Global phylogeny and biogeography of grammitid ferns (Polypodiaceae)

Accepted Manuscript Global phylogeny and biogeography of grammitid ferns (Polypodiaceae) Michael A. Sundue, Barbara S. Parris, Tom A. Ranker, Alan R. Smith, Erin L. Fujimoto, Delia Zamora-Crosby, Clifford W. Morden, Wen-Liang Chiou, Cheng-Wei Chen, Germinal Rouhan, Regina Y. Hirai, Jefferson Prado PII: DOI: Reference:

S1055-7903(14)00284-X http://dx.doi.org/10.1016/j.ympev.2014.08.017 YMPEV 5001

To appear in:

Molecular Phylogenetics and Evolution

Received Date: Revised Date: Accepted Date:

23 April 2014 13 August 2014 15 August 2014

Please cite this article as: Sundue, M.A., Parris, B.S., Ranker, T.A., Smith, A.R., Fujimoto, E.L., Zamora-Crosby, D., Morden, C.W., Chiou, W-L., Chen, C-W., Rouhan, G., Hirai, R.Y., Prado, J., Global phylogeny and biogeography of grammitid ferns (Polypodiaceae), Molecular Phylogenetics and Evolution (2014), doi: http:// dx.doi.org/10.1016/j.ympev.2014.08.017

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Global phylogeny and biogeography of grammitid ferns (Polypodiaceae) Michael A. Sunduea,c,*, Barbara S. Parrisb, Tom A. Rankerc, Alan R. Smithd, Erin L. Fujimotoc, Delia Zamora-Crosbya, Clifford W. Mordenc, Wen-Liang Chioue, Cheng-Wei Chenf, Germinal Rouhang, Regina Y. Hirai,h and Jefferson Pradoh.

a

The Pringle Herbarium, Department of Plant Biology, The University of Vermont, 27 Colchester Ave., Burlington, Vermont 05405 U.S.A. b Fern Research Foundation, 21 James Kemp Place, Kerikeri, Bay of Islands, New Zealand 0230 c Department of Botany, University of Hawaii, 3190 Maile Way, Honolulu, Hawaii 96822 U.S.A. d University Herbarium, 1001 Valley Life Sciences Bldg. # 2465, University of California, Berkeley, California 94720-2465 U. S. A. e Division of Botanical Garden, Taiwan Forestry Research Institute, 53 Nan-Hai Rd., Taipei 100, Taiwan f Institute of Molecular and Cellular Biology, National Tsing Hua University, Hsinchu 30013, Taiwan. g Muséum national d’Histoire naturelle, UMR CNRS 7205 ‘Origine, Structure et Evolution de la Biodiversité’, Botanique, 16 rue Buffon CP 39, 75005 Paris, France h Instituto de Botânica, Caixa Postal 68041, CEP 04045-972, São Paulo-SP, Brazil

*

Corresponding author: Phone 1-646-247-8992. E-mail address: [email protected] (Michael Sundue)

ABSTRACT We examined the global historical biogeography of grammitid ferns (Polypodiaceae) within a phylogenetic context. We inferred phylogenetic relationships of 190 species representing 31 of the 33 currently recognized genera of grammitid ferns by analyzing DNA sequence variation of five plastid DNA regions. We estimated the ages of cladogenetic events on an inferred phylogeny using secondary fossil calibration points. Historical biogeographical patterns were inferred via ancestral area reconstruction. Our results supported four large-scale phylogenetic and biogeographic patterns: 1) a monophyletic grammitid clade that arose among Neotropical polypod ancestors about 31.4 Ma; 2) a paraphyletic assemblage of clades distributed in the Neotropics and the Afro-Malagasy region; 3) a large clade distributed throughout the AsiaMalesia-Pacific region that originated about 23.4 Ma; and, 4) an Australian or New Zealand origin of the circumaustral genus Notogrammitis. Most genera were supported as monophyletic except for Grammitis, Oreogrammitis, Radiogrammitis, and Zygophlebia. Grammitid ferns are a well-supported monophyletic group with two biogeographically distinct lineages: a primarily Neotropical grade exhibiting several independent successful colonizations to the Afro-Malagasy region and a primarily Paleotropical clade exhibiting multiple independent dispersals to remote Pacific islands and temperate, austral regions.

1. Introduction With close to 1500 species, the cosmopolitan Polypodiaceae are among the largest families of ferns (Smith et al., 2006). Most species are epiphytes and represent the fourth largest family of epiphytic vascular plants (Gentry and Dodson, 1987). The family’s prominence was established during a Cenozoic radiation in which leptosporangiate ferns diversified into new niches in otherwise angiosperm-dominated forests (Schuettpelz and Pryer, 2009). The family includes a large monophyletic clade referred to as the “grammitids”, which were often treated as a separate family (Grammitidaceae) prior to molecular phylogenetic evidence (Schneider et al., 2004). With ca. 900 species (Perrie and Parris, 2012) treated in 33 genera, the grammitids comprise close to two-thirds of the diversity in the family; the nongrammitid Polypodiaceae (referred to here as the polypods) include some 450 species treated in 40 genera (Smith et al., 2006). Whereas the polypods are characterized by often round exindusiate sori, reniform, monolete spores usually lacking chlorophyll at maturity, usually scaly leaves, and dorsiventral rhizomes with well-developed phyllopodia, synapomorphies for the grammitid clade include a reduction in the number of cells in the middle of the sporangial stalk from three to one (Wilson, 1959), a reduction in the number of vascular bundles in the petiole from several (like most eupolypod I ferns) to a single bundle or two that fuse above the base (Parris, 1990; Sundue, 2010a), and globose, trilete spores that contain fully developed chlorophyll at maturity (Parris, 1990; Sundue, 2010a). Although both groups often have minute branched hairs (c. 0.1 mm long), polypods most often have broad basifixed or peltate scales whereas grammitids do not have these scales but have pluricellular uniseriate setae (Sundue et al., 2010). Occasional intermediates between scales and setae suggest that these two types of indument may in fact be homologous (Sundue, unpublished). Perhaps owing to their relatively larger size, ease of cultivation, and conspicuous adaptations (e.g., myrmecophily, detritus catching leaves, desiccation tolerance), it is the polypods, as defined above, that have received most attention in the literature. The grammitids, which are generally small, inconspicuous, and not amenable to cultivation, are less well known. This has begun to change recently, with grammitids becoming the focus of many systematic studies. Polypodiaceae have undergone massive generic re-circumscription following molecular phylogenetic studies (e.g., Ranker et al., 2004; Schneider et al., 2004; Kreier et al., 2008; Wang et al., 2010), which demonstrated that many genera were not monophyletic. Within the grammitids, the classical genera Grammitis, Xiphopteris, and Ctenopteris were largely based on blade dissection, a character that has proven nearly useless in defining monophyletic clades. More recently described genera such as Terpsichore A. R. Sm. and Lellingeria A. R. Sm. & R. C. Moran, have relied instead on suites of morphological characters including microscopic and anatomical features (Moran and Smith, 1991; Smith, 1993). These characters have more reliably defined monophyletic groups, but are still subject to cases of morphological homoplasy resulting in polyphyletic genera (Ranker et al., 2004; Labiak et al. 2010b). Consequently, the polyphyly of Lellingeria R. C. Moran & A. R. Sm. (Labiak et al., 2010b) and Terpsichore A. R. Sm. (Ranker et al., 2004; Sundue et al., 2010) has led to the segregation of Alansmia M. Kessler et al. (Kessler et al., 2011), Ascogrammitis Sundue (Sundue, 2010b), Galactodenia Sundue & Labiak (Sundue et al., 2012), Leucotrichum Labiak (Labiak et al., 2010a), Moranopteris R.Y. Hirai & J. Prado (Hirai et al., 2011), Mycopteris Sundue (Sundue, 2013), and Stenogrammitis Labiak (Labiak, 2011). Similarly, our understanding of Paleotropical grammitids has changed radically. Two names that had been widely applied are now recognized as synonyms of other genera:

Ctenopteris Blume ex Kunze = Prosaptia C. Presl (Price, 1982, 1987), and Xiphopteris Kaulf. = Cochlidium Kaulf. (Bishop, 1978). Moreover, an understanding that traditional generic boundaries were not well defined (Parris 1977, 1984), and the determinations of polyphyly of such large historically recognized genera as Grammitis Sw. (in the strict sense including only those species with a distinct black, sclerified leaf margin, sensu Parris, 2007), Ctenopteris, and Xiphopteris, which were defined mainly on the basis of blade dissection, led to the recent coining of Archigrammitis Parris (Parris, 2013), Chrysogrammitis Parris (Parris 1998a), Dasygrammitis Parris (Parris, 2007), Notogrammitis Parris (Perrie and Parris, 2012), Radiogrammitis Parris (Parris, 2007), Themelium (T. Moore) Parris (Parris, 1997), and Tomophyllum (E. Fourn.) Parris (Parris, 2007), and the reinstatement of Oreogrammitis Copel. (Parris, 2007). With the exception of Notogrammitis, which Perrie and Parris (2012) distinguished from Grammitis s.l. using plastid DNA markers, the circumscription of Paleotropical genera has not had the benefit of densely sampled molecular phylogenetic studies. Consequently, phylogenetic relationships among Paleotropical grammitids remain largely uncertain. Grammitids are found most abundantly in montane forests (Parris, 2005; Parris et al., 1992). At higher elevations they can be an important component of the fern flora, for example, on Mt. Kinabalu (Sabah, Malaysia) grammitid ferns constitute 25–35% of total fern diversity between 2000–4000m (Kessler et al., 2001). Likewise, Notogrammitis crassior occurs up to 2600 m in New Zealand (Parris and Given 1976), higher than any other fern species there. Among vascular epiphytes, the two species thought to grow at highest recorded elevations both belong to Melpomene (Sylvester et al. 2014). The majority of grammitid diversity is confined to the tropics, although some groups extend to the temperate zones, reaching 40°N and 56°S (Parris, 2003). Parris (2003) assigned grammitid distributions to two major phytogeographic zones: 1) Neotropics-Africa-Madagascar (including also the Mascarenes, Seychelles, and Comores) where there are about 300 species, and 2) Asia-Malesia-Pacific with about 450 species. Current species counts are c. 400 species for Neotropics-Africa-Madagascar, and c. 500 for Asia-Malesia-Pacific (Parris, unpubl.), reflecting the progress in grammitid systematics in the last decade. Dispersal within each zone appears to be common; all but one of the genera present in Africa-Madagascar also occur in the Neotropics, and at least three species are found in both places (Moran and Smith, 2001). Likewise, the Asian, Malesian, and Pacific areas contain overlapping diversity. Phylogenetic analyses further corroborate these two phytogeographic zones, with both Ranker et al. (2004) and Sundue et al. (2010) finding evidence that most species from the Asia, Malesia, and the Pacific belong to a single clade. Migration between these two zones is rare. As currently understood, only four genera cross these phytogeographic zones: 1) Stenogrammitis, a primarily Neotropical genus that reaches Africa-Madagascar, the Hawaiian Islands, and a few southPacific islands (Labiak et al., 2011; Ranker et al., 2010); 2) Ctenopterella, a mainly Malesian Pacific genus that extends to Africa (Parris, 2007); 3) Notogrammitis, an austral genus most diverse in Australia and New Zealand that reaches South Africa and southern South America, and 4) Grammitis s.s., ranging from the western Pacific through the Neotropics to Africa, Madagascar, and the Mascarenes. Notably the Hawaiian Islands represent an area of overlap, appearing to have been colonized via migrations from both phytogeographic zones (Geiger et al., 2007). The historical biogeography of ferns includes examples of both vicariance and longdistance dispersal, and discerning between these has been a goal of biogeographic research (Barrington, 1993; Wolf et al., 2001; Korall and Pryer, 2014; Labiak et al., 2014). Copeland

