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Straminergon y Calliergon (Calliergonaceae, Bryopsida) en la Peninsula Iberica.

Straminergon and Calliergon (Calliergonaceae, Bryopsida) in the Iberian Peninsula

INTRODUCTION

TAXONOMIC HISTORY OF STRAMINERGON AND CALLIERGON

The genus Calliergon (Sull.) Kindb., including Straminergon stramineum (Dicks. ex Brid.) Hedenas, has been traditionally placed in the family Amblystegiaceae G.Roth (e.g. Crum & Anderson, 1981; Nyholm, 1965; Karczmarz, 1971; Kanda, 1976; Smith, 1978). As many phylogenetic works have demonstrated (e.g. Hedenas, 1995, 1998; Vanderpoorten & al., 2001, 2002a), the family Amblystegiaceae, as traditionally circumscribed, is polyphyletic, although some monophyletic groups have been found within it. Tuomikoski & Koponen (1979) differentiated the natural group Calliergon-Warnstorfia, result that was later supported by Hedenas (1993), Hedenas & Kooijman (1996) and more recently by Vanderpoorten & al. (2001, 2002a) and Hedenas & al. (2005).

Based on previous phylogenetic works, Vanderpoorten & al. (2002b) described a new family, the Calliergonaceae (Kanda) Vanderpoorten et al., containing Calliergon, Hamatocaulis, Loeskypnum, Straminergon and Warnstorfia. The relationships among the species belonging to these genera, as well as with Scorpidium species and Hygrohypnum ochraceum, which appeared to be closely related with the Calliergonaceae family in the preceding analyses, have been further studied in a recent work (Hedenas & al., 2005), showing that there are two big well supported clades within the family Calliergonaceae, one with Hamatocaulis and Scorpidium, and the other one with Calliergon, Loeskypnum, Straminergon and Warnstorfia. The inclusion of Hygrohypnum ochraceum in the Calliergonaceae is questioned, and further analyses including the rest of Hygrohypnum species are currently being undertaken.

The genus Straminergon was described by Hedenas (1993) to place S. stramineum, which up to then had been traditionally placed in Calliergon (Sull.) Kindb. (e.g. Crum & Anderson, 1981; Nyholm, 1965; Karczmarz, 1971; Kanda, 1976; Tuomikoski & Koponen, 1979). The decision to describe a new genus to place S. stramineum was based on the autoapomorphies present in this species and in the lack of synapomorphic morphological characters joining S. stramineum with the rest of species considered to belong to Calliergon. The placement of S. stramineum in a distinct genus has been later supported by phylogenetic studies based on morphological, anatomical and ecological data (Hedenas & Kooijman, 1996; Hedenas, 1998), on molecular data (Vanderpoorten & al., 2001), and on morphological and molecular data (Vanderpoorten & al., 2002a; Hedenas & al., 2005). Nowadays it is mainly accepted in modern floras that Straminergon is a separate and monotypic genus (e.g. Ignatov & Ignatova, 2004; Smith, 2004; Hedenas, 2003a, 2003b).

Calliergon was a very heterogeneous and artificial genus, containing many unrelated species (check Karczmarz (1971) for details about the taxonomic history of the genus), until Tuomikoski & Koponen (1979) re-circumscribed it on the basis of morphological features that had so far escaped the attention of taxonomists. They kept C. cordifolium (Hedw.) Kindb., C. giganteum (Schimp.) Kindb., C. megalophyllum Mikut., C. richardsonii (Mitt.) Kindb. and Straminergon stramineum (as Calliergon stramineum) in the genus, although pointing out the differences between the latter and the rest of the species, and suggesting its possible placement in a subgenus. This circumscription of the genus Calliergon was later supported by HEDEnas (1993), except for S. stramineum, which, as explained above, was left out. Currently it is widely accepted that Calliergon includes four species: C. cordifolium, C. giganteum, C. megalophyllum and C. richardsonii (Mitt.) Kindb. However, recent results based on morphological and molecular evidence (Hedenas & al., 2005) question the monophyly of the genus, since C. megalophyllum appears, in some of the analyses, out of the well supported clade containing the other three Calliergon species, and within a poorly supported clade as sister of the Warnstorfia-StraminergonLoeskypnum group. More phylogenetic analyses seem to be necessary to decide whether C. megalophyllum belongs to Calliergon or not.

HISTORY OF STRAMINERGON AND CALLIERGON IN THE IBERIAN PENINSULA

Straminergon stramineum was first reported in the Iberian Peninsula by Jeanbernat (1864), who collected it in the Spanish Pyrenees. Machado gathered it for the first time in Portugal, in Serra da Estrela (Central Range) (Machado, 1917). Since then S. stramineum has been reported mainly in the Cantabrian Range, Iberic System and Central Range.

The first record for Calliergon cordifolium in the Iberian Peninsula was provided by Allorge (1928), who collected it in the Cantabrian Range. Later there have only been a few more records for this species in the Pyrenees (Casas, 1986), Central Range (Elias-Rivas, 1988) and Iberian Range (Casas & al., 1984; Garcia-Alvaro & al., 2001).

Calliergon giganteum has been reported in the French Pyrenees since the end of the nineteenth century (Boulay, 1884; Jeanbernat & Renauld, 1885). However, it was not cited in the Spanish Pyrenees until one century later (Brugues & al., 1999). Besides this record, it was also reported in the Cantabrian Range (Simo & al., 1978).

The main goals of the present study have been: Firstly, to assess the taxonomic status of the Iberian specimens of Straminergon and Calliergon in the light of modern taxonomic works. Secondly, to determine the current distribution and ecology of Straminergon and Calliergon species in the Iberian Peninsula, and finally to provide descriptions and iconography specific for the Iberian specimens of Straminergon and Calliergon, since there is not a complete Iberian bryophyte flora.

