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Phytotaxa 73: 37–59 (2012) ISSN 1179-3155 (print edition) www.mapress.com/phytotaxa/ PHYTOTAXA Copyright © 2012 Magnolia Press Article ISSN 1179-3163 (online edition)

Common but new: rosamrosiae, a “new” widespread of apple (, Bryophytina) from the Mediterranean and western North America

LIA DAMAYANTI1, JESÚS MUÑOZ2,3, SUSANN WICKE1,4,5, LARS SYMMANK5, BLANKA SHAW6, JAN-PETER FRAHM1 & DIETMAR QUANDT1,5 1 Nees Institute of Biodiversity, Rheinische Friedrich-Wilhelms-Universität Bonn, Meckenheimer Allee 170, 53115 Bonn, Germany. Email: [email protected] 2 Real Jardín Botánico, CSIC, Plaza de Murillo 2, E-28014 Madrid, Spain. Email: [email protected] 3 Centro de Investigación de la Biodiversidad y Cambio Climático (BioCamb), Universidad Tecnológica Indoamérica, Quito, Ecuador 4 Institute for Evolution and Biodiversity Westfälische Wilhelms-Universität, Hüfferstrasse 1, 48149 Münster, Germany. 5 Institute of , Technische Universität Dresden, Zellescher Weg 20 b, 01217 Dresden, Germany. 6 Department of Biology, Duke University, Durham, North Carolina 27708, USA.

Abstract

Recent phylogenetic analyses challenged the traditional generic concept of the (apple mosses), especially with regard to the largest Bartramia. Although molecular analyses revealed the three Bartramia sections (Bartramia, Strictidium and Vaginella) to be monophyletic, they appeared in different parts of the inferred phylogenies and thus rendered the genus itself polyphyletic. In addition, Anacolia laevisphaera, a tropical montane species, appeared in the section Strictidium, weakening its character as a typical Mediterranean element. Although morphologically similar to members of the section, such as B. stricta, A. laevisphaera has been generally treated within the genus Anacolia based on sporophytic characters. In fact, Anacolia laevisphaera is often confused with B. stricta in sterile condition. A thorough analysis revealed another surprise, in placing A. laevisphaera sister to the Southern South American samples of Bartramia stricta, while the Mediterranean populations of “B. stricta” cluster with the Australian B. breutelii. Subsequent morphometric studies revealed three morpho-species in accordance with the polyphyletically resolved B. stricta. Here we describe the new species Bartramia rosamrosiae to accommodate the Mediterranean and western North American populations of what was traditionally treated as B. stricta.

Key words: Bartramiaceae, , Chile, mosses, phylogeny,

Introduction

Anacolia Schimper (1876: 513) and Bartramia (Hedwig 1801: 164–165) are two members of the Bartramiaceae, the only family of the order Bartramiales (Quandt et al. 2007), which belongs to the diplolepideous-alternate mosses. Two species of these genera have puzzled American bryologists for a long time, due to their similar gametophyte: Anacolia laevisphaera (Taylor 1846: 56) Flowers (1935: 155), described from the Pichincha volcano in Ecuador, and Bartramia stricta Bridel (1803: 132) described from the Magellan’s Strait after materials collected by Commerson. They could only be differentiated using sporophytic characters, as Anacolia laevisphaera has gymnostomous, rugulose capsules, while in Bartramia stricta they are striate and have a peristome. However, they are usually sterile, which lead to the assumption that the range of Anacolia laevisphaera reached the southern tip of South America. Nevertheless, Matteri (1983) was able to show gametophytic differences between the strictly southern South American B. stricta and the northern South American/Central American/African populations of Anacolia laevisphaera. But

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Matteri (1983) did not compare the South American populations of B. stricta with the northern temperate populations of the taxon so identified. “Bartramia stricta” had been considered a prominent member of the Mediterranean bryoflora since Schwägrichen (1816) identified some specimens collected by Desfontaines in “Barbaria” (Maghreb, North Africa) as Bartramia stricta, although he did not have the opportunity to study Commerson’s type material in Bridel’s herbarium. A year later, Bridel (1817), who indeed had Commerson’s on hand, endorsed Schwägrichen’s (1816) view that the Mediterranean plants pertain to Bartramia stricta, and so this name became firmly rooted in European (e.g., Bridel 1827, De Notaris 1838, Casares Gil 1915, Mönkemeyer 1927, Smith 2004, Ederra 2007) and North American bryology (e.g., Lawton 1971). In the frame of a phylogenetic analysis of the Bartramiaceae, Symmank (2007) performed a detailed analysis of Anacolia laevisphaera and Bartramia stricta s.s. to confirm Matteri’s (1983) conclusions, and also to resolve the situation of the northern temperate “B. stricta”. Symmank (2007) found that the “B. stricta” populations formed a strongly supported clade independent from the strongly supported clade of the southern South American populations, the origin of the type of B. stricta. Another interesting result was the sister relationship between Anacolia laevisphaera and B. stricta s.s. From Symmank’s (2007) results, it became clear that the three taxa pertain to Bartramia section Strictidium (Müller 1900: 352) Brotherus (1904: 640), along with Bartramia breutelii Müller (1858: 162), an Australian species recently monographed for the Flora of Australia (Bell 2006), but their relationships have to be ascertained. The aim of this study is, therefore, to morphologically characterize the northern temperate “Bartramia stricta”, to unravel its phylogenetic relationships with other members of Bartramia section Strictidium, and determine its precise geographic distribution.

Materials and methods

Taxon and DNA sampling

For the phylogenetic analyses, we selected 36 specimens representing, on the one hand Bartramia sect. Strictidium, and on the other Anacolia and Flowersia Griffin & Buck (1989: 372), a recent segregate of Anacolia. Within Bartramia sect. Strictidium, the sampling aimed at covering the geographic breadth of each taxon. The sampling was complemented with an additional 17 members of the family, representing the major lineages. In accordance with recent phylogenetic analyses (Quandt et al. 2007, Cox et al. 2010), two members of the Hedwigiaceae (Hedwigiales), Rhacocarpus purpurascens (Bridel 1812: 121) Paris (1900: 292) and Hedwigia ciliata (Hedwig 1801: 40) Palisot de Beauvois (1805: 15), were chosen as outgroups. For this selection of taxa, we sequenced the internal transcribed spacers of nuclear ribosomal DNA (ITS1 and 2), and the plastid trnS-rps4-trnT-trnL-trnF cluster (thereafter trnS-trnF), including the rps4 gene, four intergenic spacers separating the genes and a group I intron residing in trnL. All sequences are deposited in EMBL (European Molecular Biology Laboratory); accession numbers are listed together with voucher information in Table 1.

DNA Isolation, PCR-amplification and Sequencing

DNA was extracted using commercially available spin columns (Macherey-Nagel, Germany) following the manufacturer’s protocol. Methods for cleaning and grinding of plants prior to extraction and PCR protocols followed Olsson et al. (2009). Gel cleaned PCR products were sequenced by Macrogen Inc., South Korea (www.macrogen.com).