(1939) hypothesized that Grammitis s.l. (including all species with simple and entire leaves) were ancestrally Antarctic in distribution, attaining their present distribution via migration through New Zealand, South Africa and Madagascar, and the southern Andes. While this austral source for the origin of grammitids cannot be ruled out, migration via all three routes conflicts with current inferences. Based on the results of Schuettpelz and Pryer (2009), it appears that the Polypodiaceae diverged from Paleotropical ancestors ~55.8 Ma, and migrated into the Neotropics ~43 Ma. Grammitids appear to have evolved from these Neotropical polypod ancestors ~30.6 Ma. Subsequent migration to Asia/Africa occurred later, and must have operated via long-distance dispersal, at least at the intercontinental level, because the continents have not moved appreciably since the clade evolved. Long-distance dispersal of spores by wind was also invoked by Moran and Smith (2001) to explain floristic similarities between the Neotropics and Africa/Madagascar. The timing and patterns of these dispersals remain largely unknown, but putatively, it would seem that they are quite recent; some of the same species occur in both areas, and there is very little diversification in several of the genera that migrated to Africa, namely Melpomene A. R. Sm. & R. C. Moran, Alansmia, Cochlidium, Leucotrichum, and Ceradenia L. E. Bishop. The pattern is similar in some other genera, such as Zygophlebia L. E. Bishop, Enterosora Baker, Stenogrammitis, and Grammitis s.s., but in these genera more diversification has occurred in Africa and Madagascar. The origin of the Asian-Malesian-Pacific grammitids; however, is particularly unclear. With these issues in mind, we undertook a phylogenetic study of grammitid ferns aiming establishing a global picture of phylogeny and historical biogeography. Our sampling included 50% of Paleotropical species and 31 of the 33 currently recognized genera (Archigrammitis Parris and Luisma M. T. Murillo & A. R. Sm. remain unsampled). We addressed two primary questions: Are the Neotropical and Afro-Malagasy floras closely related as hypothesized by Parris (2003)? Do the Asian-Malesian-Pacific species constitute a single clade, or is this Old World diversity the result of multiple migrations? 2. Materials and Methods 2.1. DNA extraction and amplification Ingroup sampling included 190 species (202 accessions) from 31 of the 33 currently recognized genera; samples of Archigrammitis and Luisma were unavailable. Sampling included the type species for 20 genera (Table 1). Outgroups included five species of Neotropical polypods found to be closely related in previous studies (e.g., Schuettpelz and Pryer, 2009). This choice was further informed by our unpublished analysis of available sequences in GenBank that also find these Neotropical polypods as the closest relative of the grammitid clade. Sequences not generated by us were downloaded from GenBank. DNA extraction and PCR amplification protocols followed those of Labiak et al. (2010b). We PCR-amplified five plastid DNA markers: the atpß and rbcL coding regions, and the trnL-trnF, rps4-trnS, and trnG-trnR intergenic spacers, and generated 67, 71, 106, 92, and 71 sequences of each, respectively. DNA sequencing was performed at the Greenwood Molecular Biology Facility at the University of Hawai‘i at Mānoa, and all sequences were submitted to GenBank (Appendix 1).

2.2. Alignment and analyses Sequences were edited and contigs were produced using Geneious 6.17 (Biomatters Ltd., San Francisco, CA), and the MAAFT plug-in was used to produce alignments (Katoh, 2013). Alignments were visually inspected and no areas appeared to be aligned ambiguously. For each aligned marker, optimal data partitioning and models of substitution evolution were estimated using AICc in PartitionFinder (Lanfear et al., 2012). We provided PartitionFinder with subsets for each marker, and for the three coding regions we provided subsets for each nucleotide position. The resulting best scheme was a single GTR+I+G model for the dataset. This was implemented in the Bayesian and likelihood tree searches. We conducted tree searches using maximum parsimony (MP), maximum likelihood (ML) and Bayesian (BI) analyses. Maximum parsimony tree searches were performed using TNT (Goloboff et al., 2008) employing two approaches. First, we conducted traditional heuristic searches with 1000 parsimony ratchet replicates (Nixon, 1999) (200 iteration ratchet, the up and down weights set to 5% each), holding 20 trees per ratchet, followed by tree-bisection-reconnection (TBR)-max branch swapping. We then implemented a New Tech search strategy set at level 15 implementing tree fusing, sectorial search, and the parsimony ratchet finding the minimum length tree 10 times. In each case, support for nodes was calculated by bootstrap analyses (BS), with 1000 replicates using the same methods. Maximum likelihood tree searches were conducted using RAxML (Stamatakis, 2006) through the CIPRES portal (Miller et al., 2010). Five independent searches for the ‘best tree’ and 10,000 BS replicates were generated implementing the best partition scheme determined by PartitionFinder. Bayesian tree searches were conducted using MrBayes (Huelsenbeck and Ronquist, 2001; Ronquist and Huelsenbeck, 2003) through the Oslo Lifeportal (https://lifeportal.uio.no/root). We conducted five runs implementing the best partition scheme determined by PartitionFinder for 10 million generations. Each run included four chains (one cold, three heated) with unlinked parameters, and chain temperature set to 0.2. Priors were uniform except that rates were allowed to vary among loci (ratepr = variable). The posterior was sampled every 1000 generations, and the first 25% discarded as “burn-in”. Convergence was estimated by examining the standard deviation of split frequencies, plotting the output parameters in TRACER v 1.5 (Rambaut et al., 2013), and examining tree files in AWTY (Wilgenbush et al., 2004; Nylander et al., 2008) 2.3. Fossil Calibration Known fossil calibration points for the Polypodiaceae are limited. Consequently, we chose to calibrate our chronogram using node ages generated by a fossil-calibrated analysis of all leptosporangiate ferns (Schuettpelz and Pryer, 2009). Grammitis succinea L. D. Gómez, a fossil in Dominican amber, is the only known fossil that appears to belong to the grammitids (Gómez, 1982). While purported to have several synapomorphies for the grammitids (namely, stiff, erect uniseriate, pluricellular setae and uniseriate sporangial stalks), this fossil is relatively young (approximately 25 Ma) and cannot be placed with confidence in any clade within grammitids; consequently, it was not useful to our study. We estimated divergence times using a relaxed molecular clock as implemented in BEAST, using a Markov chain Monte Carlo strategy (Drummond et al., 2006; Drummond and Rambaut, 2007; Drummond et al., 2013). We partitioned the dataset by plastid DNA region, and specified the optimal model for each region as determined by PartitionFinder. We implemented a

Yule speciation tree prior and an uncorrelated lognormal model of rate change, with clock models unlinked between partitions and a GTR+G substitution model in all cases. We used the following calibration points from Schuettpelz and Pryer (2009) with normal distribution priors: the most recent common ancestor (MRCA) of the grammitid clade (31.2 Ma, 3.12 s.d.), the MRCA of the large Neotropical clade including Mycopteris–Stenogrammitis (23.3 Ma, 2.33 s.d.), the large primarily Paleotropical clade including Moranopteris–Oreogrammitis (23.4 Ma, 2.34 s.d.), and the MRCA of Serpocaulon A. R. Sm. (15.5 Ma, 1.55 s.d.). The clade corresponding to each calibration point was constrained to be monophyletic. Three analyses were run, each for 30,000,000 generations, with parameters sampled every 1000 generations. The program LogCombiner was used to pool the resulting files. Tracer v1.5 (Drummond and Rambaut, 2007) was used to examine the posterior distribution of all parameters and their associated statistics including estimated sample sizes (ESS) and 95% highest posterior density (HPD) intervals. The program TreeAnnotator v2.0.2 (Drummond and Rambaut, 2007) was used to summarize the post burn-in trees and produce a maximum clade credibility chronogram showing mean divergence time estimates with 95% HPD intervals. Historical Biogeography—Ancestral area reconstruction (AAR) was conducted using the dispersal-cladogenesis (DEC) model as implemented in the program Lagrange (Ree and Smith, 2008). We defined the following geographic areas: A) Neotropical, B) Africa–Madagascar and islands of the Atlantic (Ascension and St. Helena) and Indian Oceans (Comores, Réunion, Mauritius, Seychelles), C) tropical Asia and the Pacific including Bougainville, Solomon Islands and Vanuatu, as well as NE Australia and New Caledonia and Micronesia (Palau, Pohnpei, Kosrae), D) a temperate circumaustral region south of 28°S, E) Hawaiian Islands and the central and eastern South Pacific, and F) Nearctic. Distributions were assigned to each species based upon known records of herbarium specimens, and species with broad ranges were scored as polymorphic. These occurred in Alansmia, Cochlidium, and Stenogrammitis. Given the recent age of grammitid ferns, and vagility of their spores, we made no restrictions in the adjacency matrix. Dispersal constraints (Q matrix) between these regions were scored as uncommon (0.25) or very uncommon (0.01) and are presented in Table 2; maximum range size was set at 2.