The present work is part of a general revision of the genera traditionally placed under Amblystegiaceae s.l. in the Iberian Peninsula.

MATERIAL 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, Herb. Martinez-Abaigar). Many specimens have also been collected by the authors in the studied area during this study; these are now deposited in MACB.

About 100 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, illustrations and SEM photographs are based on Iberian specimens. The sporophytic characters of Calliergon cordifolium and C. giganteum are also based on Iberian specimens, while those of Straminergon stramineum were described from non-Iberian specimens kept at S and BM during the visits of one of the authors to these institutions, since there were no sporophytes in any of the studied Iberian specimens belonging to this species. The type specimens kept at S and BM were checked.

Nomenclature follows Crosby & al. (1999) and Hedenas (1993). Names of authors are abbreviated according to Brummit & Powell (1992).

RESULTS

KEY AND DESCRIPTIONS OF STRAMINERGON AND CALLIERGON IN THE IBERIAN PENINSULA

Stem leaves ovate, elongate-ovate or lingulate; alar groups ovate or rectangular along basal leaf margin, reaching up to 50% of distance from leaf margin to costa. Axilary hairs scarce and small (1-2 hyaline apical cells) Straminergon

Stem leaves broadly ovate to broadly ovate-cordate; alar groups triangular, reaching the costa or at least 75% of distance from leaf margin to costa. Axilary hairs abundant and large (2-5 hyaline apical cells) Calliergon

Straminergon Hedenas, J. Bryol. 17: 462. 1993.

Type: Straminergon stramineum (Dicks. ex Brid.) Hedenas, J. Bryol. 17: 463. 1993 (Hypnum stramineum Dicks. ex Brid., Muscol. Recent. 2(2): 172. 1801 (basionym). Calliergon sect. Straminea C.E.O. Jensen, Danmarks Mosser 2: 88. 1923 (Calliergon subgen. Straminea (C.E.O. Jensen) Tuom. & T.J. Kop. Ann. Bot. Fenn.16: 223. 1979. Type: Straminergon stramineum (Dicks. ex Brid.) Hedenas, J. Bryol. 17: 463. 1993 ( Hypnum stramineum Dicks. ex Brid., Muscol. Recent. 2(2): 172. 1801 (basionym).

Plants forming loose tufts, pale yellow, greenishyellow, medium-sized. Stem erect, usually unbranched or with very few short branches; stem transverse section round-oval, with central strand, large and thin-walled medullar cells and 1-2 rows of yellowish thick-walled cells, without hyalodermis; axillary hairs scarce, small, with 1-2 hyaline apical cells and 1-2 quadrate and brownish basal cells; pseudoparaphyllia foliose, broad; paraphyllia absent. Rhizoids on back of leaves or scattered on stem, brown, scarcely branched, smooth, frequent. Stem leaves straight, concave, ovate, elongate-ovate or lingulate; [+ or -] erect and imbricate, sometimes spreading; margin plane, slightly recurved at base, entire; apex rounded or obtuse, cucullate; costa long and single (up to 80% of length leaf); median lamina cells linear, slightly incrassate, porose; marginal cells similar or narrower than adjacent median lamina cells; alar cells rectangular, shortly rectangular or quadrate, hyaline, [+ or -] inflated, [+ or -] thick-walled, forming a well delimited ovate or rectangular group along basal leaf margin, decurrent; initial cells of rhizoids abundant near apex or beside nerve in upper part of leaves, rhomboidal and hyaline. Branch leaves similar to stem ones. Dioicous. Perigonia lateral on lower part of stem, perigonial leaves ovate, abruptly and shortly acuminate, margin entire, ecostate. Perichaetia lateral on stem; inner perichaetial leaves straight, not plicate, ovate-lanceolate, erect, margin entire or irregularly denticulate near apex, apex shortly acuminate, sometimes rounded or obtuse, costa single, usually rudimentary, cells fusiform, porose and smooth; vaginula naked. Seta long (4-5 cm), brownish, [+ or -] twisted, smooth. Capsule inclined to horizontal, cylindrical, curved, brownish, smooth. Exothecial cells shortly rectangular, quadrate to rounded, thick-walled; stomata numerous in base of capsule, phaneropore. Separating annulus absent. Peristome well developed; exostome yellowish, lower outside ornamentation dotted (reticulate at high magnification), upper outside ornamentation papillose, border broad; endostome hyaline to yellowish, basal membrane high, 40-45% of endostome height, segments narrowly perforated, papillose, strongly papillose in upper part, cilia 2-3, long, nodose. Lid conical. Calyptra cucullate, naked. Spores 15.0-22.5 [micro]m, finely papillose.

Straminergon includes one species, S. stramineum.

Straminergon stramineum (Dicks. ex Brid.) Hedenas, J. Bryol. 17: 463. 1993.

Hypnum stramineum Dicks. ex Brid., Muscol. Recent. 2(2): 172. 1801 (basionym) (Hypnum sarmentosum var. stramineum (Dicks. ex Brid.) Sommerf. Suppl. Fl. Lapponicae 67. 1826 (Amblystegium stramineum (Dicks. ex Brid.) De Not., Atti Reale Univ. Genova 1: 137. 1869 (Calliergon stramineum (Dicks. ex Brid.) Kindb. Canad. Rec. Sci. 6(2): 72. 1894 (Acrocladium stramineum (Dicks. ex Brid.) P.W. Richards & E.C. Wallace, Trans. Brit. Bryol. Soc. 1(4): xxv. 1950. Lectotype: BM! (in Dickson herbarium): BM000851585.

