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Tortella rigens (Bryophyta, Pottiaceae): relationships, regional variation, and conservation aspects

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Abstract

Tortella rigens occurs in the European Baltic region and at a few localities in North America. At least 95 % of all localities are found in southern Sweden. It is morphologically similar to the more widespread T. bambergeri, which was found in Sweden during this study. Haplotype networks, based on ITS or the chloroplast markers atbB-rbcL and rps4, for T. rigens, T. bambergeri, and closely related species display significant reticulation and NeighborNet and Jacknife analyses are used to explore the species’ relationships. Significant incongruence is found between the two molecular data sets or between these and the morphologically defined species, and several species appear polyphyletic or paraphyletic. Sporophytes of T. rigens are described and appear to be a result of hybridisation, which could partly explain the reticulation. Among the four South Swedish T. rigens populations, those of Gotland (GTL) and the Stockholm archipelago (SAR) are not differentiated molecularly from each other and there are signs of significant migration between them. The ones in Västergötland (VAS) and Öland (OEL) are distinct from each other and from GTL and SAR. The haplotype pattern of GTL displays signs of population increase whereas a decrease is found for VAS. Although the time scale is uncertain, this decrease is of conservation concern, considering that VAS has one unique haplotype. OEL has both the highest number of unique haplotypes (4) and the highest haplotype diversity of all four populations and has therefore the highest importance for conservation. Tortella rigens and T. bambergeri can be differentiated by atpB-rbcL.

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Acknowledgments

I thank Keyvan Mirbakhsh for her efficient laboratory work, Tomas Hallingbäck and Heribert Köckinger for discussions and for providing important material, and the curators of CANM, NY, and TU for loans. Two reviewers provided valuable input on an earlier version of this paper. This study was funded by the Swedish Environmental Protection Agency, Naturvårdsverket (No. NV-10009-12).

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  1. Department of Botany, Swedish Museum of Natural History, Box 50007, 104 05, Stockholm, Sweden

    Lars Hedenäs

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  1. Lars Hedenäs
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Correspondence to Lars Hedenäs.

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Handling editor: Jochen Heinrichs.

Electronic supplementary material

Below is the link to the electronic supplementary material.

606_2014_1159_MOESM1_ESM.pdf

Online Resource 1 Aligned sequences for atpB-rbcL, rps4, and ITS for Tortella rigens and T. bambergeri. For each species the different haplotypes, based on the respective markers, are displayed. (PDF 36 kb)

Appendices

Appendix 1

Specimen data and GenBank accession numbers for the studied specimens. Data format: Sample No.: [for Tortella rigens: REGIONAL POPULATION (see Fig. 4b)]; Locality; Coll. Year, Collector (collector’s No.) (LH = L. Hedenäs); Herbarium acronym: registration No.; GenBank Accession numbers for ITS, atpB-rbcL, rps4. Sample numbers for specimens which sequences were downloaded from GenBank begin with ‘GB’