3. Results 3.1. Phylogenetic analyses The final aligned dataset included 5,569 sites of which 1974 (35%) were parsimony informative (Table 3). During Bayesian analyses, runs converged after the first 1 million generations, ESS values of each parameter were all well above the recommended threshold of 200, and the traces of corresponding parameters in independent runs converged to the same optimum. Maximum parsimony tree searches with TNT found shortest trees of 10,761 steps. In a strict consensus, these MP trees retained all backbone relationships, except for one large polytomy forming in the crown group of Oreogrammitis, Radiogrammitis, and Themelium. The overall topology of the MP trees was very similar to that resulting from the ML and Bayesian analyses. One minor difference was that Grammitis s.s. was recovered as monophyletic, but without strong branch support. Best ML trees resulting from RAxML shared an overall topology identical to that of the

Bayesian trees and therefore are not discussed in further detail. Results from the Bayesian analyses were generally well resolved and well supported (Fig. 1). Results from our analyses supported grammitids as monophyletic (Fig. 1). The overall topology can be explained in general as a monophyletic tropical Asian clade nested within a primarily Neotropical and African grade (Fig. 2). Our results support the Neotropical genus Moranopteris as sister to this tropical Asian clade, and the tropical Asian genus Chrysogrammitis, which has been difficult to resolve in previous analyses, as sister to the remainder of the tropical Asian clade. Relationships within the Neotropical and African grade are generally congruent with previous studies, but with improved resolution. Well-supported relationships not previously reported include: 1) Zygophlebia L. E. Bishop is paraphyletic with Enterosora Baker nested within it; 2) Ceradenia L. E. Bishop is resolved as two clades corresponding to Bishop’s (1988) subgenera Ceradenia and Filicipecten; and 3) Lomaphlebia J. Sm. is sister to the clade comprising Grammitis s.s. and Cochlidium Kaulf. In previous studies (e.g., Sundue et al. 2010), Terpsichore was resolved as sister to all other grammitids, but here it is sister to the clade of Adenophorus Gaudich., Cochlidium, Grammitis s.s., and Lomaphlebia. These five genera instead compose a clade sister to all other grammitids. Paraphyly of Grammitis s.s. with respect to Cochlidium is also a novel result supported by our Bayesian and likelihood results. The remaining tropical Asian genera resolve in two main clades, one comprising Calymmodon C. Presl, Dasygrammitis, Micropolypodium Hayata, Scleroglossum Alderw., Tomophyllum, and Xiphopterella, and the other comprising Acrosorus, Ctenopterella, Notogrammitis, Oreogrammitis, Prosaptia, Radiogrammitis, Themelium, and four species currently combined in Grammitis (listed in figures 1 and 2 as “Grammitis”) that are not placed within current generic concepts. Most Paleotropical genera were supported as monophyletic, including Calymmodon, Chrysogrammitis, Dasygrammitis, Micropolypodium, Notogrammitis, Prosaptia, Tomophyllum, Scleroglossum, and Xiphopterella. In contrast, Oreogrammitis and Radiogrammitis are polyphyletic. The three species of Themelium included were resolved as monophyletic, but nested within the large clade comprising Oreogrammitis and Radiogrammitis. Ctenopterella and Acrosorus included only one species each, so the monophyly of those genera remains untested. 3.2. Analyses of Divergence Times and Diversification Rates BEAST analyses estimated the MRCA of the grammitids at 31.4 Ma and the MRCA of the large Paleotropical clade at 23.4 Ma (Fig. 2a, b). While stem ages varied among Paleotropical genera, many of them began to diversify within the last 8.4 Ma, including Acrosorus, Calymmodon, Chrysogrammitis, Dasygrammitis, Micropolypodium, Prosaptia, Scleroglossum, and Xiphopterella. The large predominantly Neotropical clades were estimated to originate from 18–23 Ma, with subsequent diversification occurring as early as 14.5 Ma for Ascogrammitis and as recently as 3.8 Ma in Melpomene. The stem age of the Hawaiian Island endemic Adenophorus was estimated as 22.5 Ma, and subsequent diversification of the genus was estimated at 10.6 Ma. 3.3. Ancestral Area Reconstruction LAGRANGE analyses reconstructed a Neotropical ancestral area for the entire grammitid clade (Fig. 2a). The majority of early diverging lineages are distributed in the Neotropics, and a Neotropical ancestral area was retained throughout the first six backbone nodes of the tree. The

present distribution of grammitids in tropical Asia was explained by a single transition to tropical Asia at 23.4 Ma (25.0–20.7 Ma). Subsequent backbone nodes after this transition were reconstructed as having a tropical Asian ancestral area as well. There was a single transition from within the tropical Asian clade to the austral region at 14.7 Ma (21.0–8.6) by the genus Notogrammitis. Based on our sampling, the present distribution of grammitids in Africa and Madagascar is explained by at least six separate migrations from the Neotropics: 1) the Grammitis cryptophlebia clade 6.8 Ma [11.9–3.0]; 2) Alansmia elastica 4.9 Ma [9.9–2.0]; 3) Melpomene flabelliformis 0.3 Ma [2.3–0.002]; 4) Stenogrammitis oosora 5.59 Ma [8.9–2.6]; and 5) Zygophlebia 12.6 Ma [22.1–7.5]. Migrations of the Grammitis cryptophlebia clade and Zygophlebia differ from the others by involving more than a single species. The former is a single event followed by a small radiation of three species. The Zygophlebia migration involves two species (three accessions) that comprise the first two bifurcations of the ZygophlebiaEnterosora clade. This could be interpreted as a single migration followed by a subsequent migration back to the Neotropics, or as two separate migrations to Africa-Madagascar. The present distribution of grammitids in the Hawaiian Islands was explained by three migrations, two from the Neotropics (Stenogrammitis 1.5 Ma [4.7–0.38]; Adenophorus 22.5 [29.0–14.1]); and one from tropical Asia (Oreogrammitis hookeri + O. forbesiana 2.3 Ma [7.08– 0.36]). 4. Discussion 4.1. Phylogenetic results 4.1.1. Overall results Our sampling included 31 of the 33 currently recognized grammitid genera. We find that 24 of them are monophyletic (Fig. 1); however, these monophyletic genera are nested within other genera in three cases. Ctenopterella (14 spp.) and Acrosorus (11 spp.) are each represented by a single species, and so their monophyly remains untested. Monophyly among the predominantly Neotropical lineages is not surprising as many of these genera were recently circumscribed following results of molecular phylogenetic studies. Our finding that most Paleotropical genera are monophyletic is surprising. Most of these generic concepts were based on morphological characters alone, which had been shown to be prone to homoplasy among Neotropical lineages (Ranker et al. 2004; Sundue 2010; Sundue et al. 2010). 4.1.2. Cochlidium, Grammitis s.s., and Lomaphlebia Circumscription of Grammitis, typified by G. marginella, a Neotropical species not included in our analyses (Bishop, 1977), has been a central problem in the systematics of grammitid ferns. One extreme has been to use it to include all or nearly all of the diversity of the clade (e.g., Tryon and Tryon, 1982; Christenhusz and Chase, 2014). Most recent authors; however, have restricted it to include the ca. 25 species with simple, entire leaves that have black sclerotic margins. The decision to adopt a broad Grammitis comprising the entire grammitid lineage by Tryon and Tryon (1982) was conservative, but justified by their understanding that the other two widely applied names at that time, Ctenopteris and Xiphopteris, were patently

artificial. However, the problems in generic circumscription that Tryon and Tryon (1982) faced have now largely been resolved by subsequent studies. Nonetheless, Christenhusz and Chase (2014) recently advocated treating all grammitid genera at the subgeneric rank within a single genus, Grammitis, and the clade as a tribe Polypodieae within an unwieldy mega-family Polypodiaceae, subfamily Polypodioideae. We view this decision as retrogressive, uninformative to the goals of biological systematics, and contrary to all other recent advances in depiction of the relationships of ferns. It is also inconsistent with their proposed treatment of the remainder of the family. Their classification has the effect of obscuring understanding rather than clarifying it. A more informative classification can be found in Smith et al. (2008). Surprisingly, our results do not support the monophyly of Grammitis s.s., the blackmargined group. Instead, it is paraphyletic, with a Neotropical clade and an Afro-Malagasy clade, and the genus Cochlidium sister to the latter (Fig. 1). Cochlidium differs morphologically from Grammitis s.s. by having veins usually ending in hydathodes, lacking the dark sclerotic margin, and in some cases by having a coenosorus (Bishop, 1978). The coenosorus is often sunken into a thickened lamina, giving Cochlidium a very different appearance from Grammitis s.s., in spite of having a large number of similarities that include radially symmetrical rhizomes, concolorous scales, and the loss of laminar setae. These results, together with the presence of Grammitis cryptophlebia (Baker) Copel., which lacks a black sclerotic lamina margin, as sister to G. melanoloma (Cordemoy) Tardieu, which has a black margin, indicate that the black sclerotic lamina margin, previously considered to be a unique character, may in fact be homoplastic. However, our sampling of Grammitis s.s. is limited, and denser sampling is needed to accurately estimate the evolution of that character. Sister to all of these is Lomaphlebia turquina (Maxon) Sundue and Ranker (Appendix 2), one of two species in the genus endemic to the Caribbean and which has not been included in previous molecular phylogenetic analyses. Its position here is supported by its general morphology in that it is very similar to Grammitis s.s., but lacks the black sclerotic lamina margin and has a submarginal commissural vein. 4.1.3. Ceradenia, Enterosora, and Zygophlebia Our results support a monophyletic Ceradenia as sister to a clade of Zygophlebia with Enterosora nested within the latter (Fig. 1). These largely agree with previous results; however, Sundue et al. (2010) found Zygophlebia nested within Enterosora, or sister to it. Bishop (1988) anticipated the close relationship between Ceradenia and Zygophlebia and subsequent studies have questioned their circumscription (Rakotondrainibe and Deroin, 2006). In contrast, the distinction between Enterosora and Zygophlebia has received less attention. The two genera share a number of characters, and the most prominent character used to separate them, spongiose mesophyll, is homoplastic throughout grammitids (Bishop and Smith, 1992; Sundue, 2010a). These results suggest that maintaining the two as separate genera may not be tenable. Ceradenia is resolved into two clades that correspond with Bishop’s subgenera: subg. Ceradenia and subg. Filicipecten (Fig. 1). Although several of the species included here were described after Bishop’s (1988) classification, the morphological characters he used still apply. Subgenus Ceradenia is characterized by radially symmetrical rhizomes, short or absent petioles, and white wax-like glandular hairs upon the laminar surfaces and paraphyses. Dorsiventral rhizomes, elongate petioles, and white wax-like glandular hairs restricted to paraphyses characterize subg. Filicipecten.