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TABLE 1. List of investigated specimens for the molecular analyses, with GenBank accession numbers for the regions sequenced including voucher and isolate numbers and the herbaria where the specimens are kept. * = voucher label Anacolia rozea; ** = voucher label Anacolia breutelii var. breutelii; *** = voucher label Anacolia cameruniae Dixon. Taxon Isolate Herbarium Voucher Accession numbers trnS-F ITS Outgroup (Hedwigiaceae/Hedwigiales) Rhacocarpus purpurascens (Brid.) Paris B345 Buchbender Buchbender s.n. 28.10.2000 HF536608 HF536712 Hedwigia ciliata (Hedw.) P. Beauv. B411 Buchbender Buchbender s.n. 16.12.2000 HF536653 HF536705 Bartramiaceae Schwägr. (Bartramiales D. Quandt, N.E. Bell & M. Stech) Anacolia Schimp. Anacolia menziesii (Turner) Paris B401 BONN Quandt s.n. 06.08.2006 HF536626 HF536675 (California) Anacolia menziesii (Turner) Paris B912 MUB Anonymous s.n. HF536627 HF536676 (Spain) (MUB-12959) Anacolia webbii (Mont.) Schimp. B861 MUB Anonymous s.n. HF536628 HF536677 (Spain 1) (MUB-11127) Anacolia webbii (Mont.) Schimp. B860 MUB Anonymous s.n. EF107540 HF536678 (Spain 2) (MUB -6204) Flowersia D.G. Griffin & W.R. Buck Flowersia abyssinica (Müll. Hal.) B453 H Anonymous s.n. (H-3002988) HF536649 HF536701 D.G. Griffin & W.R. Buck Flowersia campylopus (Schimp.) B045 BONN Cardenas 4369 HF536650 HF536702 D.G. Griffin & W.R. Buck Flowersia campylopus (Schimp.) B084 BONN Eggers & Frahm s.n. HF536651 HF536703 D.G. Griffin & W.R. Buck 19.02.1979 Flowersia sinensis (Broth.) B612 DUKE Shaw 5625 HF536652 HF536704 D.G. Griffin & W.R. Buck Sw. ex F. Weber & D. Mohr Conostomum cleistocarpum Herzog B685 BONN Lewis 87-1868 HF536647 HF536699 Conostomum pentastichum (Brid.) B042 BONN Frahm s.n. HF536648 HF536700 Lindb. (Bruch & Schimp.) Schimp. Breutelia affinis (Hook.) Mitt. B051 BONN Frahm s.n. 18.02.1998 HF536645 HF536697 Breutelia chrysea (Müll. Hal.) B62 NY Buck 39489 HF536646 HF536698 A. Jaeger (Duke) Brid. Philonotis andina (Mitt.) A. Jaeger B055 BONN Churchill, Magombo HF536656 HF536708 & Price 19810 Philonotis fontana (Hedw.) Brid. B054 BONN Garton 20748 HF536657 HF536709 (Hedw.) Brid. B15 DUKE Shaw M4 (4800) HF536658 HF536710 (Duke) Brid. Plagiopus oederianus (Sw.) H.A. B321 H van Rooy 1340 (H-3112041) HF536659 HF536711 Crum & L.E. Anderson Leiomela (Mitt.) Broth. Leiomela ecuadorensis H. Rob. B922 NY Churchill & Rengifo M. s.n. HF536654 HF536707 (NY-01206286) Leiomela bartramioides (Hook.) Paris B323 H Sloover 18792 HF536654 HF536706 Bartramia section Bartramia Hedw. E10020 BONN Quandt E10020 EF107539 HF536683 Bartramia mossmaniana Müll. Hal B442 BONN Bryo Austral No.11-6 HF536635 HF536686 ...... continued on the next page

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TABLE 1. (Continued) Taxon Isolate Herbarium Voucher Accession numbers trnS-F ITS Hedw. AS1006 Solga Solga AS1006 HF536636 HF536687 Bartramia section Vaginella Bartramia angustifolia Mitt. B081 BONN Lewis 87684 HF536629 HF536679 Bartramia aurescens Dixon B186 BONN Menzel, Frahm & Frey s.n. HF536630 HF536680 28.8.1986 Bartramia hampeana Müll. Hal. Bh114 BONN Vanderpoorten FSA 129 HF536633 HF536684 Brid. E10024 BONN Quandt E10024 HF536634 HF536685 Bartramia section Strictidium Bartramia breutelii Schimp. ex Müll.Hal. Australia 1 B020 H Streimann 058855 HF536631 HF536681 (H-3010990) Australia 2 B029 MO Streimann 058361 HF536632 HF536682 (MO-4433936) Bartramia laevisphaera (Taylor) Müll. Hal. Bolivia 1 B662 MA Anonymous s.n. HF536614 HF536663 (MA-Musci 15495) Bolivia 2* B076 BONN Lewis 38812d-1 HF536615 HF536664 Cameroon** B184 NY Chuah-Petiot M38 b HF536612 HF536661 Cape Verde B009 BONN Lindlar 459 HF536616 HF536665 . Colombia 1 Murcia- NY Anonymous s.n. (NY 17246) HF536620 HF536669 994 Germany 1 BsDN1 BONN Quandt Bs DN 1 HF536609 HF536713 Germany 2 BsM1 BONN Korneck & Frahm s.n. HF536610 HF536714 Pommern, 03.01.1198 Kenya B211 Townsend Townsend 85/537 HF536621 HF536670 Mexico 1 B036 MO Cardenas s.n. HF536622 HF536671 (MO-4435028) Mexico 2 BN5504 BONN/MO Cardenas 5504 HF536623 HF536672 South Africa*** B014 MO van Rooy 97549001 HF536611 HF536660 (MO-5377286) Tanzania 1 B179 H Laaka 900 HF536624 HF536673 Tanzania 2 B364 BONN Ochyra No 126-150 HF536625 HF536674 Bartramia rosamrosiae Damayanti, J.Muñoz, J.-P.Frahm & D.Quandt, sp. nov. California 1 BsC1 BONN Norris 84975 HF536637 HF536688 California 2 BsC3 BONN Norris 80910 HF536638 HF536689 Canada B021 UBC Schofield 55398 (B-35139) HF536639 HF536690 Spain BS Sp 12 BONN Quandt Bs Sp 12 (X10285) HF536640 HF536691 Great Britain GBA5 BONN Ferguson s.n. Population A5 HF536641 HF536692 Morocco B837 BONN Quandt s.n. 18.03.2010 HF536642 HF536693 Saudi Arabia 1 B074 B Frey & Kürschner 1563 - HF536694 Saudi Arabia 2 BsSA2 B Frey & Kürschner 1564 HF536643 HF536695 Turkey BsT3 (2) BONN Quandt Bs T 3 (2) HF536644 HF536696 Bartramia stricta Brid. Argentina B018 H Matteri 3269/84 HF536613 HF536662 (H-3011010) Chile 1 BsChile4 DR Müller C 1088 HF536617 HF536666 Chile 2 B026 DR Müller C 445 HF536618 HF536667 Chile 3 B027 DR Müller C 1552 HF536619 HF536668

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Alignment and tree inference