Plants forming loose tufts (Figure 1), sometimes solitary shoots mixed with other mosses, pale yellow, greenish-yellow, medium-sized (5-10(12) cm). Stem erect, usually unbranched or with very few short branches (0.5-1.5 cm); stem transverse section round-oval, with central strand, large and thin-walled medullar cells and 1-2 rows of yellowish thick-walled cells, without hyalodermis; axillary hairs scarce (1-2 per leaf axil), with 1-2 hyaline apical cells; pseudoparaphyllia foliose, broadly triangular, lingulate or irregular. Rhizoids in tufts on back of leaves, specially at leaf apices, brown, scarcely branched, smooth, frequent. Stem leaves straight, concave, ovate, elongate-ovate or lingulate, 0.5-0.8 mm wide x 1.0-1.8 mm long; attitude upon the stem [+ or -] erect and imbricate, sometimes spreading; margin plane, slightly recurved at base, entire; apex rounded or obtuse, usually cucullate; costa long and single (up to 80% of length leaf); median lamina cells linear, 4.0-6.0 [micro]m wide (45.0-87.5 [micro]m long, slightly incrassate, porose, becoming shorter and wider towards apex and base; marginal cells similar or narrower than adjacent median lamina cells; alar cells rectangular, shortly rectangular or quadrate, hyaline, [+ or -] inflated, [+ or -] thick-walled, forming a well-delimited group, ovate or rectangular, consisting of 3-4 rows of cells along basal leaf margin, upper cells of group less inflated and smaller, usually quadrate or rectangular, alar group decurrent; initial cells of rhizoids abundant near apex or beside nerve in upper part of leaves, rhomboidal and hyaline. Dioicous. Perichaetia as described for the genus. Sporophyte as described for the genus. Sporophytes not known from Iberian specimens.

Habitat: Sphagnum bogs, fens, rarer on irrigated rocks and beside streams. Sometimes submerged.

General distribution: Widespread in artic and temperate areas in the Northern Hemisphere, rarer in Southern Hemisphere (Hedenas, 2003a, 2003b).

Distribution in the iberian Peninsula (Figure 2): Straminergon stramineum is widespread in the mountainous areas of the northern half of the Iberian Peninsula, while in the southern half this species is restricted to Sierra Nevada. In the Eurosiberian Region S. stramineum grows in the Cantabrian Range (Lugo, Asturias, Leon, Zamora, Palencia, Cantabria and Vasque Country) from the low montane to alpine belts (90-2000 m a.s.l.), and in the Pyrenees (Navarra, Huesca, Andorra, Lerida and Gerona), where it can be found at higher altitudes (1300-2450 m a.s.l.). In the Mediterranean Region it grows in the oromediterranean belts of the high mountains (18002225 m a.s.l.) in the Central Range (Madrid, Avila, Segovia, Salamanca and Beira Alta), Iberian Range (Burgos, La Rioja, Soria, zaragoza) and Penibetic Range (Sierra Nevada in Granada).

Specimens examined: Appendix 1.

Calliergon (Sull.) Kindb., Canad. Rec. Sci. 6(2): 72. 1894.

Hypnum sect. Calliergon Sull., Manual (ed. 2) 672. 1856 (basionym) (Amblystegium subgen. Calliergon (Sull.) Lindb. Musci Scand. 34. 1879--Hypnum subgen. Calliergon (Sull.) Lesq. & James, Man. Mosses N. America 318, 402. 1884-- Amblystegium sect. Calliergon (Sull.) Braithw., Brit. Moss Fl. 3: 17. 1896. Type: Calliergon cordifolium (Hedw.) Kindb., Can. Rec. Sci. 6(2): 72. 1894--Hypnum cordifolium Hedw., Sp. Musc. Frond. 254. 1801 (basionym).

Amblystegium sect. Obtusifolia De Not., Atti Reale Univ. Genova 1: 129. 1869. Type: Amblystegium cordifolium (Hedw.) De Not., Cronac. Briol. Ital. 2: 23. 1867--Hypnum cordifolium Hedw., Sp. Musc. Frond. 254. 1801 (basionym).

Calliergon sect. Cordifolia C.E.O. Jensen, Danmarks Mosser 2: 88. 1923. Type: Calliergon cordifolium (Hedw.) Kindb., Canad. Rec. Sci. 6(2): 72. 1894--Hypnum cordifolium Hedw., Sp. Musc. Frond. 254. 1801 (basionym).