4.1.4. Phylogenetic relationships within the tropical Asian clade A novel relationship revealed here is support of a large clade of species with radially symmetrical rhizomes comprising c. 140 spp. (Parris, unpublished) belonging to six genera: Dasygrammitis, Scleroglossum, Tomophyllum, Micropolypodium, Xiphopterella, and Calymmodon (Fig. 1). Parris (2007) argued that Dasygrammitis was closely allied with Tomophyllum, Scleroglossum, and Calymmodon based in part upon this character, and she indicated that Xiphopterella was also likely to be related. Parris (2007) also noted the morphological similarity between Micropolypodium and Xiphopterella in some characters, including radial rhizome. Radially symmetrical rhizomes occur frequently among grammitids (Sundue et al., 2010), particularly in Alansmia, Leucotrichum, and Radiogrammitis, but the clade of six tropical Asian genera resolved here is by far the largest group of taxa sharing this morphology. Polypods, in contrast, exhibit only dorsiventral rhizomes. Thus, across the family, the change from dorsiventral to radially symmetrical rhizomes is likely to be a derived trait that arose multiple times within the grammitid clade. We speculate that this change is coincident with the reduction in size and with higher niche specialization of grammitids compared to polypods. 4.1.5. Oreogrammitis, Radiogrammitis, and Themelium Copeland (1917) described Oreogrammitis to accommodate a single species from Mt. Kinabalu, Borneo, and cited its confluent sori as unique. Christensen and Holttum (1934) maintained the genus, but noted that its validity was problematical. Parris (1983, 1990) initially sunk the genus into Grammitis because confluent sori occur in other unrelated grammitids, but later resurrected it, making combinations for 107 species when it became apparent that a broadly circumscribed Grammitis was untenable (Parris, 2007). The closely related Radiogrammitis was simultaneously described and distinguished by its radially symmetrical rhizomes that sometimes lack scales. By contrast, the rhizomes of Oreogrammitis are uniformly dorsiventral and scaly. Our results find that Oreogrammitis is polyphyletic and nested in three places within Radiogrammitis (Fig. 1). In some ways, this result is not surprising since the two genera were distinguished using essentially a single character -- rhizome symmetry. It is a bit unexpected; however, because rhizome symmetry effectively circumscribes many genera of grammitid ferns (Ranker et al., 2004; Sundue et al., 2010). The number of changes in rhizome symmetry within this clade suggests that there may be an increased rate of change for this character relative to other grammitid lineages. Further sampling is needed to address that issue, however. There are 153 species of Oreogrammitis and 36 species of Radiogrammitis (Parris, unpubl.), thus the apparently large number of transitions seen in our results may be an artifact of relatively low sampling of both Oreogrammitis and Radiogrammitis. Relationships within Oreogrammitis are further complicated by Themelium (27 species; Parris, unpubl.), which nests within it (Fig. 1). Themelium is similar to Oreogrammitis in its dorsiventral rhizome with glabrous rhizome scales, but differs in having glabrous sporangia and usually subclathrate to clathrate rhizome scales. Radiogrammitis has small leaves that are uniformly simple and entire (or repand) and those of Oreogrammitis are usually also simple and entire (or repand, or even pinnate in a few species). Some species of Themelium also have simple laminae; however, most are pinnate or twice-pinnate, sometimes with rigid sclerified axes and reduced laminar tissue (Parris 1997, 2004, 2010). This transition from simple to compound leaves is surprising, but not unique. Laminar blade dissection among the 10 species of

Adenophorus alone ranges from simple to tripinnate (Ranker et al., 2003), in Ceradenia it ranges from simple to bipinnate (Bishop, 1988), and in Notogrammitis it ranges from simple to bipinnatifid (Perrie and Parris, 2012). These transitions occur outside of the grammitids as well, one example being that of Elaphoglossum sect. Squamipedia Mickel & Atehortúa (Vasco et al., 2013) in comparison to most other Elaphoglossum spp. that have simple, entire leaves. This is among the largest and least well sampled clades of grammitids. With an estimated 216 species, our sampling represents only 10% of the diversity. Our results suggest that generic recircumscription is necessary within this clade, but denser sampling, including the types of all three genera, is warranted before making taxonomic changes. If a single genus is to be retained, Oreogrammitis has nomenclatural priority, but would require alteration of its morphological definition. 4.1.6. Orphan species Notwithstanding the problems involving the circumscription of Oreogrammitis, Radiogrammitis, and Themelium, current generic concepts accommodate all but four species included in our analyses. Three of these species reside in a clade sister to Notogrammitis and could potentially be accommodated within an expanded concept of that genus. That would require, however, alteration of the morphological definition of the genus. Two of these species, Grammitis deplanchei and G. pseudaustralis, are endemic to New Caledonia. The third, G. diminuta, is endemic to Lord Howe Island. Similar phylogenetic relationships were found by Perrie and Parris (2012), who argued to exclude these species from Notogrammitis because they depart morphologically and because they lacked sufficient support values for those nodes to justify their inclusion in that genus. Grammitis stenophylla Parris is the fourth unplaced taxon. In our analyses, it forms a clade with Ctenopterella denticulata and the three accessions of Acrosorus friderici-et-pauli. Grammitis stenophylla is an Australian endemic that was previously thought to be related to species now included in Notogrammitis (Parris, 1998b). Perrie and Parris (2012) concluded that it was unrelated to Notogrammitis, but did not have sufficient sampling to resolve its phylogenetic placement. Its position is well resolved here as sister to Acrosorus and Ctenopterella (Fig. 1), from which it departs strongly morphologically, and which themselves have little in common with each other. Long branches lead to each of these genera, and this may indicate poor sampling in this clade. Acrosorus comprises nine species (Parris, unpubl.) and Ctenopterella has 20 species (Parris, unpubl.). 4.2. Historical biogeography 4.2.1 General patterns Based on the result of our divergence estimates and ancestral area reconstructions, we hypothesize that the grammitid lineage evolved in the Neotropics toward the end of the Eocene (37–44.9 Ma) close to the Oligocene boundary (Fig. 2a), when the Antarctic ice sheet began to expand rapidly. Subsequent lineages remained Neotropical through the end of the Oligocene. Frequent migration to Africa-Madagascar, and infrequent migration to the Hawaiian Islands and eastern Pacific occur among these lineages, beginning in the Miocene. Toward the end of the Oligocene, a single lineage migrated to tropical Asia. Currently, this lineage comprises well over half of grammitid diversity. Like the Neotropical lineages, migration of the tropical Asian

lineage continued during the Miocene, but these migrations were to the Hawaiian Islands and Australasian regions. Further migrations from this lineage did not occur to Africa-Madagascar. Our evidence does not support Copeland’s (1939) hypothesis of an Antarctic migration of Grammitis s.l. to each of the austral continents. Instead, we reconstruct an austral lineage as being derived from tropical Asian ancestors. 4.2.2. Origin of the Afro-Malagasy Grammitid Flora We infer that migrations to Africa and Madagascar occurred at least six times in our analyses, all within the last 14 Ma (–21 Ma) including the genera Alansmia, Cochlidium, Grammitis s.s., Melpomene, Stenogrammitis, and Zygophlebia (Fig. 2a). Considering the results of other studies and of species not sampled here, we expect that the total number of long-distance migrations from the Neotropics to Africa-Madagascar will be considerably higher and will include species of Ceradenia, Enterosora, and Leucotrichum, as well as other species of Grammitis s.s., Stenogrammitis, and Zygophlebia (Ranker et al., 2010; Labiak et al., 2010a, Rouhan et al., 2012). Because of their recent age, these are best explained by long-distance dispersal. Thus, as hypothesized by Moran and Smith (2001), the entire African-Malagasy grammitid flora appears to be the product of recent migration from the Neotropics, based on current sampling. Two of the migrations to Africa and/or Madagascar have led to radiations of species, including Stenogrammitis with 14 species in the Neotropics and 13 species in the African-Malagasy region (Parris, unpublished), and Zygophlebia with seven species in the Neotropics (Bishop, 1989) and nine species in Africa-Madagascar (Bishop, 1989; Parris, 2003; Rakotondrainibe and Deroin, 2006). The other large genera in the region, Ceradenia and Grammitis s.s., are each represented by eight species, respectively (Bishop, 1988; Parris, 2003). There is currently no phylogenetic evidence that species have migrated from tropical Asia to Africa-Madagascar or vice-versa. However, our sampling does not include any of the six species of Ctenopterella from Africa, Madagascar, the Comores or the Mascarenes (Parris, 2007; Parris, 2012; Parris, unpubl.). If Ctenopterella is monophyletic, these species would have been derived from tropical Asian ancestors; however, monophyly of Ctenopterella remains to be tested. Over half of the diversity in Zygophlebia resides in Madagascar, and our study is the first to sample taxa from that region (two spp., three accessions). Their relationships, resolved here as a grade sister to the remainder of the clade, could be interpreted as evidence for migration to Africa-Madagascar followed by a subsequent migration back to the Neotropics (Fig. 2a), a result not otherwise supported by our results. This result could also be derived from two subsequent dispersal events. The latter explanation is consistent with what appears to be the predominant direction of migration via long-distance dispersal of spores from West to East in our results. Africa has also received migrants from the austral region. South Africa is home to Notogrammitis angustifolia, which, along with N. crassior on islands to the SW and SE of Africa, is part of a southern migration out of tropical Asia. Our results, and those of Perrie and Parris (2012), suggest that N. angustifolia and N. crassior evolved from ancestors residing in Australia or New Zealand, and not the Americas. 4.2.3. Origin of the Tropical Asian and Temperate Austral Grammitid Floras The tropical Asian grammitid flora is best explained by a single migration from the Neotropics to tropical Asia, an event estimated to have occurred 24.5 Ma (22.4–27.3 Ma) (Fig.