Sequences were edited and manually aligned using PhyDE® version 0.996 (Müller et al. 2005, available at http://www.phyde.de) following alignment rules based on strict motif reconstruction as advocated in Kelchner (2000) and Quandt & Stech (2005). Primer sequences were clipped off. The reported hairpin associated inversion in P6 of the trnL group I intron (Quandt & Stech 2005) was positionally isolated in the alignment and included in the analysis as reverse complement in order to gain information from substitutions within the detected inversion, as discussed in Quandt et al. (2003) and Borsch & Quandt (2009). Alignments are available on request from the authors. Indels were incorporated as binary data using a simple indel coding (SIC) strategy (Simmons & Ochoterena 2000) as implemented in SeqState (Müller 2005, 2006). Parsimony (MP) ratchet analyses (Nixon 1999) in PAUP 4.0b10 (Swofford 2003) were performed using command files generated with PRAP2 (Müller 2004, available at http://bioinfweb.info/Software/PRAP2) implementing 10 random addition cycles of 200 iterations each, with 25% upweighting of the characters in the iterations. Maximum Likelihood (ML) reconstructions were done using RAxML (Stamatakis 2006). Internal branch support in parsimony and likelihood approaches was estimated by heuristic bootstrap searches with 1000 replicates each. Bayesian analyses were performed with MrBayes v3.1.2 (Huelsenbeck & Ronquist 2001), applying the GTR+Γ+I model for the sequence data and the restriction site model (F81) for the binary indel partition. To allow for possibly deviating model parameters among the different regions, the data set was further divided into two sequence partitions (partition 1: plastid; partition 2: nrDNA). Four runs with four chains (5 x 106 generations each) were run simultaneously, with a chain sampling every 1000 generations. Tracer v1.4 (Rambaut & Drummond 2007) was used to examine the log likelihoods to determine the effective sampling size and stationary of the MCMC search. Calculations of the consensus tree, including clade posterior probability were performed based upon the trees sampled after the chains converged (< generation 500 000). Consensus topologies and support values from the different methodological approaches were compiled and drawn using TreeGraph2 (Stöver & Müller 2010). Phylogenetic analyses were conducted on each partition separately. No incongruences were detected. Therefore, only the results from analyses based on the combined data matrix will be presented in the following. Full clade support requires a bootstrap (BS) value of 100 and a posterior probability (PP) of 1.0. Significant support is considered >0.95 for PP and >90 for BS, while the borders for strong support are a little lower: 0.95 for PP and 80 in BS. Moderate support goes with PP >0.90 and BS >70. PP values below 0.90 and BS values below 70 are not considered supportive. Support values are given in the following order: BS from ML analyses (MLBS) / PP from analyses without indel coding (PP), BS from MP analyses including indel coding (MPBSsic) / and PP from analyses with indel coding (PPsic).

Morphological analysis

The morphological characterization was based on herbarium material following standard procedures. Leaves were taken from the tips, middle part and base of mature shoots, and cross sections of leaves and stems were done manually. About 250 specimens were studied, representing three Strictidium clades, namely, Anacolia laevisphaera, southern South American populations of Bartramia stricta, and Mediterranean and North American populations of “Bartramia stricta” (Appendix 1), of which a representative set of 75 specimens (25 specimens of each taxon) were chosen randomly for statistical analysis (Appendix 2). The studied specimens are kept at B, BONN, CAVA, DR, H, K, KOCH, MA, MO, MUB, NMW, NY, and UBC. Characters considered to be important in Bartramiaceae taxonomy (compare Matteri 1983, Fransen 1995, 2004) were measured: thickness of the stem cortex cells, stem diameter, central strand diameter, proportion of central strand/stem diameter, shape of leaf base (triangular vs. oblong), leaf orientation under dry and wet conditions (appressed to spreading), basal margin curvature (recurved vs. flat), shape of marginal cells at distal leaf (linear vs. oblong), shape of distal marginal teeth (nearly isodiametric vs. linear), and number of cells between

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TABLE 2. Characters used in the morphometric analysis and their corresponding code in Figure 2. Factor 1 and 2 indicate the respective weights of each character in the PCA. The table is sorted by descending weights for F1. Character id Character description F1 F2 1 Shape of leaf base 0.967 0.175 3 Leaf base margin 0.967 0.175 4 Shape of distal marginal cells 0.967 0.175 2 Leaf tips orientation 0.959 0.115 10 Leaves orientation dry 0.866 0.087 9 Plant density 0.780 -0.170 11 Leaves’ orientation wet 0.609 -0.154 15 Thickness of cortex cell walls (µm) 0.602 -0.505 12 Stem diameter (µm) 0.381 0.510 13 Central cylinder diameter (µm) -0.124 0.947 7 Size of marginal teeth -0.321 0.352 14 Central cylinder/stem proportion -0.415 0.847 8 Maximum plant height -0.699 0.127 5 Maximum number of cells between teeth -0.949 -0.152 6 Marginal teeth orientation -0.967 -0.175

Results

Alignment and phylogenetic reconstructions

The combined alignment of the plastid trnS-F region (1988 characters, 221 pi [parsimony informative]) and the nuclear ITS (2252 characters, 277 pi) contained 4240 characters (498 pi) that were complemented by 678 binary sited obtained from a simple indel coding approach, adding another 378 pi sites. The parsimony ratchet analysis of the combined matrix retained 18 trees (length 2247, CI = 0.749, RI = 0.888), while 39 trees (length 1378, CI = 0.730, RI = 0.892) were retained once indels were excluded. Both analyses, however, resulted in congruent trees with similar support values, but with slightly less resolution and support in the latter. The obtained maximum likelihood topology tree is depicted in Figure 1, complemented with support values from the different analyses. The backbone topology is fully resolved with generally significant posterior support and moderate to strong bootstrap support (likelihood and parsimony). The only exception concerns the “monophyly” of Bartramia that gains only moderate MLBS support while all other algorithms were not supportive. In addition it has to be noted that the two included members of the genus Leiomela (Mitten 1869: 253) Brotherus (1904: 634), which was formerly regarded as a subsection of Bartramia, are nested within Bartramia section Vaginella Müller (1849: 492). However, the first branching lineage of the Bartramiaceae consists of Anacolia

42 • Phytotaxa 73 © 2012 Magnolia Press DAMAYANTI ET AL. TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. as sister to Flowersia, followed by Conostomum Weber & Mohr (1804: 121) as sister to Breutelia (Bruch & Schimper 1851: 1) Schimper (1856: 85) and Philonotis Bridel (1827: 15–28). The following crown group represents Bartramia with Leiomela nested inside and Plagiopus Bridel (1826: 596) as first branching lineage. While the relationships among the three Bartramia sections remain unsupported, each section gains full support in all analyses, corroborating that the former subsection Leiomela is nested in section Vaginella, and renders Bartramia in its current circumscription paraphyletic. Phylogenetic relationships within section Vaginella are, however, lacking support. Anacolia laevisphaera is clearly resolved (100/1/100/1) within the section Strictidium as sister to Bartramia stricta accessions from South America. In contrast, the Mediterranean populations of “B. stricta”, incl. populations from California, British Columbia and Wales (UK) cluster with Bartramia breutelii populations from Australia with significant to full support in model based approaches, and moderate support in parsimony (MPBSsic 76).

FIGURE 1. Maximum likelihood tree based on the combined trnS-F and ITS matrix. Likelihood bootstrap percentages are depicted above (bold) the branch followed by posterior probabilities. Support values from the analyses including indel coding are depicted below the branches: with MPBSsic formatted in italics and PPsic underlined. Multivariate analyses

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Both the PCA (Fig. 2) and discriminant analysis (Fig. 3) differentiate the studied specimens into three groups. The characters related to the marginal teeth shape and structure, and the leaf shape and orientation clearly separate the northern Hemisphere populations of “B. stricta” in a separate group, while the central strand width, the cortex cells walls thickness, and size of the plants, allow separating Anacolia laevisphaera from the southern South America B. stricta (compare Appendix 5).

FIGURE 2. PCA diagrams for the first two factors, F1 and F2, with the variance explained by each. Italicized numbers correspond to the characters in Table 1. A: Bartramia (Anacolia) laevisphaera; M: Bartramia rosamrosiae sp. nov.; S: Bartramia stricta.

FIGURE 3. Canonical analysis based on the two discriminant functions F1 and F2 of the discriminant analysis. Gray dots are the cen- troids of each group.