Tortella arctica (Arnell) Crundw. & Nyholm: P138: Sweden. Jämtland, Åre, 2010, LH; S: B182517; KM020640, KM020530, KM020750. Tortella bambergeri (Schimp.) Broth.: P091: Sweden. Dalsland, Skållerud, 1985, T. Hallingbäck 44376; S: B184928; KM020617, KM020506, KM020726. P095: Austria. Oberoesterreich, Koppenwinkel Alm, 1962, J. Froehlich; S: B196096; KM020611, KM020500, KM020720. P096: Austria. Kärnten, Kanziani-Berg, 1975, J. Poelt (Pl. Graec., Bryoph. 2); S: B196097; KM020612, KM020501, KM020721. P097: Austria. Steiermark, Lavanttal, 1988, H. Pittoni & J. Poelt (Pl. Graec., Bryoph. 145); S: B196098; KM020613, KM020502, KM020722. P104: Sweden. Västergötland, Djupadalen, 1974, T. Hallingbäck 44375; S: B184932; KM020620, KM020509, KM020729. P129: Hungary. (no locality provided), 1936, A. Boros; S: B186073; KM020614, KM020503, KM020723. P130: United Kingdom. Scotland. Gleann Beag, 1963, A.C. Crundwell; S: B196677; KM020615, KM020504, KM020724. P131: Austria. Nideroesterreich, Kuhschneeberg, 1951, J. Froehlich; S: B196678; KM020616, KM020505, KM020725. P151: Sweden. Dalsland, Steneby, 1986, T. Hallingbäck 44374; herb. Hallingbäck; KM020618, KM020507, KM020727. P153: Sweden. Dalsland, Skållerud, 2011, T. Hallingbäck 5755; herb. Hallingbäck; KM020619, KM020508, KM020728. P200: Sweden. Södermanland, Utö, 2013, LH; S: B198239; KM020621, KM020510, KM020730. Tortella densa (Lorentz & Molendo) Crundw. & Nyholm: P136: Sweden. Gotland, Boge, 1997, LH; S: B42917; KM020638, KM020528, KM020748. P137: Norway. Hordaland, Stord, 1990, N. Hakelier; S: B184679; KM020639, KM020529, KM020749. GB1de: Ireland. Claire, 1992, E. Wiltshire; BM; AY854412, AY950332, AY950377. Tortella fragilis (Hook. & Wilson) Limpr.: P098: Sweden. Södermanland, Utö, 2010, LH; S: B176012; KM020622, KM020511, KM020731. P099: Sweden. Pite Lappmark, Arjeplog, 2006, LH et al.; S: B114557; KM020623, KM020512, KM020732. P132: Norway. Finnmark, Söröysund, 2001, LH; S: B59844; KM020624, KM020513, KM020733. GB1fr: Greenland, F.J.A. Daniels; MSUN: 0795-869.5; AY854417, AY950337, AY950382. GB2fr: Russia. Gorno-Altai, 1991, M. Ignatov; BM; AY854416, AY950336, AY950381. Tortella inclinata (R.Hedw.) Limpr.: P113: Sweden. Södermanland, Mörkö, 2003, T. Hallingbäck 39628; S: B184931; KM020634, KM020524, KM020744. P117: Sweden. Södermanland, Mörkö, 1984, LH; S: B196111; KM020635, KM020525, KM020745. P134: Sweden, Värmland, Filipstad, 2010, LH & G. Odelvik; S: B179320; KM020636, KM020526, KM020746. P135: Sweden. Västmanland, Sala, 2011, LH et al.; S: B185169; KM020637, KM020527, KM020747. GB1in: Germany. Northrhine-Westphalia, C. Schmidt; BM: 000824494; AY854420, AY950340, AY950385. Tortella rigens Alberts.: P089: VAS; Sweden. Västergötland, Dala, 1991, N. Hakelier; S: B195852; KM020536, KM020426, KM020646. P090: VAS; Sweden. Västergötland, Högstena, 1974, T. Hallingbäck 44373; S: B184934; KM020537, KM020427, KM020647. P092: OEL; Sweden. Öland, Sandby, 2005, P. Lundquist 1198842; S: B148156; KM020549, KM020439, KM020659. P093: OEL; Sweden. Öland, Kastlösa, 2005, M. Arnesson 1198767; S: B148179; KM020550, KM020440, KM020660. P094: SAR; Sweden. Uppland, Djurö, 2011, LH; S: B187057; KM020565, KM020455, KM020675. P102: VAS; Sweden. Västergötland, Högstena, 1985, N. Hakelier; S: B195851; KM020538, KM020428, KM020648. P103: VAS; Sweden. Västergötland, Österplana, 1961, N. Hakelier; S: B195850; KM020539, KM020429, KM020649. P105: VAS; Sweden. Västergötland, Högstena alvar, 1974, T. Hallingbäck 42384; S: B184933; KM020540, KM020430, KM020650. P106: VAS; Sweden. Västergötland, Österplana hed, 1975, T. Hallingbäck 44377; S: B184936; KM020541, KM020431, KM020651. P107: VAS; Sweden. Västergötland, Dala, 1974, E. Nyholm; S: B184787; KM020542, KM020432, KM020652. P108: VAS; Sweden. Västergötland, Österplana, 1975, G. Bråvander; S: B81291; KM020543, KM020433, KM020653. P109: VAS; Sweden. Västergötland, Valtorp, 1942, A. Hülphers; S: B184759; KM020544, KM020434, KM020654. P110: VAS; Sweden. Västergötland, Vilske-Kleva, 1942, A. Hülphers; S: B184785; KM020545, KM020435, KM020655. P111: VAS; Sweden. Västergötland, Södra Kyrketorp, 1946, N. Albertson; S: B184767; KM020546, KM020436, KM020656. P112: OEL; Sweden. Öland, Gräsgård, 2005, P. Stjernfeldt 1198769; S: B148190; KM020551, KM020441, KM020661. P114: SAR; Sweden. Södermanland, Utö, 1983, LH; S: B196110; KM020566, KM020456, KM020676. P115: SAR; Sweden. Södermanland, Mörkö, 1985, LH; S: B184796; KM020567, KM020457, KM020677. P116: SAR; Sweden. Södermanland, Ornö, 1983, LH; S: B196109; KM020568, KM020458, KM020678. P118: OEL; Sweden. Öland, Södra Möckleby, 2005, P. Lundquist 1198752; S: B148185; KM020552, KM020442, KM020662. P119: OEL; Sweden. Öland, Böda, 1962, A.C. Crundwell & E. Nyholm; S: B77839; KM020553, KM020443, KM020663. P120: OEL; Sweden: Öland, Föra, 1960, A.C. Crundwell & E. Nyholm 60-4; S: B184705; KM020554, KM020444, KM020664. P121: OEL; Sweden. Öland, Ås, 2005, M. Beckman 1198759; S: B148148; KM020555, KM020445, KM020665. P122: OEL; Sweden. Öland, Vickleby, 2005, M. Aronsson 1198735; S: B148163; KM020556, KM020446, KM020666. P123: OEL; Sweden. Öland, Södra Möckleby, 2005, P. Lundquist 1198751; S: B148184; KM020557, KM020447, KM020667. P124: OEL; Sweden. Öland, Ventlinge, 2005, M. Aronsson; S: B148292; KM020558, KM020448, KM020668. P125: OEL; Sweden. Öland, Hulterstad, 2005, M. Miller 1198766; S: B148149; KM020559, KM020449, KM020669. P126: OEL; Sweden. Öland, Resmo, 2005, M. Beckman 1198764; S: B148168; KM020560, KM020450, KM020670. P127: OEL; Sweden. Öland, Bårby, 2006, T. Hallingbäck & N. Lönnell 43866; S: B184937; KM020561, KM020451, KM020671. P128: OEL; Sweden. Öland, Möckelmossen, 2006, T. Hallingbäck & N. Lönnell 43898; S: B184939; KM020562, KM020452, KM020672. P140 (Originally identified as Tortella humilis by P. M. Eckel in 1987): SAR; Sweden. Uppland, Runmarö, 1984, J.T. Johansson; S: B184798; KM020569, KM020459, KM020679. P154: VAS; Sweden. Västergötland, Västerplana, 2012, T. Hallingbäck 6183; herb. Hallingbäck; KM020547, KM020437, KM020657. P155: VAS; Sweden. Västergötland, Södra Kyrketorp, 2003, T. Hallingbäck 39536; herb. Hallingbäck; KM020548, KM020438, KM020658. P156: CAN; Canada. Ontario, Renfrew, 1987, R.R. Ireland 22737; NY: 00134127; KM020598, KM020488, KM020708. P157: CAN; Canada. Ontario, Renfrew, 1987, R.R. Ireland 22735; CANM: 304175; KM020599, KM020489, KM020709. P158: CAN; Canada. Ontario, Algoma, 1989, R.R. Ireland 24437; CANM: 314373; KM020610, KM020499, KM020719. P159: CAN; Canada. Ontario, Carleton, 1978, J. Reddoch; CANM: 260653; KM020600, KM020490, KM020710. P160: CAN; Canada. Ontario, Ottawa-Carleton, 1988, D.F. Brunton 8639; CANM: 308477; KM020601, KM020491, KM020711. P161: EST; Estonia. Saare mk, Kärla, 2012, K. Vellak; TU: 168632; KM020602, KM020521, KM020742. P162: EST; Estonia. Harju Co., Väiki-Pakri, 2008, N. Ingerpuu; TU: 162045; KM020603, KM020492, KM020712. P163: EST; Estonia. Saare Co., Kärla, 2008, K. Vellak; TU: 168765; KM020604, KM020493, KM020713. P164: EST; Estonia. Saaremaa, Lümanda, 2010, S. Pihu; TU: 168246; KM020605, KM020494, KM020714. P166: SAR; Sweden. Södermanland, Nämdö, 2013, LH; S: B196865; KM020570, KM020460, KM020680. P167: SAR; Sweden. Södermanland, Nämdö, 2013, LH; S: B196870; KM020571, KM020461, KM020681. P168: SAR; Sweden. Södermanland, Trosa-Vagnhärad, 2013, LH & I. Bisang; S: B196734; KM020572, KM020462, KM020682. P169: SAR; Sweden. Södermanland, Trosa-Vagnhärad, 2013, LH & I. Bisang; S: B196736; KM020573, KM020463, KM020683. P170: SAR; Sweden. Uppland, Djurö, 2013, LH; S: B196684; KM020574, KM020464, KM020684. P171: SAR; Sweden. Uppland, Djurö, 2013, LH; S: B196694; KM020575, KM020465, KM020685. P172: SAR; Sweden. Uppland, Djurö, 2013, LH; S: B196688; KM020576, KM020466, KM020686. P173: SAR; Sweden. Södermanland, Nämdö, 2013, LH; S: B196885; KM020577, KM020467, KM020687. P174: FIN; Finland. Åland, Eckerö, 2013, LH; S: B197025; KM020606, KM020495, KM020715. P175: FIN; Finland. Åland, Eckerö, 2013, LH & I. Bisang; S: B197029; KM020607, KM020496, KM020716. P176: FIN; Finland. Åland, Eckerö, 2013, LH & I. Bisang; S: B197037; KM020608, KM020497, KM020717. P177: GTL; Sweden. Gotland, Fleringe, 2013, LH & I. Bisang; S: B196893; KM020583, KM020473, KM020693. P178: GTL; Sweden. Gotland, Bunge, 2013, LH & I. Bisang; S: B196898; KM020584, KM020474, KM020694. P179: GTL; Sweden. Gotland, Fleringe, 2013, LH & I. Bisang; S: B196900; KM020585, KM020475, KM020695. P181: GTL; Sweden. Gotland, Hellvi, 2013, LH; S: B196908; KM020586, KM020476, KM020696. P182: GTL; Sweden. Gotland, Hellvi, 2013, LH; S: B196910; KM020587, KM020477, KM020697. P183: GTL; Sweden. Gotland, Hellvi, 2013, LH; S: B196912; KM020588, KM020478, KM020698. P184: GTL; Sweden. Gotland, Hellvi, 2013, LH; S: B196917; KM020589, KM020479, KM020699. P185: GTL; Sweden. Gotland, Hellvi, 2013, LH; S: B196923; KM020590, KM020480, KM020700. P186: GTL; Sweden. Gotland, Hangvar, 2013, LH; S: B196929; KM020591, KM020481, KM020701. P187: GTL; Sweden. Gotland, Hangvar, 2013, LH; S: B196934; KM020592, KM020482, KM020702. P188: GTL; Sweden. Gotland, Stenkyrka, 2013, LH; S: B196937; KM020593, KM020483, KM020703. P189: GTL; Sweden. Gotland, Stenkyrka, 2013, LH; S: B196945; KM020594, KM020484, KM020704. P190: GTL; Sweden. Gotland, Hangvar, 2013, LH; S: B196950; KM020595, KM020485, KM020705. P191: OEL; Sweden. Öland, Mörbylånga, 2005, M. Beckman 1198738; S: B148169; KM020563, KM020453, KM020673. P192: OEL; Sweden. Öland, Gårdby, 2005, M. Miller 1198732; S: B148152; KM020564, KM020454, KM020674. P193: GTL; Sweden. Gotland, Hellvi, 2013, LH; S: B196915; KM020596, KM020486, KM020706. P194: GTL; Sweden. Gotland, Endre, 1989, LH G89-36; S: B197080; KM020597, KM020487, KM020707. P195: SAR; Sweden. Uppland, Djurö, 2013, LH; S: B196689; KM020578, KM020468, KM020688. P196: SAR; Sweden. Uppland, Djurö, 2013, LH; S: B196691; KM020579, KM020469, KM020689. P197: SAR; Sweden. Södermanland, Trosa-Vagnhärad, 2013, LH & I. Bisang; S: B196737; KM020580, KM020470, KM020690. P198: SAR; Sweden. Södermanland, Nämdö, 2013, LH; S: B196873; KM020581, KM020471, KM020691. P199: SAR; Sweden. Södermanland, Nämdö, 2013, LH; S: B196876; KM020582, KM020472, KM020692. Tortella tortuosa (Hedw.) Limpr.: P100: Sweden. Gotland, Stenkyrka, 2001, LH; S: B62094; KM020625, KM020514, KM020734. P101: Sweden. Medelpad, Borgsjö, 2006, LH; S: B116614; KM020626, KM020515, KM020735. P133: Sweden. Pite Lappmark, Arjeplog, 2006, LH et al.; S: B116150; KM020627, KM020516, KM020736. P139: Canada. NWT (Nunavut). Bathurst Island, 1973, R.R. Ireland & I. Brodo 16585; S: B196676; KM020609, KM020498, KM020718. P146: Austria. Styria, Eisenerzer Alpen, 2012, H. Köckinger 14932; S: B196697; KM020628, KM020517, KM020737. P147: Austria. Carinthia, Karawanken, 2012, H. Köckinger 14933; S: B196698; KM020629, KM020518, KM020738. P148: Austria. Carinthia, Gailtaler Alpen, 2008, H. Köckinger 14934; S: B196699; KM020630, KM020519, KM020739. P149: Austria. Carinthia, Gurktal, 2003, H. Köckinger 12334; S: B196700; KM020631, KM020520, KM020740. P150: United Kingdom. Northern Ireland, Fermanagh, 1993, T. Hallingbäck 42536; herb. Hallingbäck; KM020632, KM020522, KM020741. P152: Sweden. Dalsland, Skållerud, 1986, T. Hallingbäck 1452; herb. Hallingbäck; KM020633, KM020523, KM020743. OUTGROUP: Oxystegus tenuirostris (Hook. & Taylor) A.J.E.Sm.: P144: Sweden. Dalarna, By, 2011, LH et al.; S: B185165; KM020644, KM020534, KM020754. P145: Sweden. Närke, Fellingsbro, 1994, N. Hakelier; S: B196695; KM020645, KM020535, KM020755. Tortella humilis (Hedw.) Jenn.: P141: Spain. Murcia, Bullas, 2002, T. Hallingbäck 39035; herb. Hallingbäck; KM020641, KM020531, KM020751. GB1hu: Paraguay. Guaira, E. Zardini & P. Aquino 32386; BM; AY854419, AY950339, AY950384. GB2hu: United States. Arkansas, 1992, P.L. Redfearn Jr.; BM; AY854418, AY950338, AY950383. Trichostomum crispulum Bruch: P142: Sweden. Västergötland, Dala, 1984, N. Hakelier; S: B196696; KM020642, KM020532, KM020752. P143: Sweden. Gotland, Langs hage, 2002, T. Hallingbäck 38746; S: B185015; KM020643, KM020533, KM020753.