2b). Because of its recent age, this distribution pattern is likely the result of long-distance dispersal. The historical biogeography of grammitid ferns in tropical Asia is quite different than that in Hawai‘i and Africa-Madagascar, which are both explained by multiple long-distance dispersal events. Tropical Asia is home to the greatest diversity of grammitid species, with c. 490 species (Parris, unpubl.), and a single origin of this diversity is a remarkable finding. Grammitids of tropical Asian origin have migrated to the Hawaiian Islands and other islands of the central and eastern Pacific, but there is no indication that they have migrated to Africa-Madagascar, or back to the Neotropics. One important migration out of tropical Asia is that of Notogrammitis (12 spp.), which dominates the grammitid flora in the southern hemisphere south of 35°30’S (Perrie and Parris, 2012). Notogrammitis is estimated to have entered austral regions 14.7 Ma (21.0–8.6 Ma) (Fig. 2b). Range expansion was likely through Zealandia rather than migration across the Australian plate via the Sahul Shelf, because the sister group to Notogrammitis comprises species currently distributed on New Caledonia and Lord Howe Island. Notogrammitis has also migrated around the southern hemisphere to South Africa and southern South America and oceanic islands, including Tristan da Cunha, Gough, Amsterdam, Marion, Crozet, Kerguelen, Falklands, and South Georgia Islands. It is the only Paleotropical lineage that has reestablished itself in the Americas. Long-distance dispersal best explains this range expansion, which according to our estimates has occurred during the last 1.5 Ma. 4.2.4. Origin of the Hawaiian Grammitid Flora The Hawaiian grammitid flora is also explained by multiple long-distance migrations (Fig. 2a, b). Unlike Africa-Madagascar and the Austral region whose floras are the product of migrations from single regions, the Hawaiian grammitid flora is the product of migrations from two regions, the Neotropics (Adenophorus and Stenogrammitis) and tropical Asia (Oreogrammitis). These results agree with the results of Geiger et al. (2007). Two of these migrations have resulted in local radiations resulting in three species of Oreogrammitis (only two sampled here), and 10 species of Adenophorus. The migration of Oreogrammitis to the Hawaiian Islands is part of a larger pattern of tropical Asian species moving into the central and eastern Pacific to Rarotonga (Radiogrammitis cheesemanii and the Marquesas Islands (O. uapensis). A species from the Moluccas and New Guinea (R. parva) also resides within this clade in our BEAST analysis (Fig. 2), complicating the larger pattern. We interpret our results as evidence for multiple migrations from tropical Asia to the eastern Pacific, rather than a reversal in direction, and this result is supported by our Bayesian analysis that supports O. uapensis further outside of this clade (Fig. 1). More sampling is needed to clarify the pattern. Nonetheless, there is no indication that grammitids have migrated from the Hawaiian Islands to other regions such as the Neotropics. Our estimated stem age of the Adenophorus clade of 22.5 Ma (29.0–14.1) is effectively identical to what Schneider et al. (2005) estimated for the Hawaiian endemic diellia clade of Asplenium L. of 24.3 Ma (27.0–21.5). Both of these ages effectively coincide with the estimate of the renewal of Hawaiian terrestrial life at c. 23 Ma (Clague, 1996; Price and Clague, 2002) after a 10 Ma lull in new island production by the Hawaiian volcanic hotspot. This suggests that these two fern lineages were among the first to colonize successfully the newly emerging islands in the early Miocene. As with the diellia lineage of Asplenium, the Adenophorus lineage is considerably older than any of the current, high Hawaiian Islands (i.e., Kaua‘i is the oldest at c. 5.2 Ma) and extant or ancestral species have arrived at the current islands by dispersing along the

island chain as new islands have been produced. Interestingly, the Adenophorus clade at 22.5 Ma and only 10 species is nearly the same age as the large, species-rich Paleotropical clade at 24.5 Ma and c. 500 species (Parris, unpubl.). Acknowledgments We thank the Curators of the following herbaria for making facilities available for examination of material and/or loans: A, AK, B, BISH, BM, BO, BR, BRI, CANB, CGE, CHR, E, FI, GH, K, KEP, KLU, L, LAE, M, MEL, MO, NHT, NSW, NY, P, PAP, PDA, PE, PTBG, S, SAN, SAR, SING, TAIF, TI, UC, US, VT, WAG, WELT, and Ewen Cameron (AK) for organizing loans to Barbara Parris. We thank the University of Oslo Lifeportal for their computational resources and the Willi Hennig Society for freely providing TNT. This material is based upon work supported by the National Science Foundation under Grant No. DEB-1119695 to T. A. Ranker and C. W. Morden]. Michael Sundue was also supported by the H. M. Burkill Fellowship provided by the Singapore Botanic Garden. References Barrington, D. S., 1993. Ecological and historical factors in fern biogeography. J. Biogeogr. 20, 275–279. Bishop, L. E., 1977. The American species of Grammitis sect. Grammitis. Amer. Fern J. 67, 101–106. Bishop, L. E., 1978. Revision of the genus Cochlidium (Grammitidaceae). Amer. Fern J. 68, 76– 94. Bishop, L. E., 1988. Ceradenia, a new genus of Grammitidaceae. Amer. Fern J. 78, 1–5. Bishop, L. E., 1989. Zygophlebia, a new genus of Grammitidaceae. Amer. Fern J. 79, 103–118. Bishop, L. E., Smith, A. R., 1992. Revision of the fern genus Enterosora (Grammitidaceae) in the New World. Syst. Bot. 17, 345–362. Christenhusz, M. J. M., Chase, M. W., 2014. Trends and concepts in fern classification. Ann. Bot. 1–24, doi: 10.1093/aob/mct299. Christensen, C., Holttum, R. E., 1934. The ferns of Mount Kinabalu. Gardens' Bull. Straits Settlements 7, 191–324. Clague, D. A., 1996. The growth and subsidence of the Hawaiian-Emperor volcanic chain, in: Keast, A., Miller, S. E. (Eds.), The Origin and Evolution of Pacific Island Biotas, New Guinea to eastern Polynesia: Patterns and Processes. SPB Academic Publishing, Amsterdam, pp. 35–50. Copeland, E. B., 1917. New species and a new genus of Borneo ferns, chiefly from the Kinabalu collections of Mrs. Clemens and Mr. Topping. Philipp. J. Sci. 12, 47–65. Copeland, E. B., 1939. Fern evolution in Antarctica. Philipp. J. Sci. 70, 157–188. Copeland, E. B., 1952. [1951]. Grammitis. Philipp. J. Sci. 80, 93–276. Drummond, A. J., Ho, S.Y. W., Phillips, M. J., Rambaut, A., 2006. Relaxed phylogenies and dating with confidence. Plos Biol. 4, E88. Drummond, A. J., and Rambaut, A., 2007. BEAST: Bayesian Evolutionary Analysis by Sampling Trees. BMC Evol. Biol. 7, 214–222. Drummond, A. J., Rambaut, A., Suchard, M., 2013. BEAST V1.8.0 2002-2013. Bayesian Evolutionary Analysis Sampling Trees. Available at

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Nixon, K. C., 1999. The parsimony ratchet, a new method for rapid parsimony analysis. Cladistics 15, 407–414. Nylander, J. A., Wilgenbusch, J. C., Warren, D. L., Swofford, D. L., 2008. AWTY (are we there yet?): A system for graphical exploration of MCMC convergence in Bayesian phylogenetics. Bioinformatics 24, 581–583. Parris, B. S., 1977. A naturally occurring intergeneric hybrid in Grammitidaceae (Filicales): Ctenopteris heterophylla × Grammitis billardierei. New Zealand J. Bot. 15, 597–599. Parris, B. S., 1983. A taxonomic revision of the genus Grammitis Sw. (Grammitidaceae: Filicales) in New Guinea. Blumea 29, 13–222. Parris, B. S., 1984. Another intergeneric hybrid in Grammitidaceae: Ctenopteris longiceps × Grammitis sumatrana. Fern Gaz. 12, 337–340. Parris, B.S., 1990. Grammitidaceae, in Kramer, K.U., Green, P.S. (Eds.), The Families and Genera of Vascular Plants, vol.1, Pteridophytes and Gymnosperms. Springer, Berlin, pp. 153–157. Parris, B. S., 1997. Themelium, a new genus of Grammitidaceae (Filicales). Kew Bull. 52, 737– 741. Parris, B. S., 1998a. Chrysogrammitis, a new genus of Grammitidaceae (Filicales). Kew Bull. 53, 909–918. Parris, B. S., 1998b. Grammitidaceae. Flora of Australia 48, 450–468. Parris, B. S., 2003. The distribution of Grammitidaceae (Filicales) inside and outside Malesia. Telopea 10, 451–466. Parris, B. S., 2004. New combinations in Acrosorus, Lellingeria, Prosaptia and Themelium. Kew Bull. 59, 223–225. Parris, B. S., 2005. Grammitidaceae (Pteridophyta) of Mount Jaya, New Guinea, and other montane Malesian localities. Fern Gaz. 17, 183–2003. Parris, B. S., 2007. Five new genera and three new species of Grammitidaceae (Filicales) and the re-establishment of Oreogrammitis. Gard. Bull. Singapore 58, 233–274. Parris, B. S., 2010. New combinations in Oreogrammitis, Prosaptia, Radiogrammitis, Themelium and Tomophyllum (Grammitidaceae: Polypodiopsida: Monilophyta) from Malesia and the Pacific Islands. Kew Bull. 65, 123–125. Parris, B. S., 2012. Ctenopterella gabonensis, a new species of grammitid fern (Polypodiaceae) from Gabon, Africa. Fern Gaz. 19, 135–138. Parris, B. S., 2013. Archigrammitis, a new genus of grammitid fern (Polypodiaceae) from Malesia and Polynesia. Fern Gaz. 19, 135–138. Parris, B. S., Beaman, R. S., Beaman, J. H., 1992. The Plants of Mount Kinabalu 1. Ferns and Fern Allies. Royal Botanic Gardens, Kew. Parris, B. S., Given, D. R., 1976. A taxonomic revision of the genus Grammitis Sw. (Grammitidaceae: Filicales) in New Zealand. New Zealand J. Bot. 14, 85–111. Perrie, L. R., Parris, B. S., 2012. Chloroplast DNA sequences indicate the grammitid ferns (Polypodiaceae) in New Zealand belong to a single clade, Notogrammitis gen. nov. New Zealand J. Bot. 50, 457–472. Price, J. P., Clague, D. A., 2002. How old is the Hawaiian biota? Geology and phylogeny suggest recent divergences. Proc. Roy. Soc. London, Ser. B, Biol. Sci. 269, 2429–2435. R Development Core Team., 2013. R: A language and environment for statistical computing. Vienna: R Foundation for Statistical Computing.