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Discussion

Taxonomic studies on Bartramia sect. Strictidium led to the discovery that the taxon that was long recognized in Europe as “Bartramia stricta” pertains in fact to an undescribed taxon. It is common in Mediterranean-type climates of the Northern Hemisphere, but it has passed unnoticed until now. The combination of molecular and morphometric analysis mutually strengthen each other, and besides supporting the recognition of this new taxon, allow confirming that Anacolia laevisphaera is a member of Bartramia section Strictidium. In addition, Anacolia cameruniae Dixon (1933: 25) should be synonymised with B. laevisphaera as neither morphological nor genetic differences could be identified. Material identified as Anacolia breutelii var. breutelii from South Africa correspond to B. laevisphaera as well. Both molecular and morphometric analyses show that the Mediterranean populations (including populations from the US, Canada, and Great Britain), commonly identified as Bartramia stricta, differ from B. stricta s.s. and from Bartramia laevisphaera, and must be recognized as a new species. If Leiomela is to be recognized on genus level it ultimately renders Bartramia paraphyletic, even if the sectional relationships are not resolved with support. This implies either elevating the three currently recognized sections to generic level or downgrading Leiomela to sectional or subsectional rank. However, the recognition of Leiomela as a subsection of Bartramia or generic rank warrants further analyses as does the broad circumscription of Bartramia. Perhaps the inclusion of further taxa such as Quathlamba Magill (1987: 421) and additional members of section Vaginella and Bartramia will help to resolve this issue in the future.

Taxonomy

Bartramia laevisphaera (Taylor) Müller (1849: 506) ≡ Glyphocarpa laevisphaera Taylor (1846: 56) ≡ Glyphocarpus laevisphaerus (Taylor) Jaeger (1875: 64) ≡ Anacolia laevisphaera (Taylor) Flowers (1935: 155). Type:—ECUADOR. Pichincha: Without locality, 1846, W. Jameson s.n. (holotype BM!). ≡ Anacolia cameruniae Dixon (1933: 25), syn. nov. Type:—CAMEROON. Southwest Province: Mount Cameroon, 11000 ft, 12 January 1932, M. Steele 23 (holotype BM!).

Bartramia rosamrosiae Damayanti, J.Muñoz, J.-P.Frahm & D.Quandt, sp. nov. (Figs. 4 & 5).

Type:−SPAIN. Málaga: Ronda road A-372 to Grazalema, 7 km after crossing with C-339, 36°45'35.2"N, 5°20'7.4"W, 8 April 2001, D. Quandt BS Sp 12 (holotype BONN X10285!, isotype MA-Musci!).

A species of Bartramia that can be distinguished from B. laevisphaera by its smaller size (<40 vs. <75 mm), its wide central strand in the stems (occupying 0.20–0.40 vs. 0.10–0.25 of the total stem diameter), triangular leaves without shoulders (vs. leaves with an oblong leaf base and recurved margins), thinner walls of the stem cortex cells (less than 2 μm vs. 2–3 μm), striate capsules with peristome on a 10–15 mm long seta (vs. wrinkled, gymnostomous capsule on a short seta <5 mm long), and distal teeth consisting of linear cells that scarcely protrude (vs. short to nearly isodiametric teeth cells). The similar B. stricta has leaves with an oblong base distally ending in weakly distinct shoulders (vs. triangular in B. rosamrosiae), and teeth formed by one, rarely two isodiametric cells that protrude from the leaf margin. (Compare Figs. 4 & 5, Table 3)

Plants moderately robust, green above and reddish brown below, in dense, seldom loose tufts. Stems up to 40 mm high, 0.2–0.4 mm in diameter, erect to ascending, simple or fastigiate and then branches 7–20 mm long, straight or curved at tip, orange, brown or light brown, with abundant reddish rhizoids covering most of basal stem. Central strand wide, occupying 0.20–0.40 of stem diameter, cortex cells large, with thin walls (<2 µm). Leaves stiffly erect, appressed, not sheathing, sometimes somewhat homomallous when dry, erect to spreading when moist, triangular, rarely lanceolate with an almost indistinguishable oblong base and weak shoulders, never sheathing, 1.5–4.5 mm long; margins flat or shortly (0.1–0.6 mm) recurved at the extreme base, unistratose proximally, 2–3-stratose distally; singly or doubly serrate distally, marginal teeth linear,

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FIGURE 4. Comparison of: A. Bartramia laevisphaera. B. B. stricta. C. B. rosamrosiae sp. nov. First row: Sporophyte dry (scale bar 2 mm). Second row: Plant dry (scale bar 2 mm). Third row: Plant wet (scale bar 2 mm).

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FIGURE 5. Comparison of: A. Bartramia laevisphaera, B. B. stricta, C. B. rosamrosiae sp. nov.. First row: Cortex cells (scale bar 20

µm). Second row: Central strand illustrating the proportional differences (scale bar 100 µm). Third row: Leaf shape (scale bar A3 = 2 mm; B3–C3 = 1 mm) & distal margin (scale bar 50 µm). Last row): Cross section at leaf base (scale bar A4/B4 = 100 µm; C4 = 50 µm).

BARTRAMIA ROSAMROSIAE, A “NEW” SPECIES OF APPLE MOSSES Phytotaxa 73 © 2012 Magnolia Press • 47 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. seldom shorter, adjacent or with 1–2 cells between teeth; costa relatively slender, short-excurrent, 60–135 µm wide and 40–50 µm thick at base, dorsally rough in distal leaf; basal cells elongate-oblong to linear for 7–24 juxtacostal rows, isodiametric to short-oblong in 4–10 marginal rows, medial and distal cells smaller and narrower, thick-walled, prorulose on both sides. Sexual condition synoicous, gametoecium terminal, appearing lateral by innovations, perichaetial leaves lanceolate, differentiated from vegetative leaves, with oblong base and more distinct shoulders, limb sometimes linear; costa slender and narrower than in vegetative leaves. Seta solitary, reddish to orange brown, 10–15 mm, straight or scarcely curved, smooth. Capsule erect, ovoid to globose, striate when dry. 22–32 µm, reddish-brown, verrucose. Etymology:—Dedicated to our friend Rosa M. Ros (Murcia, Spain), in recognition of her outstanding contribution to the study of the Mediterranean bryophytes. Selected specimens examined (paratypes) [total 167, see Appendix 1]:—CANADA. British Colombia: Vancouver Island, head of Peddar Bay, 48°20’N, 123°29’W, 16 April 1974, W.B Schofield 55398 (UBC- B35139). USA. California: Shasta County, 40°35.7’N, 122°25.3’W, 150 m, 16 Febuary 2003, D.H. Norris 105288 (UC-1767483); San Francisco County, Glen Canyon, San Miguel Hills, 37°44’15’’N, 122°26’25’’W, 250 m, 29 January 2000, J.R. Shevock 18884 (UC-1739225); Sonoma County, 38°27’0’’N, 122°39’00’’W, 122–198 m, 22 April 1998, A.T. Whittemore & T.E. Whittemore 6696 (MO). FRANCE. Var: Massif des Maures, between Plan-de-la-Tour and Col de Gratteloup (D 44), 43°21’46.7’’N, 06°33’47.5’’E, 250 m, 1 April 2001, D. Quandt Bs F 5 (1) (BONN X10247). Corse: Marsiglia, road from Lenturia to Marsiglia, 42°57’14.2’’N, 09°22’29.2’’E, 25 April 2003, S. Wanke Bs Corse 3–1 (BONN X10269). PORTUGAL. Coimbra: 2 km after Degracias on the road N348 to Soure, 40°00’44.5’’N, 08°31’46’’W, 350 m, 12 April 2001, D. Quandt Bs P 6 (BONN). Madeira: Miradouro, January 2002, A. Vanderpoorten 1132 (BONN X10260, Herbarium Vanderpoorten). ITALY. Basilikata: Polino, between San Severino and Francavilla, July 2002, L. Gartung Bs I 5 (BONN). MOROCCO. Taza: Bab Taza, 35°04’45’’N, 05°10’05’’W, 1100 m, 17 March 1997, M.J Cano, M.T. Gallego & R.M. Ros 10516 (MUB: 10516). TURKEY. Antalya: Alanya, road from Payallar to Güzelbag, 1 km after Payallar, April 2003, D. Quandt Bs T 3 (BONN). SAUDI ARABIA. Makkah: Between Mekka and Taif, 1450–1600 m, W. Frey & H. Kürschner 1564 (B).