Plants forming loose tufts, green, pale green, green, yellowish-green to brownish, never reddish (outside the Iberian Peninsula sometimes pinkish), medium-sized to large and robust (outside the Iberian Peninsula very large). Stem usually erect, radially branched, pinnately or irregularly branched; stem cross section round-oval, with central strand, large and thin-walled medullar cells and cortex of yellowish thick-walled cells, without hyalodermis; axillary hairs abundant, large, with 2-5 hyaline apical cells (outside de Iberian Peninsula to 10) and 1-2 quadrate to shortly rectangular and brownish basal cells; pseudoparaphyllia foliose, broad; paraphyllia absent. Rhizoids on back of leaves or sparse on stem, brown-reddish, scarcely branched, smooth. Stem leaves straight, concave, [+ or -] broadly ovate, ovate-cordate (outside the Iberian Peninsula also rounded-ovate or broadly rounded-ovated); attitude upon the stem erect to spreading; margin plane, entire or slightly sinuate; apex rounded or obtuse; costa single, almost reaching the apex (outside the Iberian Peninsula also branched, up to 80% of length leaf); median lamina cells linear, smooth, becoming wider and shorter towards apex and base; marginal cells narrower; alar cells rectangular, hyaline, inflated, thin-walled, forming a triangular group (outside the Iberian Peninsula also oval-rounded), reaching or not the costa, sometimes excavate, broadly decurrent, transition from alar cells to adjacent basal cells gradual or abrupt; initial cells of rhizoids very common near leaf apex and beside costa, rhomboidal and hyaline. Branch leaves smaller and narrower (or only smaller outside the Iberian Peninsula). Autoicous or dioicous. Perigonia lateral on lower part of stem, below perichaetia in autoicous species, perigonial leaves ovate to ovate-oblong, abruptly and shortly acuminate or apiculate, margin entire, ecostate. Perichaetia lateral on stem; inner perichaetial leaves straight, not plicate, ovate-lanceolate, erect, margin entire or slightly sinuate, apex abruptly and shortly to longly acuminate, costa long and single, almost reaching the apex, cells linear, thin-walled, smooth, sometimes initial cells of rhizoids present in upper part of perichaetial leaves; vaginula naked. Seta long (4-5 cm), reddish, slightly twisted, smooth. Capsule horinzontal, cylindrical, brownish, smooth. Exothecial cells shortly-rectangular, quadrate to rounded, thickwalled, smooth; stomata at base of capsule, phaneropore kind. Separating annulus absent. Peristome well developed; exostome yellowish, lower outside ornamentation dotted (reticulate at high magnification), upper outside ornamentation papillose, border broad; endostome hyaline, basal membrane high, 40-45% of endostome height, segments narrowly perforated, finely papillose, cilia 2-4, long, nodose. Lid conical. Calyptra cucullate, naked. Spores 12.5-23 pm, finely papillose.

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Calliergon includes four species, from which two occur in the Iberian Peninsula, C. cordifolium and C. giganteum.

Key to the species of Calliergon in the Iberian Peninsula

Stems slightly and irregularly branched or unbranched. Alar groups (diffusely to well defined, usually reaching the costa or almost so; transition from alar cells to adjacent basal cells gradual. Autoicous ... C. cordifolium

Stems densely and pinnately branched. Alar groups very well defined, usually not reaching the costa (75-90% of distance from leaf margin to costa); transition from alar cells to adjacent basal cells abrupt. Dioicous ... C. giganteum

Calliergon cordifolium (Hedw.) Kindb., Canadian Record of Science 6(2): 72. 1894.

Hypnum cordifolium Hedw., Sp. Musc. Frond. 254. 1801 (basionym) (Stereodon cordifolius (Hedw.) Brid., Bryol. Univ. 2: 824. 1827 (Amblystegium cordifolium (Hedw.) De Not., Cronac. Briol. Ital. 2: 23. 1867 (Acrocladium cordifolium (Hedw.) P.W. Richards & E.C. Wallace, Transac. Brit. Bryo. Soc. 1(4): xxv. 1950. Lectotype: "In Svecia lectam acceptiunque a D. O. Swartz" (plant no. 3 from the left of the sheet), G, selected by Hedenas (1993), amending previous selection by Karczmarz (1971).

Full synonymy in Hedenas (1993).

Plants in loose tufts (Figure 3, 4(1-2)), green or pale green, medium-sized to large (7-12(15) cm). Stem erect, slightly and irregularly branched or unbranched, branches 2-3 cm; stem cross section round, with central strand, large and thin-walled medullar cells and 2-3 rows of yellowish thick-walled cells, without hyalodermis; axillary hairs abundant (3-5 per leaf axil), with 2-5 hyaline apical cells; pseudoparaphyllia foliose, semi-orbicular or broadly triangular, margin irregular. Rhizoids forming tufts on back of leaves or scattered on stem, brown-reddish, scarcely branched, smooth, quite abundant. Stem leaves straight, concave, broadly ovate or ovate-cordate, 1.5-2.0 mm wide (3.0-4.0 mm long; attitude upon the stem erect-spreading to spreading; margin plane, convolute when dry, entire or slightly sinuate; apex rounded or obtuse; costa long and single, almost reaching the apex, (60)80-100(120) pm wide at base and gradually narrowed along leaf; median lamina cells linear, straight or slightly flexuose, 5.0-8.75 pm wide (75.0-112.5 [micro]m long, thin-walled, smooth, becoming shorter, wider and flexuose towards apex, shorter and wider, longly rhomboidal to rectangular and porose towards base; marginal cells narrower than adjacent median lamina cells; alar cells rectangular, hyaline, inflated, thin-walled, forming a (diffuse to well defined group, triangular, usually reaching the costa, sometimes excavate and broadly decurrent, transition from alar cells to adjacent basal cells gradual; initial cells of rhizoids very common in apex and beside nerve, specially in the upper fifth of leaf. Branch leaves ovate, smaller and narrower than stem leaves, 1.0-1.6 mm wide (2.4-2.7 mm long. Autoicous. Perichaetia lateral on stem above usually numerous perigonia; inner perichaetial leaves straight, not plicate, ovate-lanceolate, erect, margin entire or slightly sinuate, apex abruptly and shortly to longly acuminate, costa long and single, almost reaching the apex, cells linear, thin-walled, smooth, sometimes initial cells of rhizoids present in upper part of perichaetial leaves; vaginula naked. Sporophyte typical for the genus. Spores 12.5-20 [micro]m, finely papillose. Sporophytes very rare in Iberian specimens.

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Habitat: Rich fens, wet high-mountain meadows, beside mountain pools and streams and on soil in wet forests. Rarely submerged.

General distribution: Temperate areas in the Northern Hemisphere (Hedenas, 2003a).