Appendix 2

The 68 specimens of Tortella rigens that were searched for male plants. Five arbitrarily chosen shoots were studied in each plant. All specimens are from Sweden or Finland (Åland) and were collected by Lars Hedenäs (LH), sometimes together with Irene Bisang (IB). The specimens are in S, and the ‘B’ numbers are S registration numbers

Gotland: Bunge: ca. 1 km W of Bunn, 12 May 2013, LH & IB: B196898; ca. 500 m E of Strå, 11 May 2013, LH & IB: B196886. Fleringe: ca. 1.2 km WSW of Tvärlingsmyr, 12 May 2013, LH & IB: B196900; ca. 1.6 km WSW of Tvärlingsmyr, 12 May 2013, LH & IB: B196907; ca. 400 m SW of Hau farm, 11 May 2013, LH & IB: B196889; ca. 800 m WSW of Hau farm, 11 May 2013, LH & IB: B196893; W end of Kyrkgatmyr, S side, 11 May 2013, LH & IB: B196896. Hangvar: central portion of Hajdhagen, 14 May 2013, LH: B196933; S portion of Hajdhagen, 14 May 2013, LH: B196934; Skarphagen, 14 May 2013, LH: B196950; Skarphagen, 14 May 2013, LH: B196950; W portion of Hajdhagen, 14 May 2013, LH: B196929. Hellvi: ca. 1 km SE of Sudergårde, 13 May 2013, LH: B196921; ca. 1,5 km SSE of Sudergårde, 13 May 2013, LH: B196923; ca. 200 m N of Lörge, 13 May 2013, LH: B196917; ca. 750 m WNW of Lörge, 13 May 2013, LH: B196920; just SW of Kyllaj, 13 May 2013, LH: B196915; NW end of Kyllajhajdar, 13 May 2013, LH: B196908; NW end of Kyllajhajdar, 13 May 2013, LH: B196909; NW end of Kyllajhajdar, 13 May 2013, LH: B196910; S end of Kyllajhajdar, 13 May 2013, LH: B196912. Lärbro: ca. 750 S of Stora Ire, 13 May 2013, LH: B196924. Stenkyrka: ca. 1 km N of Ekebys, 14 May 2013, LH: B196941; ca. 1.2 km N of Ekebys, 14 May 2013, LH: B196942; ca. 1.6 km N of Ekebys, 14 May 2013, LH: B196945; ca. 2 km NNW of Ekebys, 14 May 2013, LH: B196946; ca. 2 km NNW of Ekebys, 14 May 2013, LH: B196947; ca. 750 m NW of Bromyr, 14 May 2013, LH: B196949; Myrskogen, ca. 600 m W of Kvie, 14 May 2013, LH: B196937. Öland: Borgholm: N of Borgholm, 21 May 1980, LH: B198240. Södermanland: Nämdö:between Klippudden and Östanvik, 8 May 2013, LH: B196873; between Sandviksudden and Klippudden, 8 May 2013, LH: B196872; just ENE of Nämdö church, 8 May 2013, LH: B196885; Klockberget, 8 May 2013, LH: B196876; Kyrknäset, 8 May 2013, LH: B196877; Nämdö Böte, just inside N margin of summer house area, 8 May 2013, LH: B196865; Nämdö Nature Reserve, SW of Östanvik, 8 May 2013, LH: B196875; Sandviksudden, 8 May 2013, LH: B196868; SW of Sandviksudden, 8 May 2013, LH: B196870; Västanvik, 8 May 2013, LH: B196880; Västanvik, 8 May 2013, LH: B196883. Ornö: Sundby Nature Reserve, E of S end of Lake Långträsk, 29 July 2013, LH: B198246; Sundby Nature Reserve, SW end of peninsula E of Västra Varpet, 29 July 2013, LH: B198244. Trosa-Vagnhärad: SSE of Fänsåker, 1 May 2013, LH & IB: B196734; SSE of Fänsåker, 1 May 2013, LH & IB: B196735; SSE of Fänsåker, 1 May 2013, LH & IB: B196736; SSE of Fänsåker, 1 May 2013, LH & IB: B196737. Utö: Ängsbergen, 24 July 2013, LH: B198241; Ängsbergen, 24 July 2013, LH: B198242; Källviksberget, 24 July 2013, LH: B198243. Uppland: Djurö: Runmarö, between Noreträsk and Viträsk, 24 April 2013, LH: B196692; Runmarö, between Noreträsk and Viträsk, 24 April 2013, LH: B196693; Runmarö, between Noreträsk and Viträsk, 24 April 2013, LH: B196694; Runmarö, between Storsten and Nore, 24 April 2013, LH: B196688; Runmarö, between Storsten and Nore, 24 April 2013, LH: B196689; Runmarö, between Storsten and Nore, 24 April 2013, LH: B196690; Runmarö, N of Storsten, 24 April 2013, LH: B196684; Runmarö, N of Storsten, 24 April 2013, LH: B196685; Runmarö, N of Storsten, 24 April 2013, LH: B196686; Runmarö, N of Storsten (10 m from B196682), 24 April 2013, LH: B196683; Runmarö, N of Storsten (10 m from B196683), 24 April 2013, LH: B196682; Runmarö, W of Byholmen, 24 April 2013, LH: B196681; Runmarö, WNW of Nore, 24 April 2013, LH: B196691; Runmarö, WNW of Storsten, 24 April 2013, LH: B196687. Åland: Eckerö: Enskär, 350 m SE of old Coast Guard Station, 27 May 2013, LH: B197025; Signilskär, along small brooklet 300 m S of Mjölkarskatan, 25 May 2013, LH & IB: B197029; Signilskär, N of Bird Station, 24 May 2013, LH & IB: B197028; Signilskär, Söderskatan, 26 May 2013, LH & IB: B197037.

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Hedenäs, L. Tortella rigens (Bryophyta, Pottiaceae): relationships, regional variation, and conservation aspects. Plant Syst Evol 301, 1361–1375 (2015). https://doi.org/10.1007/s00606-014-1159-9

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