TABLE 3. Differential characters among the Bartramia (Anacolia) laevisphaera, B. stricta and B. rosamrosiae sp. nov. Characters B. laevisphaera B. stricta B. rosamrosiae Plant length <75 mm <45 mm <40 mm Ratio central strand to stem 0.10–0.25 0.20–0.40 0.20–0.40 diameter cortex cell wall thickness 2–3 μm<2 μm<2 μm Leaf shape lanceolate lanceolate triangular, rarely lanceolate Leaf base oblong oblong to rarely triangular triangular Leaf margin at base mostly recurved mostly recurved flat Leaf margin in upper half dentate, recurved dentate, recurved serrate, recurved Teeth cells short to nearly isodiametric short to nearly isodiametric linear Capsule surface in dry state rugulose striate striate Seta short (<5 mm) long (8–15 mm) long (10–15 mm) Peristome lacking present present

Ecology and distribution:—Bartramia rosamrosiae plants grow mostly at low elevations, at 50–1100 m. They are mostly found on soil, thin soil over rocks, or crevices in boulders, in shady and moist places, but also in dry to very dry or in temporarily/seasonally moist places, as in California. The nature of the substrate includes sedimentary rocks, slate, limestone, and also artificial concrete walls. They can be found near rivers, in open ‘shrubland’, open grasslands or in forests.

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The species is distributed in the northern hemisphere (Fig. 6), mostly occurring in areas with Mediterranean climate. In the western part of North America it is known from British Columbia (Canada) and California (USA). In Europe and Northern Africa it is relatively common in Mediterranean areas, and rare in cooler places with strong Atlantic influence, like Wales (UK) and the Bailiwick of Guernsey in the English Channel. It is known from Azerbaijan, Cyprus, France (including Corsica), Great Britain, Greece, Italy, Morocco, Portugal (including Madeira), Spain (including the Canary Islands), and Turkey. In addition it occurs in Saudi Arabia and Israel.

FIGURE 6. Distribution of the species of Bartramia sect. Strictidium: Cross = B. breutelii; Triangles = B. rosamrosiae sp. nov.; Stars = B. (Anacolia) laevisphaera; Squares = B. stricta.

Acknowledgments

This research was jointly funded by the German Science Foundation (DFG Fr 481/11-1 & Fr 481/11-2 to JPF), the Spanish Ministry of Education (CGL2009-09530-BOS to JM), the European Commission’s BIODIBERIA HUMAN POTENTIAL PROGRAMME (to DQ) and SYNTHESYS (to SW & DQ), which is financed by the European Community Research Infrastructure Action under the FP6 “Structuring the European Research Area” Programme (http://www.synthesys.info). We are grateful to the curators of the herbaria cited in the text for loan of specimens. Additional specimens were provided by C. C. Townsend (Twickenham) and Andrew Ferguson (Wales), which is greatly acknowledged. We are especially indebted to Frank Müller (Dresden), Bernard Goffinet (UConn) and Bill Buck (NYBG) for providing us with critical material from Chile and Jean-Pierre Hébrard (Marseille) for hosting and guiding field work in France during spring 2000 and 2001. We are grateful to Sven Fransén (Gothenburg) for discussion and providing us with a second opinion on critical specimens, and to Lars Hedenäs and Katia Cezón for critical comments on the manuscript. We also thank our long-lasting friends Olaf Werner and Rosa María Ros (both Murcia) for their help in innumerable occasions.

References

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Appendix 1. Specimens examined

Bartramia laevisphaera (Taylor) Müll. Hal. Anonymous (BONN X10286), (H-N.520), (MA-Musci 15495) Arts 12/005 (BONN), 12/010 (BONN) Cardenas 07/1995 (MO-4435028), S 1956 (BONN), S 5484 (BONN), S 5504 (BONN) Chuah-Petiot M38 b (NY) Churchill & Betancur-B. s.n. (MO-3953991), 18000 (HUA) De Sloover 19385 (Herb. Townsend) del Carman Ramirez & del Carman Ramirez 20613 (MO) Delgadillo M 1141 (BONN), M 2025 (BONN), 5803 (MO) Frahm 935 (BONN), 4561 (BONN), 6410 (BONN), 823921 (BONN) Frahm, Gradstein, Philippi, Schultze-Motel 866 (NY) Futschig TK 5515 (BONN), s.n. "16 September1967" (BONN) Garcia-M. s.n. " 29 May 1976" (BONN) Hinz s.n. "17 February 1975" (BONN) Jameson 30, 31 (BM, Type) Korneck & Frahm Elztal 1998 (BONN) Korneck & Frahm Pommern, 03. 01. 1998 (BONN) Laaka 868 (H-3194026), 900 (H-3194027) Lewis 38812 d-1 (MO-3960874), 78-2249 (H) Linares & Churchill 3717 (H) Lindlar 459 (BONN) Pocs et al. 88148/A (BONN, MO-4423125) Quandt Bs DE 1 (1) (BONN X10272), Bs DE 1 (2) (BONN X10273), Bs DE 2 (1) (BONN X10274), Bs DE 2 (2) (BONN X10275), Bs DN 1 (BONN X10276) Ramírez P. 9770 (MO) Shevock 17853 (MO-5211290) Stark 976 (MO) Steere 23115 (H) Townsend 85/426 (K), 85/537 (Herb. Townsend) van Rooy 3448 (BONN), 97549001 (MO-5377286), 97549001 (PRE-561175.0)

Bartramia rosamrosiae Damayanti, J. Muñoz, J.-P. Frahm & D. Quandt, sp. nov. Allen 344 (MO), s.n. (NY-438139), s.n. (NY-438141), s.n. (NY-438142), s.n. (NY-438156), s.n. (NY-438160) Baker MS 727 (UC) Baker s.n. (NY-438145) Bolander 256 (MO), 3191/21421 (MO), s.n. (NY-438150), s.n. (NY-438152) Cano, Gallego & Ros 10516 (MUM) Crundwell 43 (NMW) Cuynet 2I (BONN) De Merey s.n. (L-0279793) de Sloover 31316 (BONN) Düll & I. Düll Bs Gr 1 (BONN X10258), Bs Gr 2 (BONN X10257)