Distribution in the Iberian Peninsula (Figure 5): Calliergon cordifolium grows in the montane to alpine belts (1200-2000 m a.s.l.) in the Pyrenees (Huesca),

Cantabrian Range (Asturias and Leon) and northwestern part of the Iberian Range (La Rioja and Soria).

Specimens examined: Appendix 1.

Specimens erroneously identified as Calliergon cordifolium: Appendix 3.

Calliergon giganteum (Schimp.) Kindb., Canadian Record of Science 6(2): 72. 1894.

Hypnum giganteum Schimp., Syn. Musc. Eur. 642. 1860 (basionym) (Stereodon giganteus (Schimp.) Mitt., J. Proc. Linn. Soc. 8: 43. 1864 (Amblystegium giganteum (Schimp.) De Not., Cronac. Briol. Ital. 2: 23. 1867 (Amblystegium aduncum var. giganteum (Schimp.) De Not., Cronac. Briol. Ital. 2: 24. 1867 (Hypnum cordifolium var. giganteum (Schimp.) Sanio, Verh. Bot. Prov. Brandenburg 24: 84. 1882 (Acrocladium giganteum (Schimp.) P.W. Richards & E.C. Wallace, Trans. Brit. Bryol. Soc. 1(4): xxv. 1950. Syntypes: "Suecia", "pedale et altius Wiszniewo Borussiae occid. a cl. Dr Klinggraff lectum", "Vogeso", "Helvetia" and "Salisburgia amic. Sauter misit" (not seen). Calliergon subsarmentosum Kindb., Rev. Bryo. 36: 42. 1909. Holotype: "Noth America, Canada, Vancouver Island, Shawnigan Lake, wet earth, 18 june 1908 J. Macoun", S!: B8239. Full synonymy in Hedenas (1993).

Plants forming loose tufts (Figure 6, 4(3-4)), shiny yellowish-green or brownish, large and robust (10-20 cm). Stem erect, pinnately and radially branched, branches 0.5-1.0 cm; stem cross section round-oval, with central strand, large and thin-walled medullar cells and 2-3 rows of yellowish thick-walled cells, without hyalodermis; axillary hairs abundant (3-5 per leaf axil), with 2-5 hyaline apical cells; pseudoparaphyllia foliose, broad, margin irregular. Rhizoids on back of leaves or scattered on stem, red-brown, scarcely branched, smooth, infrequent. Stem leaves straight, plane, broadly ovate-cordate, 1.7-3.0 mm wide x 2.25-4.0 mm long; attitude upon the stem erect-spreading to spreading; margin plane, entire or slightly sinuate or finely denticulate at apex; apex rounded or obtuse; costa long and single, almost reaching the apex, 90-190 [micro]m wide at base and narrowing very little, keeping a stout appearance throughout its length; median lamina cells linear, 3.7-4.5 [micro]m wide x 50.0-70.0 [micro]m long, thin-walled, becoming wider beside costa, shorter, wider and more flexuose towards apex, shorter, wider, thick-walled and porose towards base; marginal cells narrower than adjacent median lamina cells; alar cells rectangular, hyaline, inflated, thin-walled, forming a very well defined group, triangular, usually not reaching the costa (up to 70-90% of distance from leaf margin to costa), strongly excavate and broadly decurrent, transition from alar cells to surrounding cells abrupt; initial cells of rhizoids very common in the upper third of leaves and beside costa, rhomboidal and hyaline. Branch leaves ovate-lanceolate, 0.5-1.0 mm wide (1.5-2.0(3.0) mm long; apex acute and blunt, usually incurved, giving a tubular appearance. Dioicous. Perichaetia lateral on stem; inner perichaetial leaves straight, not plicate, ovate-lanceolate, erect, margin entire or slightly sinuate, apex abruptly and shortly to longly acuminate, costa long and single, almost reaching the apex, cells linear, thin-walled, smooth; vaginula naked. Sporophyte typical of the genus. Spores 20-23 [micro]m, finelly papillose. Sporophytes very rare in Iberian specimens.

Habitat: Rich fens and wet meadows beside mountain pools and streams. Sometimes submerged.

General distribution: Temperate areas in the Northern Hemisphere, southernmost South America and New Zealand (Hedenas, 2003a).

Distribution in the Iberian Peninsula (Figure 7): Calliergon giganteum is restricted to the Eurosiberian Region, in the Pyrenees (Lerida) and Cantabrian Range (Asturias). In the Pyrenees it grows only in the alpine region (1800 m a.s.l.), while in the Cantabrian Range it has been collected at lower altitudes, from 90 to 1870 m a.s.l.

Specimens examined: Appendix 1.

Specimens erroneously identified as Calliergon giganteum: Appendix 3.

DISCUSSION

TAXONOMIC REMARKS

Straminergon stramineum is a very distinct species, usually easily recognized, since it is not very variable. However, in the Iberian Peninsula it has occasionally been confused with Calliergon cordifolium, which can be more variable, especially in branching pattern and leaf shape. Therefore, C. cordifolium forms with unbranched stems and narrow leaves were confused with S. stramineum, while forms with branched stems and very broad leaves were sometimes identified as C. giganteum and more rarely as Brachythecium rivulare (see Appendix 3). From the latter species C. cordifolium can be differentiated by the apex acute and apiculate or shortly acuminate, and leaf margin dentate that B. rivulare usually shows. Table 1 summarize the differences among S. stramineum, C. cordifolium and C. giganteum for Iberian specimens.

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HABITAT AND DISTRIBUTION IN THE IBERIAN PENINSULA

Straminergon stramineum grows usually in fens in the mountainous areas of the northern half of the Iberian Peninsula and in Sierra Nevada, mainly from 1200 to 2200 m.a.s.l., except in the Cantabrian Range, where it can be found at lower altitudes, in fens close to the sea, about 100 m.a.s.l.