Rakotondrainibe, F., Deroin, T., 2006. Comparative morphology and rhizome anatomy of two new species of Zygophlebia (Grammitidaceae) from Madagascar and notes on the generic circumscription of Zygophlebia and Ceradenia. Taxon 55, 145–152. Rambaut, A., Suchard, M. A., Xie, D., Drummond, A. J., 2013. Tracer v 1.5. Available from http://beast.bio.ed.ac.uk/Tracer. Ranker, T. A., Geiger, J. M. O., Kennedy, S. C., Smith, A. R., Haufler, C. H., Parris, B. S., 2003. Molecular phylogenetics and evolution of the endemic Hawaiian genus Adenophorus (Grammitidaceae). Molec. Phylogen. Evol. 26, 337–347. Ranker, T. A., Smith, A. R., Parris, B. S., Geiger, J. M. O., Haufler, C. H., Straub, S. C. K., Schneider, H., 2004. Phylogeny and evolution of grammitid ferns (Grammitidaceae): A case of rampant morphological homoplasy. Taxon 53, 415–428. Ranker, T. A., Sundue, M. A., Labiak, P., Parris, B. S. Rouhan, G., 2010. New insights into the phylogeny and historical biogeography of the Lellingeria myosuroides clade (Polypodiaceae). PLOS Currents Tree of Life., 2010 Nov 18. Edition 1. Doi: 10.1371/currents.RRN1197. Ree, R.H., Smith, S. A., 2008. Maximum likelihood inference of geographic range evolution by dispersal, local extinction, and cladogenesis. Syst. Biol. 57, 4–14. Revell, L., 2012. Phytools: An R package for phylogenetic comparative biology (and other things). Meth. Ecol. Evol. 3, 217–223. Ronquist, F., Huelsenbeck, J. P., 2003. MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19, 1572–1574. Rouhan, G., Labiak, P. H., Randrianjohany, E., Rakotondrainibe, F., 2012. Not so Neotropical after all: The grammitid fern genus Leucotrichum (Polypodiaceae) is also Paleotropical, as revealed by a new species from Madagascar. Syst. Bot. 37, 331–338. Schneider, H., Smith, A. R., Cranfill, R., Hildebrand, T. J., Haufler, C. H., Ranker, T. A., 2004. Unraveling the phylogeny of polygrammoid ferns (Polypodiaceae and Grammitidaceae): Exploring aspects of the diversification of epiphytic plants. Molec. Phylogen. Evol. 31, 1041–1063. Schneider, H., Ranker, T. R., Russell, S. J., Cranfill, R., Geiger, J. M. O., Aguraiuja, R., Wood, K. R., Grundmann, M., Kloberdanz, K., Vogel, J. C., 2005. Origin and diversification of the Hawaiian fern genus Diellia Brack. (Aspleniaceae, Polypodiidae). Proc. Roy. Soc. London, Ser. B, Biol. Sci. 272, 455–460. Schuettpelz, E., Pryer, K. M., 2009. Evidence for a Cenozoic radiation of ferns in an angiosperm-dominated canopy. Proc. Natl. Acad. Sci. U.S.A.106, 11200–11205. Smith, A. R., 1993. Tersichore, a new genus of Grammitidaceae (Pteridophyta). Novon 3, 478– 489. Smith, A. R., Pryer, K. M., Schuettpelz, E., Korall, P., Schneider, H., Wolf, P. G., 2006. A classification for extant ferns. Taxon 55, 705–731. Smith, A. R., Pryer, K. M., Schuettpelz, E., Korall, P., Schneider, H., Wolf, P. G., 2008. Fern Classification in Ranker, T. A, Haufler, C. H. (Eds.), Biology and Evolution of Ferns and Lycophytes. Cambridge University Press, Cambridge, pp. 417–462. Stamatakis, A., 2006. RAxML-VI-HPC: Maximum Likelihood-based Phylogenetic Analyses with Thousands of Taxa and Mixed Models. Bioinformatics 22, 2688–2690. Sundue, M. A., 2010a. A morphological cladistic analysis of Terpsichore (Polypodiaceae). Syst. Bot. 35, 716–729.

Sundue, M. A., 2010b. A monograph of Ascogrammitis, a new genus of grammitid ferns (Polypodiaceae). Brittonia 62, 357–399. Sundue, M. A., 2013. Mycopteris, a new neotropical genus of grammitid ferns (Polypodiaceae). Brittonia 66, 174–185. Sundue, M. A., Islam, B., Ranker, T. A., 2010. Systematics of grammitid ferns (Polypodiaceae): Using morphology and plastid sequence data to resolve the circumscription of Melpomene and the polyphyletic genera Lellingeria and Terpsichore. Syst. Bot. 35, 701– 715. Sundue, M. A., Labiak, P. H., Mostacero, J., Smith, A. R., 2012. Galactodenia, a new genus of grammitid ferns segregated from Terpsichore (Polypodiaceae). Syst. Bot. 37, 339–346. Sylvester, S. P., Sylvester, M. D.P.V., Kessler, M. 2014. The world’s highest vascular epiphytes found in the Peruvian Andes. Alpine Botany: 1–7. Tryon, R.M., Tryon, A.F., 1982. Ferns and Allied Plants, with Special Reference to Tropical America. Springer-Verlag, New York. Vasco, A., Mickel, J. T., Moran, R. C., 2013. Taxonomic revision of the Neotropical species of Elaphoglossum sect. Squamipedia (Dryopteridaceae). Ann. Missouri Bot. Gard. 99, 244– 286. Wang, L., Wu, Z. Q., Xiang, Q. P., Heinrichs, J., Schneider, H., Zhang, X.-C., 2010. A molecular phylogeny and a revised classification of tribe Lepisoreae (Polypodiaceae) based on an analysis of four plastid DNA regions. Bot. J. Linn. Soc. 162, 28–38. Wilgenbush, J. C., Warren, D. L., Swofford, D. L., 2004. AWTY: A system for graphical exploration of MCMC convergence in Bayesian phylogenetic inference. Available at http://ceb.csit.fsu.edu/awty. Wilson, K. A., 1959. Sporangia of the fern genera allied with Polypodium and Vittaria. Contrib. Gray Herb. 185, 97–127. Wolf, P. G., Schneider, H., Ranker, T. A., 2001. Geographic distributions of homosporous ferns: Does dispersal obscure evidence of vicariance? J. Biogeogr. 28, 263–270.

Table 1. Type species of grammitid genera sampled Genus Species Acrosorus exaltatus (Copel.) Copel. = friderici-et-pauli (Christ) Copel. Adenophorus tripinnatifidus Gaudich. Alansmia lanigera (Desv.) Moguel & M. Kessler Archigrammitis friderici-et-pauli (Christ) Parris Ascogrammitis athyrioides (Hook.) Sundue Calymmodon cucullatus (Nees & Blume) C. Presl Ceradenia curvata (Sw.) L. E. Bishop Chrysogrammitis glandulosa (J. Sm.) Parris Cochlidium graminoides (Sw.) Kaulf. Ctenopterella blechnoides (Grev.) Parris Dasygrammitis mollicoma (Nees & Blume) Parris Enterosora campbellii Baker Galactodenia delicatula (M. Martens & Galeotti) Sundue & Labiak Grammitis marginella (Sw.) Sw. Lellingeria apiculata (Kunze ex Klotzsch) A. R. Sm. & R. C. Moran

Sampled √ √ √ √ √ √ √ √

Leucotrichum Lomaphlebia Luisma Melpomene Micropolypodium Moranopteris Mycopteris Notogrammitis Oreogrammitis Prosaptia Radiogrammitis Scleroglossum Stenogrammitis Terpsichore Themelium Tomophyllum Xiphopterella Zygophlebia

organense (Gardner) Labiak linearis (Sw.) J. Sm. = graminea (Sw.) Parris bivascularis M. T. Murillo & A. R. Sm. moniliformis (Lagasca ex Sw.) A. R. Sm. & R. C. Moran pseudotrichomanoides (Hayata) Hayata = okuboi (Yatabe) Hayata basiattenuata (Jenman) R. Y. Hirai & J. Prado taxifolia (L.) Sundue billardierei (Willd.) Parris clemensiae Copel. contigua (G. Forst.) C. Presl setigera (Blume) Parris pusillum (Blume) Alderw. myosuroides (Sw.) Labiak asplenifolia (L.) A. R. Sm. tenuisectum (Blume) Parris subsecundodissectum (Zoll.) Parris hieronymusii (C. Chr.) Parris sectifrons (Kunze ex Mett.) L. E. Bishop

Table 2. Dispersal Constraints (Q matrix) A B C A 1 0.25 0.01 B 0.25 1 0.25 C 0.01 0.25 1 D 0.25 0.25 0.25 E 0.01 0.01 0.01 F 0.25 0.01 0.01

D 0.25 0.25 0.25 1 0.01 0.01

Table 3. Summary of Molecular Sequences.

E 0.01 0.01 0.01 0.01 1 0.01

F 0.25 0.01 0.01 0.01 0.01 1

√ √ √ √ √ √ √ √ √ √ √ √

Region atpB rbcL rps4trnS trnGtrnR trnLtrnF Total

Aligned bases 1174 1329

Variable bases 394(34%) 503(38%)

Parsimony informative characters 295(25%) 320(24%)

No. of samples 173 200

778

537(69%)

393(51%)

144

1572

877(56%)

652(41%)

117

661 5514

419(61%) 2730(49%)

305(45%) 1974(35%)

178 207

Appendix 2. New Combinations.

Lomaphlebia graminea (Sw.) Parris, comb. nov., based on Polypodium gramineum Sw., Prodr.: 130, 1788. Type: Jamaica, Swartz s. n. (S, frag. from S in US; B 20 014912 image!, SBT, UPS as herb. Thunb. 24444). Lomaphlebia turquina (Maxon) Sundue & Ranker, comb. nov., based on Polypodium turquinum Maxon, Proc. Biol. Soc. Wash. 52: 115. 1939. Type: Cuba, E. L. Ekman 14558 (US n.v., S image!).