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Een & Persson s.n. (BONN) Ferguson Bs GB 5D (BONN X10279), Bs GB A5 (BONN X10278), Bs GB C2 (BONN X10277) Frahm 6410 (BONN), 2009683 (BONN), s.n. “4 April 1970” (BONN), s.n. “5 April 1973” (BONN) Frey & Kürschner 1563 (B), 1564 (B) Gartung Bs I 5 (BONN X10267) Gibbons s.n. (NY-438147), s.n. (NY-438148), s.n. (NY-438148) Haring 199 (NY-438164), 423 (MO), s.n. (NY-438165), s.n., (NY-438167), (NY-438168) Hébrard & Quandt Bs F 1 (1) (BONN X10234), Bs F 1 (2) (BONN X10235), Bs F 1 (3) (BONN X10236), Bs F 1 (4) (BONN X10237), Bs F 2 (1) (BONN X10238), Bs F 2 (2) (BONN X10239), Bs F 3 (1) (BONN X10240), Bs F 3 (2) (BONN X10241), Bs F 3 (3) (BONN X10242), Bs F 4 (1) (BONN X10243), Bs F 4 (2) (BONN X10244), Bs F 4 (3) (BONN X10245), Bs F 4 (4) (BONN X10246), Bs F 5 (1) (BONN X10247), Bs F 5 (2) (BONN X10248), Bs F 5 (3) (BONN X10249) Hébrard Bs F 1 (BONN), Bs F 2 (BONN), Bs F 3 (BONN), Bs F 4 (BONN) Hébrard, Frahm & Quandt Bs Var 1 (1) (BONN X10164), Bs Var 1 (2) (BONN X10165), Bs Var 1 (3) (BONN X10166), Bs Var 2 (1) (BONN X10167), Bs Var 2 (2) (BONN X10168), Bs Var 2 (3) (BONN X10169), Bs Var (3) (BONN X10170), Bs Var 4 (BONN X10171) Holmen 15.007 (BONN) Howe s.n. (NY-438143), s.n. (NY-438144) Hübschmann s.n. (BONN) Hugonnot Bs F-HL-1 (BONN X10259) Illata 1493 (BONN) INF [sic] 1434 (BONN), 4040 (BONN) Kimeridze & Gognidze Bs Azer 1 (BONN X10268) Koch 387 (NY-438158), 421 (MO), 1480 (NY-438157) Kürschner 940 (B) Loiseau CLF008247 (BONN) MacFadden 16871 (NY-438140), 16872 (NY-438155), 16873 (NY-438161), 16874 (NY-438162), 3193/17024 (MO), 3194/16871 (MO), 3195/16871 (MO), 3196/17024 (MO), 871/16871 (MO) Müller 1998 (DR) N.61-960 [sic] (BONN) Norris 47607 (MO, NY-438138), 101004 (UC), 105288 (UC), 108748 (UC), 108765 (UC) Nyholm & Crundwell 442 (NMW-C96.14 2408) Parry s.n. (NY-438151) Parsons s.n. (NY-438159) Quandt Bs U 3 (BONN X10231), Bs U 2 (BONN X10230), Bs U 1 (2) (BONN X10229), BS U 1 (1) (BONN X10228), Bs T 1 (BONN), Bs T 2 (BONN), Bs T 3 (1) (BONN), Bs T 3 (2) (BONN), Bs T 3 (3) (BONN), Bs Sp 1 (BONN), Bs Sp 2 (BONN), Bs Sp 3 (BONN), Bs Sp 4 (BONN), Bs Sp 5 (BONN), Bs Sp 6 (BONN), Bs Sp 7 (BONN), Bs Sp 8 (BONN), Bs Sp 9 (BONN), Bs Sp 10 (BONN), Bs Sp 11 (BONN), Bs Sp 12 (BONN X10285), Bs P 1 (BONN), Bs P 2 (BONN), Bs P 3 (BONN), Bs P 4 (BONN), Bs P 5 (BONN), Bs P 6 (BONN), Bs I 4 (2) (BONN X10233), Bs I 4 (1) (BONN X10232) Quandt & Frahm Bs Aude 1 (BONN X10162), Bs M 3 (1) (BONN X10161), Bs PO 1 (BONN X10172), Bs PO 2 (1) (BONN X10173), Bs PO 2 (2) (BONN X10174), Bs PO 2 (3) (BONN X10175), Bs PO 2 (4) (BONN X10176), Bs PO 2 (5) (BONN X10177), Bs PO 2 XXL (BONN X10178), Bs PO 3 (BONN X10179), Bs PO 4 (1) (BONN X10180), Bs PO 4 (2) (BONN X10181)

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Quandt & Ros Bs Sp 1 (1) (BONN X10250), Bs Sp 1 (2) (BONN X10251), Bs Sp 1 (3) (BONN X10252), Bs Sp 1 (4) (BONN X10253), Bs Sp 1 (5) (BONN X10254), Bs Sp 1 (6) (BONN X10255), Bs Sp 1 (7) (BONN X10256) Reed s.n. (NY-438146) Sapaly 84/088 (BONN), 84/667 (BONN), 92/004 (BONN) Sch430 [sic] (BONN) Schiffner s.n. (BONN) Schofield 68459 (NY-152754), 71733 (UBC) Schumm & Schwarz 3606 (BONN) Shevock 17853 (MO), 18884 (UC) Shevock & York 12897 (MO) Siegel s.n. (BONN) Stark & Castetter 1155-A (MO) Stark 580-B (MO), 784 (MO), 786 (MO), 796 (MO), 798 (MO), 833-A (MO), 971 (MO), 1251 (MO) Torrey 3 (NY-438153) Townsend 82/352 (BONN) Vanderpoorten Bs P 9 (BONN X10260), Bs P 10 (BONN X10264) Wallace s.n. (NMW-C96.14 2406) Wanke Bs Corse 3–1 (BONN X10269), Bs Corse 3–2 (BONN X10270) Weber 873/21296 (MO) Weber et al. 91959 (MO) Whittemore 5246 (NY-438137) Whittemore & Whittemore 3354 (MO), 5420 (MO), 6696 (MO) Whittemore & et al. 5347 (MO) Yurky 827 (NY-438163)

Bartramia stricta Bridel Bonavia 2269/84 (H) Buck 41544 (NY) Deguchi 25686 (KOCH), 26413 (KOCH) Gelain et al. 3394 (NY) Goffinet 7088 (H) Hatcher & Engel 239 (H), 726 (NY) Ireland & Bellolio s.n. (NY-30891), (NY-30929), (NY-30931), (NY-30992), (NY-31702), (NY-32175), (NY-32205), (NY-32436), (NY-32942), (NY-33226), (NY-33678), (NY-33871) Mahú 4827 (NY), 21350 (MO), 22081 (MO), 22127 (MO), 22316 (MO), 22618 (MO) Mahú & Mahú 24054 (MO), 24078 (MO) Matteri 3269/84 (H-3011010) Müller 818 (H), C1088 (DR), C1552 (DR), C445 (DR) Skottsberg 164 (H) Steere 60-262 (NY) Tapia 21135 (MO) Taylor & Gereau 11022 (H, MO, NY)