This distribution pattern is very common in many bryophyte species in the Iberian Peninsula. These occur in the alpine regions, above 1200 m.a.s.l., in the interior mountainous areas of the Peninsula, while grow under 100 m.a.s.l. in the Cantabrian Range thanks to the Atlantic influence, which provides higher humidity conditions, allowing many plants with high humidity requirements to grow at lower altitude than they do in the interior mountain ranges, which have more continental conditions.

Calliergon cordifolium was recorded in two localities in the Central Range (Elias-Rivas, 1988), but the specimens from these localities were erroneously identified (see Appendix 3). Thus, the distribution of this species in the Iberian Peninsula is restricted to the northern third of the peninsula, occurring in rich fens or wet meadows in the alpine regions (12002000 m.a.s.l.) in the Pyrenees, Cantabrian Range and north-western part of the Iberian Range.

Even rarer than Calliergon cordifolium is C. giganteum, which despite having very similar habitat preferences, is only present in one locality in the Catalonian Pyrenees and in other four in the Cantabrian Range. Just like Straminergon stramineum, Calliergon giganteum grows at lower altitudes in the Cantabrian Range than in the Pyrenees.

The three species studied in the present paper grow in similar habitats, mainly fens in communities of the class Scheuchzerio palustris-Caricetea nigrae Tuxen (cf. Rivas-Martinez & al., 2002). However, Straminergon stramineum occurs in poor to rich fens, while Calliergon cordifolium and C. giganteum seem to prefer rich fens, with more vascular plants. According to Hedenas & Kooijman (1996), who studied the habitat adaptations within the Calliergonaceae, based on morphological and habitat data measured in Sweden, Straminergon is adapted to less mineral-rich habitats and drier conditions than Calliergon species. This could explain why Straminergon has a larger distribution area in the Iberian Peninsula, since it grows in a wider range of pH and nutrient contents and can better bear dry periods. on the other hand the fact that Calliergon species need high contents of minerals and constant wet conditions restrict their growth to a few localities in the northern part of the Iberian Peninsula, where they can find such environmental conditions. There are some works about bryophyte vegetation in fens in the Eurosiberian Region of Spain (e.g. Ballesteros & al., 1983; Fernandez-Prieto & al., 1987). However, more habitat data, specifically collected in the sites where the Calliergon species grow, are needed to clarify the reasons of their distribution.

CONSERVATION

Calliergon cordifolium and C. giganteum are considered as vulnerable species in the Iberian Peninsula and Spain (Sergio & al., 1994), but not throughout Europe (Schumacker & Martiny, 1995).

Applying the criteria to evaluate the threat category of mosses by Hallingback & al. (1998), C. cordifolium is a Vulnerable (VU) species in the Iberian Peninsula, as defined by 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). The occurrence of C. cordifolium has been confirmed in the present work for seven 10 km x 10 km squares and seven localities in three different areas (Figure 5).

Even more critical is the situation of C. giganteum in the Iberian Peninsula, since it has been reported from five 10 km x 10 km and five localities, one of them isolated in the southern slope of the Pyrenees. According to Hallingback & al. (1998) C. giganteum could be considered as an endangered (EN) species in the Iberian Peninsula, as defined by criterion B (species recently recorded in five or fewer 10 km x 10 km squares and found in two to five localities/severely fragmented and in decline).

APPENDIX 1. Specimens examined. Straminergon stramineum

ANDORRA. 31TCH72, Tristaina, riu de la Coma del Rat, BCB 10667, C. Casas, 1978; 31TCH90, Cercle de Pessons, BCB 47816, R. M. Cros, 1983; 31TCH91, Vall d'Incles, Pont de l'Orry, MACB, E. Fuertes & G. Olivan, 2001.

