Casas, 1979, BCB 1002. Leon: 30TUN17, Puerto de las Senales, Fuertes, Acon & Olivan, 17-05-2002, MACB; Laguna de Isoba, Fuertes, Acon & Olivan, MACB; 30TUN07, Puerto de San Isidro, Fuertes, Acon & Olivan, 2002, MACB; 30TUN4580, Posada de Valdeon; pr. Cordinanes, J. J. Aldasoro, 1994, MA-Musci 19254; 30TTN76, Puerto de Pajares, P. Allorge, 1928, PC; 29TQH3366, Cabrillanes; Laguna de las Verdes de Babia, J. J. Aldasoro, 1994, MA-Musci 19249; 29TPH84, Valle del Cuina de Leon, Casas & Cros, 1984, BCB 25263; 29TQH21, Manzanal del Puerto. Branuelas, P. Allorge, 1927, PC; 30TUN4580, Posada de Valdeon, pr. Cordinanes, J. J. Aldasoro, 1995, MA-Musci 13336; 30TUN26, Picos de Mampodre, supra Acevedo, Gandoger, 1905, PC; 29TQH25, subida al Puerto de La Magdalena, E. Fuertes, M. Acon, E. Munin & G. Olivan, 1999, MACB; 30TUN4173, Posada de Valdeon, monte Gildar, Horcada del Oro, pr. Caldevilla, J. Munoz, 1994, MA-Musci 15993; 29TPH81, Toral de los Vados, P. Allorge, 1927, PC. Lerida: 30TCH23, Vall d'Aran, camino a Artiga, Casas, 1966, MACB 2187. Lugo: 29TPH38, Carretera de Oviedo a Lugo, cerca de Meira, Sierra de Meira, V. Allorge, PC; 29TPJ52, Ribadeo, P. Allorge, 1933, PC. Madrid: 30TVL36426, Puerto de Canencia, Vicente & Ron, 1984, MACB 27797; 30TVL55, Hayedo de Monte jo, Fuertes, 1980, MACB 7963; 30TVL32, Sierra de Guadarrama, Puerto de Canencia, Fuertes et al., 2000, MACB 75172; Puerto de Canencia, camino de la Tejera, J. Vicente & E. Ron, 1984, MACB 27796 pp.; 30TVL33, Sierra de Guadarrama. Puerto de Navafria, fuente del Reajo Bajo, Fuertes & Acon, 20-06-2001, MACB; 30TVL55, Dehesa Boyal de Somosierra, Fuertes & Olivan, 2001, MACB; Dehesa Boyal de Robregordo, Fuertes & Olivan, 2001, MACB; 30TVL55, Montejo de la Sierra, E. Fuertes, 1980, MAMusci 153. Navarra: 30TXN52, Sierra de Leyre, barranco de Fuente Fria, Simo, 28-09-1972, BCB 37228; 30TXN74, Valle del Roncal. Uztarroz, Casas, 1960, BCB 15095; 30TXN16, Camino de Sayoa, junto al convento, Fuertes, 1974, MACB 62112; 30TXN09, Endarlaza, P. Allorge, 31-05-1927, PC; 30TXN65, Larrau, P. Allorge, 1938, PC; 30TXN07, Santesteban, P. Allorge, 1930, PC; 30TXN7055, Isaba, pr. Isaba, Mintxate, A. Ederra, 1996, MA-Musci 21318; Isaba, Belabarce, A. Ederra, 1996, MA-Musci 21313. Orense: 29TPG37, Sierra de Queija, Gandoger, 1898, PC; 29TPG57, Viana del Bollo, margen del rio Bibey, P. Allorge, 1933, PC; 29TPG24, Verin, P. Allorge, 1933, PC; 29TNH80, Pinor P. Allorge, 1933, PC; 29TNG98, Laguna de Antela, cerca de Ginzo de Limia, P. Allorge, 1928, PC; 29TPG5383, O Bolo, C. Aedo, 1991, MA-Musci 17218. Palencia: 30TUN66, Curavacas, Fuertes & Acon, 2003, MACB. Salamanca: 30TTK67, Carretera cerca de Bejar, V. Allorge, 1956, PC; 29TQE17, Puerto Descargamaria, Casas, Cros & Brugues, 1985, BCB 21404. Segovia: 30TVL28, San Miguel de Bernuy, prob. E. Guinea, 1940, MA-Musci 6868; 30TVL66, Riofrio de Riaza, Macizo de Ayllon, Pto. de la Quesera, Fuertes & Bermejo, 1982, MACB 56621; Ibidem, Fuertes & Olivan, 2001, MACB; 30TVL06, Zarzuela del Pinar. Borde del rio Cega, P. Allorge, 1931, PC. Soria: 30TWM25, Sierra de Cebollera, E. Fuertes & E. Munin, 1999, MACB. Tarragona: 31TCF37, Montes de Prades, F. Masclaus, 1951, BCB 15097. Teruel: 30TYK08, Barranco del Tajal, Fuertes, 1976, MACB 23136, MA-Musci 7573; 30TXK28, Pinar del Puerto Bronchales, Casas, 1974, BCB 981; 30TXK18, Puerto de Orihuela, Casas, 1974, BCB 980; 30TXK19, Orihuela del Tremedal, Turbera Los Ojos, C. Casas, 1974, MA-Musci 19384 pp. Vizcaya: 30TWP40, Lequeitio, P. Allorge, 1933, PC; 30TWP10, Bakio, E. Guinea, 1941, MA-Musci 14098; Monte Sollube, cerca de Bermeo, P. Allorge, 1932, PC; Bakio, San Miguel, E. Guinea, 1941, MA Musci 14096; Monte Jata, E. Guinea, 1941, MA-Musci 14077; Bakio, Barranco del Infierno, E. Guinea, 1941, MA-Musci 13999, MA-Musci 14010; 30TWN08, Penascal, prob. E. Guinea, 1930, MA-Musci 6888; Penascal, prob. E. Guinea, 1930, MA-Musci 6887; 30TWN49, Montes de urberuaga, Zubia, MA-Musci 3677-2. Zamora: 29TQF19881, Arribes del Duero, Fernandez-Mendoza, 2000, MACB 80609.