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APPENDIX 2. Specimens used for the morphometric analysis. JFI = Juan Fernandez Island. Codes for final identifications: BL = Bartramia laevisphaera; BR = Bartramia rosamrosiae; BS = Bartramia stricta. Scientific name on the Elevation Collection Nr. Country Date Collectors Herbarium voucher (masl) or Field Nr. BL 1 Anacolia laevisphaera Mexico 2180 29. Oct 1993 C. Delgadillo M 5803 MO BL 2 Anacolia laevisphaera Colombia 2860 11. May 1991 S. P. Churchill 18000 MO & J. Betancur B. BL 3 Anacolia laevisphaera Colombia 2850 05. Mar 1996 B. R. Ramírez P. 9770 MO BL 4 Anacolia laevisphaera Colombia 2900–2950 25. Aug 1992 E. Linares C. 3717 H & S. P. Churchill BL 5 Anacolia laevisphaera Dominican 2840 14. May 1992 W.C. Steere 23115 H Republic BL 6 Anacolia laevisphaera Ecuador 3980 1–2. Nov 1978 M. Lewis 782249 H BL 7 Anacolia laevisphaera Venezuela 3200 22. July 1984 N.A. 520 H BL 8 Bartramia stricta Germany 03. Jan 1998 Korneck & Frahm “Pommern BONN 1998” BL 9 Anacolia laevisphaera Kenya 3000 08. Feb 1985 C. C. Townsend 85/426 K BL 10 Anacolia laevisphaera Bolivia 3700 28 June 2001 S. P. Churchill & M 20613 MO del Carman Ramirez BL 11 Anacolia laevisphaera New 1935 11. Aug 1979 L. R. Stark 976 MO Mexico BL 12 Anacolia laevisphaera Mexico 2320 30. Nov 1994 A. Cardenas S 5484 BONN BL 13 Anacolia laevisphaera Mexico 2500 29. May 1976 E. Garcia M s.n. BONN BL 14 Anacolia laevisphaera Mexico 27. July 1968 C. Delgadillo M 2025 BONN BL 15 Anacolia laevisphaera Mexico 2980 23. Oct. 1966 C. Delgadillo M 1141 BONN BL 16 Anacolia laevisphaera Ecuador 2700 12. July 1991 T. Arts 12/010 BONN BL 17 Anacolia laevisphaera Ecuador 3000 17. Feb 1975 W. Hinz s.n. BONN BL 18 Anacolia laevisphaera Ecuador 2700 12. July 1991 T. Arts 12/005 BONN BL 19 Bartramia stricta Germany 03. Jan 1998 Korneck & Frahm s.n. BONN BL 20 Bartramia stricta Germany 15. April 1967 J. Futschig s.n. TK 5515 BONN BL 21 Bartramia stricta Germany 05. Nov 1933 N.A. X10286 BONN BL 22 Bartramia stricta Germany 12. April 2003 D. Quandt X10272 BONN BL 23 Bartramia stricta Germany 14 Dec 2003 D. Quandt X10274 BONN BL 24 Anacolia laevisphaera Mexico 3030–3070 02. Dec 1994 A. Cardenas S 5504 BONN BL 25 Anacolia laevisphaera Mexico 25. Sep 1982 A. Cardenas S 1956 BONN BR 26 Bartramia stricta USA 122–198 22. April 1998 A.T. Whittemore 6696 MO & T.E. Whittemore BR 27 Bartramia stricta Azerbaijan 01. Oct 1963 Kimeridze & X10268 BONN Gognidze BR 28 Bartramia stricta Spain 04. April 1970 J. P. Frahm s.n. BONN BR 29 Bartramia stricta Turkey 04. Mai 1982 C. C. Townsend 82/352 BONN BR 30 Bartramia stricta France 15. Mai 1976 A. v. Hübschmann s.n. BONN BR 31 Bartramia stricta USA 150 22. Feb 2001 D. H. Norris 101004 CAVA BR 32 Bartramia stricta USA 100 20. Feb 2006 D. H. Norris 108765 CAVA BR 33 Bartramia stricta USA 400 ft 29. Jan 2000 J. R. Shevock 18884 CAVA BR 34 Bartramia stricta USA 150 16. Feb 2003 D. H. Norris 105288 CAVA BR 35 Bartramia stricta USA 100 20. Feb 2006 D. H. Norris 108748 CAVA BR 36 Bartramia stricta Cyprus 1180 20. Feb 2009 J. P. Frahm 2009683 BONN BR 37 Bartramia stricta Spain 770 8. April 2001 D. Quandt X10285 BONN BR 38 Bartramia stricta Turkey 50 01. April 1971 E. Nyholm 442 NMW & A. C. Crundwell BR 39 Bartramia stricta Guernsey 17. Mar 1953 A. C. Crundwell 43 NMW BR 40 Bartramia stricta Cyprus 25. Mar 1982 E. C. Wallace s.n. NMW ...... continued on the next page

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APPENDIX 2. (Continued) Scientific name on the Elevation Collection Nr. Country Date Collectors Herbarium voucher (masl) or Field Nr. BR 41 Bartramia stricta Saudi 1450–1600 22. Nov 1981 W. Frey & H. 1564 B Arabia Kürschner BR 42 Bartramia stricta Tenerife 500 22. Feb 1976 H. Kürschner 940 B BR 43 Bartramia stricta Morocco 27. Feb 1998 F. Müller 1998 DR BR 44 Bartramia stricta Morocco 1100 17. Mar 1997 M. J Cano, M. T. 10516 MUB Gallego, R. M. Ros BR 45 Bartramia stricta Canada 02. May 1979 W. B. Schofield 71733 UBC BR 46 Bartramia stricta USA Apr. 1898 M. S. Baker 727 CAVA BR 47 Anacolia laevisphaera USA 19. Feb 1941 I. M. Haring 423 MO BR 48 Bartramia stricta Tuscany Feb 1963 Schw. 430 BONN BR 49 Bartramia stricta Italy July 2002 L. Gartung X10267 BONN BR 50 Bartramia stricta Canary Isl. 600 1990 C. Illata 1493 BONN BS 51 Anacolia laevisphaera Chile 130 26 Jan 1988 M. Mahú 22316 MO BS 52 Anacolia laevisphaera Chile 1450 23 Oct 1985 S. Tapia 21135 MO BS 53 Anacolia laevisphaera Chile 340 17 Sept 1987 M. Mahú 22127 MO BS 54 Anacolia laevisphaera Chile 670 02. Apr 88 M. Mahú 22618 MO BS 55 Bartramia stricta Chile 560 09. Oct 2001 R. R. Ireland 32436 NY & G. Bellolio BS 56 Bartramia stricta Chile 90 28. Oct 2001 R. R. Ireland 32942 NY & G. Bellolio BS 57 Bartramia stricta Chile 350 05. Nov 2001 R. R. Ireland 33226 NY & G. Bellolio BS 58 Bartramia stricta Chile 130 12. Nov 2001 R. R. Ireland 33678 NY & G. Bellolio BS 59 Bartramia stricta Chile 85 20. Sep 2001 R. R. Ireland 31702 NY & G. Bellolio BS 60 Anacolia laevisphaera Chile 300 20. Mar 2000 B. Goffinet 7088 H BS 61 Anacolia laevisphaera Chile, JFI 02. Feb 1955 C. Skottsberg 164 H BS 62 Anacolia laevisphaera Chile, JFI 30. Nov 65 R. Hatcher & J. 239 H Engel BS 63 Bartramia stricta Chile 220 20. Mar 1999 F. Müller 818 H BS 64 Bartramia stricta Argentina 01. July 1984 A. Bonavia 2269/84 H BS 65 Bartramia stricta Chile 750–900 21. Jan 1993 C. M Taylor 11022 H & R. E Gereau BS 66 Bartramia stricta Chile 150–220 27. Nov 1981 H. Deguchi 25686 KOCH BS 67 Bartramia stricta Chile 230 14. Dec 1981 H. Deguchi 26413 KOCH BS 68 Bartramia stricta Chile, JFI 29. Nov 1965 R. Hatcher 726 NY & J. Engel BS 69 Anacolia laevisphaera Brazil 24. Oct 1987 V. Gelain et al. 3394 NY BS 70 Anacolia laevisphaera Chile 1400 17. Dec 1968 M. Mahu 4827 NY BS 71 Anacolia laevisphaera Chile 20 17. Feb 1978 M. Mahú 24054 MO & H. Mahú BS 72 Anacolia laevisphaera Chile 1170 23. Jul 87 M. Mahú 22081 MO BS 73 Bartramia stricta Chile 750–900 21. Jan 93 C. M. Taylor 11022 MO & R. E. Gereau BS 74 Bartramia stricta Chile 1650 01. Dec 01 W. R. Buck 41544 NY BS 75 Bartramia stricta Chile 19 17. Nov 01 R. R. Ireland 33871 NY & G. Bellolio

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APPENDIX 3. Character states of morphological characters scored. No Characters Description of character states 1 Shape of leaf base 0 = all triangular, 1 = most leaves oblong 2 Leaf tip orientation 0 = straight, 1 = curved 3 Leaf base margin curvature 0 = all flat, 1 = most recurved 4 Shape of distal marginal cells 0 = linear, 1 = oblong 5 Maximum number of cells between teeth (number) 6 Marginal teeth orientation 0 = whole to 1/2 of cell protruding, 1 = only cell tip (1/3 or less of the cell length) protruding 7 Shape of marginal teeth cells 0 = nearly isodiametric to short rectangular, 1 = linear, long-rectangular 8 Maximum plant height Measured from basal stem attached to the soil to apex (mm) 9 Plant density (tufts) 0 = loose, 1 = dense 10 Leaf orientation in dry state 0 = erect to erect-spreading, 1 = appressed 11 Leaf orientation in wet state 0 = erect-spreading, 1 = spreading 12 Stem diameter (µm) 13 Central cylinder diameter (µm) 14 Central cylinder/stem proportion Central cylinder width divided by stem diameter 15 Thickness of cortex cells walls (µm)