Figure 1. Fifty-percent majority rule consensus phylogram from the Bayesian analysis using the combined dataset. Numbers represent the posterior probability (PP) values of each branch. The scale bar indicates the number of substitutions per site. The dotted lines indicate where the two portions of the tree are connected. Figure 2. Biogeographical hypothesis for the grammitid ferns inferred by the DEC model using the maximum clade credibility chronogram from the BEAST analysis. Blue bars depict the median divergence time estimates with 95% HPD intervals of each node age. The most likely ancestral areas are indicated at each node by colored squares corresponding to areas indicated on the map. Two colored squares are present when ancestral ranges include two regions. Numbers at each node represent the probabilities of ancestral ranges. Areas depicted are: Neotropical (dark blue), Africa–Madagascar and islands of the Atlantic and Indian oceans (purple), tropical Asia and the Pacific (red), a temperate circumaustral region south of 28°S (orange), Hawaiian Islands and the central and eastern South Pacific (green), and the Nearctic (light blue). Scale bar indicates divergence times in millions of years.

Taxon Acrosorus friderici-etpauli (Christ) Copel. 1 Acrosorus friderici-etpauli 2 Acrosorus friderici-etpauli 3 Adenophorus abietinus (D. C. Eaton) K. A. Wilson Adenophorus epigaeus (L. E. Bishop) W. H. Wagner Adenophorus haalilioanus (Brack.) K. A. Wilson Adenophorus periens L. E. Bishop Adenophorus tenellus (Kaulf.) Ranker Alansmia aff. cultrata (Bory ex Willd.) Moguel & M. Kessler Alansmia cultrata Alansmia elastica (Bory. ex Willd.) Moguel & M. Kessler Alansmia glandulifera (A. Rojas) Moguel & M. Kessler Alansmia heteromorpha (Hook. & Grev.) Moguel & M. Kessler Alansmia lanigera (Desv.) Moguel & M.Kessler Alansmia senilis (Fee) Moguel & M. Kessler Alansmia stella (Copel.) Moguel & M. Kessler Ascogrammitis anfractuosa (Kunze ex Klotzsch) Sundue Ascogrammitis angustipes (Copel.)

Voucher Cheng-Wei Chen 1630 (TAIF) Molesworth Allen 2282 (US) J. H. Beaman 8041 (US)

Collection Locality Vietnam

atpB KM218817

rbcL KM218751

rps4-trnS KM106104

trnG-trnR no data

trnL-trnF no data

Peninsular Malaysia

no data

KM218752

KM106105

KM105963

KM106046

Sabah, Malaysia

no tata

KM218753

KM106106

KM105964

KM106047

Ranker 1100 (COLO)

Hawaii, USA

AF469778

AF468202

no data

no data

AF469791

Ranker 1103 (COLO)

Hawaii, USA

AF469779

AF468203

no data

no data

AF469792

Ranker 1561 (COLO)

Hawaii, USA

AF469775

AF468200

no data

no data

AF469788

Ranker 1114 (COLO)

Hawaii, USA

AF469774

AF468199

no data

no data

AF469787

Ranker 1352 (COLO)

Hawaii, USA

AF469773

AF468198

no data

no data

AF469786

Sundue 1162 (NY)

Ecuador

GU376477

GU376495

JN654942

JN654955

JN654968

Sundue 1214 (NY) J.Kluge 7863 (Z)

Colombia Madagascar

GU376476 KM218858

GU376496 KM218795

KM106107 *

JN654956 KM105965

JN654969 KM106048

Sundue 1765 (NY)

Costa Rica

GU376472

GU376497

JN654944

JN654957

JN654970

Sundue 2609 (NY)

Ecuador

no data

KM218803

no data

KM105966

no data

B. León 3647 UC

Peru

AY459505

AY460672

no data

no data

GU476718

Sundue 1156 (NY)

Ecuador

GU376474

GU376498

JN654941

JN654954

JN654967

Sundue 1083 (NY)

Ecuador

GU376473

GU376499

JN654939

JN654952

JN654965

Lehnert 1035 (UC)

Bolivia

GU476783

GU476853

KM106108

KM105967

GU476675

Sundue 1237(NY)

Colombia

KM218837

GU476891

KM106109

KM105968

GU476703

Sundue Ascogrammitis athyrioides (Hook.) Sundue Ascogrammitis clathrata (Sundue & M. Kessler) Sundue Ascogrammitis clavigera A. R. Sm. ex Sundue Ascogrammitis colombiense Sundue Ascogrammitis cuencana (Hieron.) Sundue Ascogrammitis davidsmithii (Stolze) Sundue Ascogrammitis dilatata (Sundue & M. Kessler) Sundue Ascogrammitis loxensis Sundue Ascogrammitis nana (Sundue & M. Kessler) Sundue Ascogrammitis pichinchae (Sodiro) Sundue Ascogrammitis pichinchensis (Hieron.) Sundue Calymmodon clavifer (Hook.) T. Moore Calymmodon gracilis (Fee.) Copel. Calymmodon gracilis Calymmodon luerssenianus (Domin) Copel. Calymmodon mnioides Copel. Calymmodon pallidivirens Parris ined. Ceradenia aff kalbreyeri (Baker) L. E. Bishop

Lehnert 261 (UC)

Peru

KM218840

GU476856

KM106110

KM105969

GU476704

Kromer 1237(UC)

Bolivia

KM218838

GU476843

KM106111

KM105970

GU476708

Schneider 2400 (UC)

Venezuela

KM218839

GU476925

KM106112

no data

GU476709

Dassler 94-7-13-1 (OS)

Colombia

GU476805

GU476827

KM106113

no data

GU476711

Lehnert 1164 (GOET)

Ecuador

no data

GU476851

KM106114

KM105971

GU476714

Sundue 785 (NY)

Bolivia

GU376639

GU387012

GU387122

GU387205

GU387284

Labiak 4728 UUPCB)

Bolivia

GU376640

GU387033

GU387124

GU387206

GU387285

Sundue 1073 (NY)

Ecuador

GU376641

GU386995

GU387125

GU387207

GU387286

Labiak 4725 (UPCB)

Bolivia

GU376642

GU387031

GU387126

GU387208

GU387287

Wilson 2816a (UC)

Ecuador

no data

GU476928

KM106115

no data

GU476730

Lehnert 1577 (UC)

Ecuador

GU476816

GU476854

KM106116

no data

GU476732

Sundue 2235 (VT)

Sabah, Malaysia

KM218818

KM218804

KM106117

no data

KM106049

Fei Wei Li 980 (TAIF)

Philippines

no data

no data

no data

KM105972

no data

Chiou 97-09-12-01 (TAIF) Kessler 14264 (VT)

Taiwan Australia

AY459451 KM218819

AY362341 KM218806

no data KM106118

no data KM105973

GU476618 KM106050

Karger 1675 (Z)

West Papua, Indonesia

KM218820

KM218807

KM106120

no data

KM106052

Sundue 2218 (VT)

Sabah, Malaysia

no data

KM218805

KM106119

no data

KM106051

Vasco & Sundue 701 (NY)

Colombia

GU476747

GU476825

KM106121

no data

GU476622

Ceradenia aulaeifolia L. E. Bishop ex A. R. Sm. Ceradenia ayapoyana M. Kessler & A. R. Sm. Ceradenia curvata (Sw.) L. E. Bishop Ceradenia farinosa (Hook.) L. E. Bishop Ceradenia fucoides (Christ) L. E. Bishop Ceradenia intonsa L. E. Bishop ex Leon-Parra & Mostacero Ceradenia intricata (C. V. Morton) L. E. Bishop ex A. R. Sm. Ceradenia jungermannioides (Klotzsch) L. E. Bishop Ceradenia kalbreyeri (Baker) L. E. Bishop Ceradenia madidiensis M. Kessler & A. R. Sm. Ceradenia pearcei (Baker) L. E. Bishop Ceradenia pilipes (Baker) L. E. Bishop Chrysogrammitis glandulosa (J. Sm.) Parris Chrysogrammitis musgraviana (Baker) Parris 1 Chrysogrammitis musgraviana 2 Cochlidium punctatum (Raddi) L. E. Bishop Cochlidium rostratum (Hook.) Maxon ex C. Chr. Cochlidium seminudum (Willd.) Maxon Cochlidium serrulatum (Sw.) L. E. Bishop Ctenopterella denticulata

A. Rojas & al. 3232 (CR)

Costa Rica

AY459453

AY460619

KM106122

KM105974

GU476623

I. Jiménez 1114 (GOET, LPB, UC) I. Jiménez 1559 (UC, LPB)

Bolivia

no data

KM218811

KM106123

KM105975

KM106053

Bolivia

KM218821

KM218788

no data

KM105976

KM106054

David Neill 11985 (MO)

Ecuador

KM218823

KM218790

KM106124

KM105977

KM106055

Sundue 1766 (NY)

Costa Rica

GU476749

GU476907

KM106125

no data

GU476625

Sundue 1321 (NY)

Colombia

GU476750

GU476901

KM106126

no data

GU476626

Lehnert 1108 (UC, LPB)

Bolivia

KM218833

KM218791

KM106127

KM105978

KM106056

A. R. Smith 2576 (UC)

Costa Rica

AY459454

AY460620

no data

no data

no data

Sundue 1759 (NY)

Colombia

GU476744

GU476905

KM106128

KM105979

GU476617

I. Jiménez 1089 (UC, LPB)

Bolivia

KM218834

no data

KM106129

no data

KM106057

A.F. Fuentes & Villalobos 13997 (MO) A. Rojas & al. 3233 (INB)

Bolivia

KM218822

KM218789

KM106130

KM105980

KM106058

Costa Rica

AY459456

AY460622

no data

no data

GU476620

Ranker 2195 (COLO)

Sabah, Malaysia

JF514082

JF514014

no data

no data

JF514048

Kessler 12570 (UC)

Sabah, Malaysia

AY459458

AY460624

KM106131

no data

GU476630

Sundue 2234 (VT)

Sabah, Malaysia

KM218825

KM218797

KM106132

KM105981

KM106059

Silva 3914 (UC)

Brazil

JF514057

JF513987

no data

no data

GU476631

I. Valdespino & J. Aranda 180 (UC) S. R. Hill 29102A no voucher** Hirai & Swartzburd 541 (SP) Ranker 2113 (COLO)

Panama

AY459459

AY460626

no data

KM105982

no data

Dominican Republic

AY459460

AY460627

no data

no data

KM106060

Brazil

JF514078

JF514010

KM106133

KM105983

JF514044

Sabah, Malaysia

JF514081

JF514013

no data

no data

JF514047

(Blume) Parris 1 Ctenopterella denticulata 2 Dasygrammitis brevivenosa (Alderw.) Parris 1 Dasygrammitis brevivenosa 2 Dasygrammitis crassifrons (Baker) Parris Dasygrammitis fuscata (Blume) Parris Dasygrammitis malaccana (Baker) Parris Dasygrammitis mollicoma (Nees & Blume) Parris Enterosora enterosoroides (Christ) A. Rojas Enterosora percrassa (Baker) L. E. Bishop 1 Enterosora percrassa 2 Enterosora trichosora (Hook.) L. E. Bishop Enterosora trifurcata (L.) L. E. Bishop Enterosora trifurcata Enterosora trifurcata Galactodenia delicatula (M. Martens & Galeotti) Sundue & Labiak Galactodenia parrisiae Sundue & Labiak Galactodenia subscabra (Klotzsch) Sundue & Labiak 1 Galactodenia subscabra 2 Grammitis bryophila (Maxon) F. Seym. Grammitis cryptophlebia (Baker) Copel.