Calliergonella lindbergii

ANDORRA. 31TCH71, Grau Roig. Pleta de Morato, Casas, 1975, BCB.

SPAIN. Asturias: 30TUP1914, Colunga, Santiago de Gobiendas, Obaga, J. Munoz, 1986, MA-Musci 17527. Cantabria: 30TUN68, Sierra de Beges, E. Fuertes & Martinez-Conde, 1983, MACB 22159. Gerona: 31TDG18, La Molina, Casas, 1950, BCB 16651; Ibidem, C. Casas, 1948, BCB; La Molina, Torrent de la Molina, Casas, 1956, BCB 25833; 31TDG48, Torrent del Roda, Faitus Llanars, F. Lloret, 1984, BCB 23609; 31TDG49, Riera de Catllar, Vilallonga del Ter, F. Lloret, 1983, BCB 23610; Setcases, Casas, 1955, BCB 16649; 31TDG29, Nuria, camino a la Font Alba, Casas, 1968, MACB 61835; Ibidem, Casas, 1949, BCB 16648; Nuria, hacia el Coll de Finestrelles y cerca del Santuario, Casas, 1949, BCB 14899; 31TDG4384, Lels Adous d'Abella, LLamars, F. Lloret, 1984, BCB 25837; 31TDG57, Vall de Bianya, Casas, 1952, BCB 14900. Huesca: 31TBH51, Valle de Anisclo, Casas, 1955, BCB 47976. Lerida: 31TCH12, Hospital de Vielha, Casas, 1961, BCB 43662; 31TCH21, Boi, Estany Llebreta, Casas, 1991, BCB 43649; 31TCH90, Maranges, Casas, 1958, BCB; 31TCH31, Boi. Sobre el Estany de Colomers, Casas, 1994, BCB 39867; 31TCG99, Montmalus, Viella, P. Monserrat, 1950, BCB 16650. Navarra: 30TXN2482, Arizcun, Ederra, 1985, MACB 61835.

Appendix 2. Morphological and anatomical characters studied.