SPAIN. Asturias: 29TQH06, Laguna de Arbas, PC, P. Allorge, 1927; Ibidem, MACB, E. Fuertes, M. Acon & G. Olivan, 1999; 29TQH16, Puerto de Leitariegos, MACB, E. Fuertes, M. Acon & G. Olivan, 1999; 29TQJ22, Cudillero, turbera <<Las Duenas>>, FCObriof 1978, C. Fernandez-Ordonez, 1993.; 30TUN60, Pico de Tres Mares. Turberas en la orilla del rio Nansa, PC, R. Heim, 1926; Ibidem, MACB, E. Fuertes, M. Acon & G. Olivan, 2000. Avila: 30TUK16, Sierra de Gredos. De la Plataforma al Puerto de Candeleda, BCB 41898, M. Brugues & al., 1993. Burgos: 30TVM9554, Palacios de la Sierra, Laguna Negra de Neila, VIT 27869, P. Heras & al., 2001. Cantabria: 30TUN87, Pena Sagra, MACB 20673, E. Fuertes, 1985; Ibidem, MACB, E. Fuertes & G. Olivan, 1997. Gerona: 29TDG29, Nuria, Coll de Finestrellas, BCB 15113, C. Casas, 1949. Granada: 30SVG60, Sierra Nevada, cerca de los albergues, PC, R. Maire, 1931. Guipuzcoa: 30TWN45, Monte Garagarza, PAMP 3267, J. Arraiza, 1984. Huesca: 31TCH02, La Maladeta, Lago de Paderna en La Reclusa, MACB 2195, FCO-bryof 356, MAF 9, C. Casas, 1966; Alta Ribagorza. Benasque. Ibon Coronas, BCB 51269, R. M. Cros, 1975; Benasque, VIT 6892, P. Heras, 1985; 30TYN23, Balneario de Panticosa, pequeno lago cerca del balneario, PC, P. Allorge, 1935; 30TYN2741, Panticosa, cerca del Ibon Brachimana, VAB 1978, BCB 15110, MACB 62115, PAMP 5039, MAF 10, C.Casas, 1965. La Rioja: 30TWM35, Puerto de Piqueras, MACB, G. Olivan, 2000. Leon: 29TPH74, Sierra de Ancares, lago Cruina, MACB 17515, E. Fuertes, 1984; 29TQH22, Branuelas, PC, P. Allorge, 1927; 29TQH35, Subida a Villabandin, FCO-briof 1924,1925, S. Rivas-Martinez, J. L. Fernandez Prieto, T. E. Diaz & A. Penas, 1985;. 30TUN07, San Isidro. Turbera junto al lago del Ausente, PC, P. Allorge, 1934; Ibidem, MACB, E. Fuertes, M. Acon & G. Olivan, 2002. Lerida: 31TCH12, Vall d'Aran, Estany Redo, PAMP 5054, MACB 62116, VAB 1962, E. Ballesteros, 1986; 31TCH21, Vall de San Nicolau, BCB 15114, C. Casas, 1959; 31TCH32, Val d'Aran. Puerto de Bonaigua, PC, O. Bolos & O.H. Folk, 1958; Puerto de la Bonaigua, BCB 15111, C. Casas, 1954; 31TCH90, Maranges, lago Malniu, BCB 15104, C. Casas, 1958; Baixa Cerdanya. Clots del Port. Sobre refugi dels Estanys de la Pera, BCB 51089, M. Brugues & al., 1999. Lugo: 29TPH63, Puerto de Piedrafita, PC, P. & V. Allorge, 1927; Piedrafita, PC, P. & V. Allorge, 1927; 29TPJ20, Sierra de Xistral, MACB, J. R. Oubina, 1983; Ibidem, MACB, E. Fuertes, G. Olivan & A. Sallent, 2000. Madrid: 30TVL11, Sierra de Guadarrama. Pequena turbera cerca del puerto de Cotos, PC, P. Allorge, 1953; 30TVL12, Penalara, Sierra de Guadarrama, PC, P. Allorge, 1953; Sierra de Guadarrama, cerca del refugio Zabala, MACB, E. Fuertes & A. Sallent, 2000; 30TVL22, Sierra de Guadarrama, Parque Natural de Penalara, Laguna de Los Claveles, MACB, E. Fuertes, M. Acon & G. Olivan, 2000. Navarra: 30TXN68 Zurian, PAMP 3268, J. Arraiza, 1984. Palencia: 30TUN56, Valle de las Lomas, prope Cardano de Arriba, MACB, I. Arroyo, 1995; 30TUN66, Pozo de Curavacas, BCB 27372, P. Monserrat, 1950; Ibidem, MACB, E. Fuertes & M. Acon, 2003. Salamanca: 30TTK67, Sierra de Bejar, Circo de la Pena Negra, SALA-BRYO 470, BCB 22981, M. J. Elias, 1984; Herguijuela, MACB 62117, GDAC 7569, PAMP 2674, MAF 2288, M. Ladero, 1973. Soria: 30TWM15, Laguna Negra, Sierra de Urbion, BCB 968, M. Brugues, 1974. Vizcaya: 30TVN67, Carranza, Salduero, VIT 27470, P. Heras & M. Infante, 2001. Zamora: 29TPG87, Sierra de Segundera, Laguna de los Peces supra San Martin de Castaneda, MACB 68699, E. Fuertes, 1998; 29TPG87, Galande, Laguna de los Peces, VIT 24676, M. Infante & P. Heras, 1999. Zaragoza: 30TWM92, Moncayo, MAF 8, B. Lazaro-Ibiza.

PORTUGAL. Beira Alta: 29TPE16, Serra da Estrela, Fonte da Canoriza, pr. Lagoa Redonda, LISU 176125, C. Sergio, A. Seneca & J. Jansen, 1992; Serra da Estrella, Lagunillas de la Torre, MACB, E. Fuertes, M. Acon & G. Olivan, 2000.

Calliergon cordifolium

SPAIN. Asturias: 29TQH16, Puerto de Leitariegos, lagunas turbosas sobre la laguna de Arbas, PC, P. Allorge, 1927; 29TQH39, Grao, Cuellagar, MA-Musci 15981, J. Munoz, 1990; 29TQH16, Puerto de Leitariegos, MA-Musci 9712, P. Allorge, 1911; Ibidem, MAMusci 3041, P. Allorge, 1927. Huesca: 31TCH02, Benasque, Plan d'Estan, VIT 29310, M. Infante & P. Heras, 2002; 30TXN94, Arangues del Preesto. Paul de Bernera, BCB 22980, P. Monserrat & D. Gomez, 1985. La Rioja: 30TWM05, Laguna de Urbion, Herbario de Martinez-Abaigar, Garcia Alvaro, 1995. Leon: 30TUN35, Riano, orillas del Esla, BCB 15115, 15108, M. Losa & P. Monserrat, 1952. Soria: 30TWM39, Sierra del Moncayo, vertiente norte, MACB 62129, E. Fuertes, 1982.

Calliergon giganteum

SPAIN. Asturias: 29TQH26, Puerto de Somiedo, VIT 24519, P. Heras & M. Infante, 1999; 29TQH27, Puerto de Somiedo, vega Penonta, FCO-briof 355, J. L Fernandez-Prieto, 1978; Puerto de la Cubilla, FCO-briof 354, C. Fernandez-Ordonez & Vigon, 1978; 29TQJ12, Cadavedo, PC, V. Allorge, 1956; 30TTN66, Lena, Puertos de la Cubilla, MA-Musci 15980, C. Fernandez-Ordonez, 1978. Lerida: 31TCH23, Pla de Beret, Vall d'Aran, BCB 31840, C. Casas, 1975.