APPENDIX 4. Matrix data of 15 morphological characters of the 75 taxa included in the analysis. (1) Shape of leaf base; (2) Leaf tip orientation; (3) Leaf base margin curvature; (4) Shape of distal marginal cells; (5) Maximum number of cells between teeth; (6) Marginal teeth orientation; (7) Shape of marginal teeth cells; (8) Maximum plant height; (9) Plant density (tufts); (10) Leaf orientation in dry state; (11) Leaf orientation in wet state; (12) Stem diameter (µm); (13) Central cylinder diameter (µm); (14) Central cylinder/stem proportion; (15) Thickness of cortex cells walls (µm). Codes for final identifications: BL = Bartramia laevisphaera; BR = Bartramia rosamrosiae; BS = Bartramia stricta. Taxon Nr (1) (2) (3) (4) (5) (6) (7) (8) (9) (10) (11) (12) (13) (14) (15)

BL 1 1 ? 1 1 4 0 0 50 ? 0 1 390.3 83.3 0.2134 2.0 BL 2 1 1 ? 1 6 0 0 26 0 0 1 355.1 50.1 0.1411 1.9 BL 3 1 1 ? 1 6 0 0 50 1 ? 1 453.5 77.1 0.1700 2.2 BL 4 1 1 ? 1 6 0 0 65 0 0 1 331.5 33.7 0.1017 3.2 BL 5 1 ? ? 1 6 0 0 28 0 0 ? 258.6 51.5 0.1991 2.0 BL 6 1 ? ? 1 4 ? 0 61 0 1 1 401.1 54.7 0.1364 4.0 BL 7 1 1 ? 1 5 0 0 75 0 0 1 403.4 60.6 0.1502 2.0 BL 8 1 1 1 ? 5 ? 0 20 1 0 1 362.2 64.4 0.1778 2.2 BL 9 1 1 ? 1 5 0 0 65 1 0 1 256.1 46.6 0.1820 2.0 BL 10 1 1 ? 1 5 0 0 75 1 0 1 415.4 61.5 0.1481 2.2 BL 11 1 ? 1 1 4 0 0 25 1 0 1 280.2 61.0 0.2177 2.1 BL 12 1 1 ? 1 3 0 0 33 1 0 1 397.1 95.6 0.2407 2.3 BL 13 1 1 ? 1 5 0 0 70 0 ? 1 435.1 68.4 0.1572 2.7 BL 14 1 1 ? 1 5 0 0 55 1 0 1 384.1 70.7 0.1841 2.4 BL 15 1 1 ? 1 7 0 0 55 0 0 1 346.5 50.0 0.1443 2.3 BL 16 1 ? ? 1 5 0 0 60 1 1 ? 374.4 86.2 0.2302 3.0 BL 17 1 ? ? 1 5 0 0 30 1 0 ? 307.1 75.4 0.2455 2.4 BL 18 1 ? ? 1 5 0 0 35 1 0 0 314.8 56.2 0.1785 2.7 BL 19 ? 1 ? ? 5 0 0 20 1 1 1 251.1 47.6 0.1896 2.2 BL 20 1 1 ? 1 6 ? 0 30 0 0 1 266.0 37.1 0.1395 1.9 ...... continued on the next page

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APPENDIX 4. (Continued) Taxon Nr (1) (2) (3) (4) (5) (6) (7) (8) (9) (10) (11) (12) (13) (14) (15)

BL 211?1130025100 308.963.20.20462.8 BL 221?1160015001 322.372.60.22532.0 BL 23 1 1 1 ? 5 0 0 29 0 0 1 361.1 79.8 0.2210 2.6 BL 241?114004510? 299.459.80.19972.3 BL 251?1160022001 369.370.00.18952.4 BR 1?00021125100 396.196.90.24461.2 BR 2 ? 0 0 ? 3 1 1 17 0 0 0 301.7 70.5 0.2337 0.8 BR 3?00021117110 296.164.50.21781.4 BR 4?00021122110 332.995.20.28601.2 BR 5?00021130110 390.1108.70.27861.0 BR 6?00021112110 150.134.80.23181.1 BR 7?00021120110 190.657.80.30330.7 BR 8?00021140110 312.771.90.22991.1 BR 9?00021120110 315.880.70.25551.0 BR 10?00021115110 331.9113.90.34321.0 BR 11?00021125110 232.053.40.23020.9 BR 12?00021123110 278.880.80.28981.0 BR 13 ? 0 0 ? 2 1 1 12 0 1 0 255.8 63.2 0.2471 0.8 BR 14?00021125110 326.084.10.25801.0 BR 15?00031120110 359.599.40.27651.1 BR 16?00031115110 409.4158.70.38760.8 BR 17?00021115110 298.292.50.31021.1 BR 18?00021120110 356.7116.00.32521.0 BR 19?00031110110 348.397.30.27941.1 BR 20?00021130110 293.962.30.21200.9 BR 21?00021120110 322.370.90.22001.3 BR 22?00021115110 312.382.20.26321.0 BR 23?00031120110 394.0126.40.32081.0 BR 24?00031113110 334.993.90.28040.9 BR 25?00031110010 189.843.00.22660.6 BS 1 1 ? ? 1 6 0 0 45 1 1 0 335.3 68.0 0.2028 1.0 BS 2 ? ? ? 1 5 0 0 15 1 1 0 326.3 99.3 0.3043 0.9 BS 3 1 ? ? 1 5 ? 0 50 1 1 0 435.7 108.2 0.2483 1.1 BS 4 ? 1 ? 1 6 1 0 24 0 1 ? 347.6 109.5 0.3150 1.1 BS 5 1 1 ? 1 3 0 0 30 0 1 ? 415.6 124.5 0.2996 1.2 BS 6 1 1 ? 1 6 0 0 45 1 0 ? 363.7 90.2 0.2480 0.9 BS 7 1 1 ? 1 5 0 0 35 1 0 0 382.2 111.3 0.2912 1.3 BS 8 ? ? ? 1 5 ? 0 30 1 0 0 361.9 115.0 0.3178 0.9 BS 9 ? ? ? 1 5 0 0 32 1 0 ? 329.7 97.8 0.2966 0.9 BS 1011?150040001 419.5130.10.31011.1 BS 11 1 ? ? 1 4 0 0 35 1 1 ? 329.9 112.7 0.3416 1.0 BS 12 1 ? ? 1 5 0 0 20 1 1 ? 317.5 83.2 0.2620 1.0 ...... continued on the next page

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APPENDIX 4. (Continued) Taxon Nr (1) (2) (3) (4) (5) (6) (7) (8) (9) (10) (11) (12) (13) (14) (15)

BS 1311?17?030001 395.1114.20.28900.9 BS 14 1 ? ? ? 6 0 0 25 1 0 ? 294.7 89.6 0.3040 1.0 BS 151?115?055101 320.983.40.25991.1 BS 16 1 1 ? ? 4 ? 0 35 1 0 ? 312.8 81.4 0.2602 0.8 BS 17 ? 0 ? ? 4 ? 0 40 1 1 0 378.8 145.6 0.3844 1.2 BS 181??140027111 346.1109.20.31551.2 BS 1911?16?015101 397.0139.30.35091.2 BS 2011?150046101 415.7112.20.26990.9 BS 21 1 ? ? 1 5 0 0 25 1 1 ? 443.1 162.5 0.3667 1.3 BS 2211?13?017001 374.0109.50.29280.8 BS 2311?150075101 422.9115.60.27341.0 BS 24 ? 1 ? 1 5 ? 0 10 1 0 1 259.3 86.6 0.3340 0.9 BS 25 1 ? ? 1 5 0 0 25 1 1 1 337.9 81.6 0.2415 1.3

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