GAMETOPHYTE

Plant

1. Habit

3. Colour

4. Size

Stem

4. Orientation in relation to the substrate

9. Branching pattern

10. Size of branches

Anatomy

11. Shape of the transverse section

12. Central strand

13. Basic tissue

14. Cortex

15. Hyalodermis

Axillary hairs

16. Number hairs/leaf axil

17. Number of apical cells

18. Colour of apical cells

19. Description of basal cells

Pseudoparaphyllia

20. Present/absent. Shape

Paraphyllia

21. Present/absent. Shape

Rhizoids

22. Position

23. Colour

24. Branching

25. Ornamentation

26. Abundance

Stem leaves

27. Symmetry

28. Rugosity/ presence of undulations/ concavity

29. Shape

30. Size

Orientation in relation to the stem

31. Wet condition

32. Dry condition (if different)

Margin

33. Curvature

34. Denticulation

Apex

35. Description

Costa

36. Length, double, single, etc

37. Surface cells

38. Proration/papillosity?

Areolation

Mid-leaf cells

39. Shape

40. Size

41. Cell walls

42. Papillosity

Apical cells

43. Shape

44. Size

45. Cell walls

Basal cells

46. Shape

47. Cell walls

Marginal cells

48. Shape

49. Size

Alar cells

50. Appearance

51. Cell walls

52. Alar group appearance

53. Alar group decurrency

Initial cells of rhizoids

54. Position

55. Appearance

Branch leaves (if different)

56. Symmetry

57. Shape

58. Size

Sexuality

59. Sexual condition

Perigonia

60. Position

61. Leaf shape

62. Leaf margin

63. Leaf costa

Perichaetia

64. Insertion

65. Paraphyses

Inner perichaetial leaves

66. Symmetry

67. Plication

68. Shape

69. Orientation

70. Margin

71. Apex

72. Costa

Areolation

Upper cells

73. Shape

74. Cells walls

75. Papillosity/proration

Lower cells

76. Shape

77. Cell walls

78. Papillosity/proration

Vaginula

79. Description

SPOROPHYTE

Seta

80. Length

81. Colour

82. Twisting

83. Ornamentation

Capsule

84. Orientation

85. Shape

86. Size

87. Colour

88. Ornamentation (wet/dry)

Exothecial cells

89. Shape

90. Size

91. Papillosity/mammillosity

92. Differentiation (apophysis to mouth)

Stomata

93. Number

94. Position

95. Structure

Separating annulus

96. Absent/present. Description

Peristome

Exostome

97. Colour

98. Orientation (wet/dry)

99. Lower outside ornamentation

100. upper outside ornamentation

101. Margin

102. Border

Endostome

103. Colour

104. Basal membrane height (%)

Segments

105. Perforation

106. Ornamentation

Cilia

107. Number

108. Development

109. Appendiculate/nodose

Lid

110. Description

Calyptra

111. Shape

112. Ornamentation

Spores

113. Size

114. Ornamentation

Appendix 3: Specimens erroneously identified as Calliergonella cuspidata.

SPAIN: Avila: UK0477, Puerto de la Pena Negra, Piedrahita, MACB 11260 is Pleurozium schreberi. Leon: 29TPH74, Tejedo de Ancares, MACB 17491, MA-Musci 4165 is Pleurozium schreberi. Madrid: 30TVL54, Montejo de la Sierra, MACB 19953 is Pleurozium schreberi. Navarra: 30TXN64, urzainqui, PC is Pleurozium schreberi. Segovia: 30TVL66, Riofrio de Riaza, Puerto de la Quesera, MACB 53238 is Pleurozium schreberi; Ibidem, MA-Musci 13377 is Pleurozium schreberi.

ACKNOWLEDGEMENTS

This study has been partially funded by the Spanish Ministry of Science and Technology, D.G.I.C.Y.T, PB 98/0792. The stays of G. O. at S in Stockholm and BM in London were respectively funded by HIGH LAT RESOURCE and SYS-RESOURCE under the EC funded IHP programme. We wish to express our gratitude to the directors and curators of all the herbaria listed above for lending the specimens. Special thanks to Dr. Lars Hadenas and Dr. Len Ellis, curators of the bryophyte collections at S and BM, for their assistance during the visits of G. O. to these institutions.

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Gisela Olivan *, Esther Fuertes * and Margarita Acon **

* Departamento de Biologia Vegetal I, Facultad de Biologia, Universidad Complutense de Madrid, E-28040 Madrid, Spain. giselaolivan@hotmail.com.

** Departamento de Biologia (Botanica), Facultad de Ciencias, Universidad Autonoma de Madrid, E-28049 Madrid, Spain

Recibido: 25 mayo 2005. Aceptado: 13 junio 2005
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Author:Olivan, Gisela; Fuertes, Esther; Acon, Margarita
Publication:Botanica Complutensis
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Date:Jan 1, 2006
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