APPENDIX 2. Morphological and anatomical characters studied.

GAMETOPHYTE

Plant

1. Habit

2. Colour

3. Size

Stem

4. Orientation to the substrate

5. Branching pattern

6. Size of branches Anatomy

7. Shape of the transversal section

8. Central strand

9. Basic tissue

10. Cortex

11. Hyalodermis Axillary hairs

12. Number hairs/leaf axil

13. Number of apical cells

14. Colour of apical cells

15. Description of basal cells Pseudoparaphyllia

16. Present/absent. Shape Paraphyllia

17. Present/absent. Shape

Rhizoids

18. Position

19. Colour

20. Branching

21. Ornamentation

22. Abundance

Stem leaves

23. Symmetry

24. Rugosity/ undulosity/ concavity

25. Shape

26. Size

Attitude upon the stem

27. Wet condition

28. Dry condition (if different)

Margin

29. Curvature

30. Denticulation

Apex

31. Description

Costa

32. Length, double, single, etc

33. Surface cells

34. Proration/papilosity? Areolation

Mid-leaf cells

35. Shape

36. Size

37. Cell walls

38. Papillosity Apical cells

39. Shape

40. Size

41. Cell walls Basal cells

42. Shape

43. Cell walls Marginal cells

44. Shape

45. Size Alar cells

46. Appearance

47. Cell walls

48. Alar group appearance

49. Alar group decurrency Initial cells of rhizoids

50. Position

51. Appearance

Branch leaves (if different)

52. Symmetry

53. Shape

54. Size

Sexuality

55. Sexual condition

Perigonia

56. Position

57. Leaf shape

58. Leaf margin

59. Leaf costa

Perichaetia

60. Insertion

61. Paraphyses Inner perichaetial leaves

62. Simmetry

63. Plication

64. Shape

65. Attitude

66. Margin

67. Apex

68. Costa Areolation Upper cells

69. Shape

70. Cells walls

71. Papilosity/proration Lower cells

72. Shape

73. Cell walls

74. Papilosity/proration

Vaginula

75. Description

SPOROPHYTE

Seta

76. Length

77. Colour

78. Twisting

79. Ornamentation

Capsule

80. Orientation

81. Shape

82. Size

83. Colour

84. Ornamentation (wet/dry)

Exothecial cells

85. Shape

86. Size

87. Papillosity/ mamillosity

88. Differentiation (apophysis to mouth) Stomata

89. Number

90. Position

91. Structure

Separating annulus

92. Absent/present. Description

Peristome

Exostome

93. Colour

94. Orientation (wet/dry)

95. Lower outside ornamentation

96. Upper outside ornamentation

97. Margin

98. Border

Endostome

99. Colour

100. Basal membrane height (%) Segments

101. Perforation

102. Ornamentation Cilia

103. Number

104. Development

105. Appendiculate/nodose

Lid

106. Description

Calyptra

107. Shape

108. Ornamentation

Spores

109. Size

110. Ornamentation

APPENDIX 3: Specimens erroneously identified.

Specimens erroneously identified as Calliergon cordifolium.

SPAIN. Caceres: 30TTK56, Castanar de Hervas, SALA-Bryo 248 is Brachythecium rivulare (Elias Rivas, 1988). Guipuzcoa: 30TWN45, Monte Garagarza (Monte Gartzaga), PAMP 3267 is Straminergon stramineum. Navarra: 30TXN68, Zurian, PAMP 3268 is Straminergon stramineum. Salamanca: 3OTTK67, Sierra de Bejar, Circo de la Pena Negra, SALA-Bryo 241 is Brachythecium rivulare (Elias Rivas, 1988).

Specimens erroneously identified as Calliergon giganteum. SPAIN. ASTURIAS: 30TQH39, Grao, Cuellagar, MA-Musci 15981 is Calliergon cordifolium.

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 SYSRESOURCE 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. Many thanks to Dr. Len Ellis and Dr. Lars Hedenas, curators of the bryophyte collections at BM and S, for their assistance during the visits of G. O. to these institutions, and to the latter for his comments on the manuscript and his invaluable guidance.

Recibido 23 Mayo 2005

Aceptado 19 Julio 2005

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Gisela Olivan, Esther Fuertes (*) & Margarita Acon (**)

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

(**) Departamento de Biologia (Botanica). Facultad de Ciencias. Universidad Autonoma de Madrid. E-28049 Madrid, Spain.
Tabla 1
Differential characters among Straminergon and Calliergon species
in the Iberian Peninsula

                Straminergon       Calliergon         Calliergon
                stramineum         cordifolium        giganteum

Branching       Unbranched or      Sparsely           Densely and
pattern         sparsely           branched to        pinnately
                branched           irregularly        branched
                                   branched

Axilary hairs   Scarce and small   Abundant and       Abundant and
                                   large              large

Leaf shape      Ovate,             Broadly ovate or   Broadly
                elongate-ovate     ovate-cordate      ovate-cordate
                or lingulate

Costa           Single, up to      Single, almost     Single, almost
                80% of length      reaching           reaching the
                leaf               the apex,          apex, 90-190 pm
                                   (60)80-100(120)    wide at base
                                   [micro]m wide
                                   at base

Alar groups     Longitudinally     Triangular,        Triangular
                ovate or           usually reaching   usually not
                rectangular        the costa          reaching the
                along margin (up                      costa (75-90% of
                to 50% of                             distance from
                distance from                         leaf margin to
                leaf margin to                        costa)
                costa)

Sexuality       Dioicous           Autoicous          Dioicous
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