Cheng-Wei Chen 2010 (TAIF) Sundue 2222 (VT)

Java, Indonesia

no data

no data

KM106134

KM105984

no data

Sabah, Malaysia

KM218828

KM218774

KM106135

KM105985

no data

Sundue 2206 (VT)

Sabah, Malaysia

KM218829

KM218775

KM106136

KM105986

KM106061

John Game 95-80 (UC)

Fiji

JF514062

KM218777

KM106137

no data

KM106062

Parris 12784 (AK)

Peninsular Malaysia

no data

KM218778

KM106138

KM105987

no data

Parris 12796 (AK)

Peninsular Malaysia

no data

KM218779

KM106139

KM105988

no data

Parris 12797 (AK)

Peninsular Malaysia

no data

KM218776

no data

KM105989

KM106063

Sundue 1172 (NY)

Ecuador

GU476756

no data

KM106140

no data

GU476634

Sundue 1665 (NY)

Costa Rica

GU476757

GU476882

KM106141

no data

GU476882

Moraga & Rojas 508 (UC) Moran 7569 (NY)

Costa Rica Ecuador

AY459468 no data

AY460635 GU476920

no data no data

no data no data

GU476636 GU476637

Ranker 1608 (COLO)

Puerto Rico

AY459521

AY460636

KM106142

no data

no data

Moran 7515 (NY) Sundue 1774 (NY) Mickel 4639 (NY)

Ecuador Costa Rica Mexico

no data no data no data

no data no data no data

no data no data GU387123

no data no data no data

GU476633 GU476632 KM106064

Monro & Knapp 5721 (NY) A. Rojas & al. 3211 (CR, INB, MO, UC)

Panama

no data

KM218794

no data

KM105990

no data

Costa Rica

AY459511

AY460677

no data

no data

GU476740

Moran 8078 (NY)

Costa Rica

GU476821

GU476860

GU387127

GU387209

GU476739

A. Rojas & al. 3240 (UC)

Costa Rica

AF469784

AF468208

KM106143

no data

AF469797

Kluge 7936 (Z)

Madagascar

KM218815

KM218799

KM106144

KM105992

KM106065

Grammitis deplanchei (Baker) Copel. Grammitis diminuta (Baker) Copel. 1 Grammitis diminuta 2 Grammitis kyimbilensis (Brause ex Brause & hieron.) Copel. Grammitis melanoloma (Cordemoy) Tardieu Grammitis paramicola L. E. Bishop Grammitis pseudaustralis E. Fourn. Grammitis stenophylla Parris Lellingeria apiculata (Kunze ex Klotzsch) A. R. Sm. & R. C. Moran Lellingeria bishopii Labiak Lellingeria flagellipinnata M. Kessler & A. R. Sm. Lellingeria hirsuta A. R. Sm. & R. C. Moran Lellingeria humilis (Mett.) A. R. Sm. & R. C. Moran Lellingeria major (Copel.) A. R. Sm. & R. C. Moran Lellingeria randallii (Maxon) A. R. Sm. & R. C. Moran Lellingeria simacensis (Rosenst.) A. R. SM. & R. C. Moran Lellingeria subsessilis (Baker) A. R. Sm. & R. C. Moran Lellingeria suprasculpta (Christ) A. R. Sm. & R. C. Moran Lellingeria suspensa (L.) A. R. Sm. & R. C. Moran

Hodel 1450 (UC)

New Caledonia

AY459471

AY460639

KM106145

no data

GU476639

Papadopulos AP845 (NSW) Papadopulos AP794 (NSW) Kessler 12773 (UC)

Lord Howe Island

no data

JF950809

no data

no data

JF950910

Lord Howe Island

no data

JF950810

no data

no data

JF950911

Madagascar

EF178641

EF178624

KM106147

KM105993

EF178659

Ranker 1504 & Adsersen (COLO) I. Jiménez & S. Gallegos (UC, LPB) L. Tibell 19878 (AK)

La Reunion

AY459475

AY460643

KM106148

no data

GU476641

Bolivia

KM218816

KM218801

KM106149

KM105994

KM106067

New Caledonia

no data

KM218785

KM106150

no data

KM106068

Parris 12576 (AK)

Australia

JX499239

JQ904084

no data

no data

JQ911714

Labiak 4223 (UPCB); Salino 3009 (UC)

Brazil

GU376573

GU387025

GU387047

GU387133

GU387215

Sundue 1159 (NY) Labiak 4726 (NY)

Ecuador Bolivia

GU376574 GU376582

GU387000 GU387032

GU387048 GU387056

GU387134 GU387142

GU387216 GU387224

A. Rojas & E. Fletes 3145 (UC) Sundue & Vasco 1289 (NY) Sundue 1147 (NY)

Costa Rica

AY459482

AY460649

no data

no data

no data

Colombia

GU476763

GU476897

no data

no data

GU476649

Ecuador

GU376592

GU386998

GU387069

GU387155

GU387233

Mickel 9495 (NY)

Trinidad

KM218860

GU386992

GU387090

KM105996

GU387253

Sundue 1169 (NY)

Ecuador

GU376610

GU387001

GU387092

GU387175

GU387255

Sundue 1139 (NY)

Ecuador

GU376613

GU386997

GU387095

GU387178

GU387258

Lellinger 989 (US)

Costa Rica

GU376614

GU386987

GU387096

GU387179

GU387259

Sundue 1047 (NY)

Ecuador

GU376618

GU386994

GU387100

GU387183

GU387263

Leucotrichum mitchelliae (Baker ex Hemsl.) Labiak Leucotrichum mortonii (Copel.) Labiak Leucotrichum organense (Gardner) Labiak Leucotrichum pseudomitchelliae (Lellinger) Labiak Leucotrichum schenckii (Hieron.) Labiak Lomaphlebia turquina (Maxon) Sundue & Ranker Melpomene allosuroides (Rosenst.) A. R. Sm. & R. C. Moran Melpomene anazalea Sundue & Lehnert Melpomene erecta (C. V. Morton) A. R. Sm. & R. C. Moran Melpomene firma (J. Sm.) A. R. Sm. & R. C. Moran Melpomene flabelliformis (Poir.) A. R. Sm. & R. C. Moran Melpomene melanosticta (Kunze) A. R. Sm. & R. C. Moran Melpomene moniliformis (Lag. ex Sw.) A. R. Sm. & R. C. Moran Melpomene xiphopteridoides (Liebmann) A. R. Sm. & R. C. Moran Microgramma bifrons (Hook.) Lellinger Microgramma bifrons Micropolypodium okuboi (Yatabe) Hayata Micropolypodium

Breedlove & Almeda 48307 (NY) Liogier 16026 (NY)

Mexico

GU376479

GU376488

JN654937

JN654950

JN654963

Dominican Republic

GU376478

GU376489

no data

no data

no data

Labiak 4302 (UPCB)

Brazil

GU376485

GU376492

JN654946

JN654959

JN654972

A. Rojas 3005 (MO)

Costa Rica

AY459484

AY460652

no data

no data

no data

Salino 4538 (BHCB, UC)

Brazil

AY459483

AY460651

no data

no data

KM106072

C. Sanchez 82061 (HAJB)

Cuba

KM218814

KM218800

no data

no data

KM106069

Solomon 1289 (NY)

Bolivia

GU376628

GU387038

GU387109

GU387192

GU387273

Sundue 1290 (NY)

Colombia

GU476773

GU476898

KM106152

KM105997

GU476662

Nunez 25 (NY)

Bolivia

GU376629

GU387013

GU387110

GU387193

GU387274

Labiak 4733 (UPCB)

Bolivia

GU376630

GU387035

GU387112

GU387195

GU387276

Sundue 1303 (NY)

Colombia

GU376631

GU387005

GU387113

GU387196

GU387277

Labiak 4156 (NY)

Brazil

GU376633

GU387024

GU387115

GU387198

GU387279

M. Moraga & A. Rojas 446 (INB)

Costa Rica

AY459486

AY460654

no data

no data

GU476664

Sundue 1300 (NY)

Colombia

GU376637

GU387004

GU387119

KM105998

GU387282

van der Werff 18062 (UC) Peru

EF463499

no data

no data

no data

DQ642224

Neill 8309 (UC) Parris 12154 (AK)

Ecuador Japan

KM218862 JF514064

AY362582 JF513994

AY362654 no data

no data no data

KM106073 JF514028

Miehe 00-093-32 (UC)

Bhutan

JF514068

JF513999

no data

no data

JF514032

sikkimense (Hieron.) X. C. Zhang 1 Micropolypodium sikkimense 2 Moranopteris achilleifolia (Kaulf.) R. Y. Hirai & J. Prado Moranopteris blepharidea (Copel.) R. Y. Hirai & J. Prado Moranopteris caucana (Hieron.) R. Y. Hirai & J. Prado Moranopteris cookii (Underw. & Maxon ex Maxon) R. Y. Hirai & J. Prado Moranopteris hyalina (Maxon) R. Y. Hirai & J. Prado Moranopteris longisetosa (Hook.) R. Y. Hirai & J. Prado Moranopteris plicata (A. R. Sm.) R. Y. Hirai & J. Prado Moranopteris truncicola (Klotzsch) R. Y. Hirai & J. Prado Mycopteris amphidasyon (Kunze ex Mett.) Sundue Mycopteris attenuatissima (Copel.) Sundue Mycopteris leucosticta (J. Sm.) Sundue Mycopteris longicaulis (Sundue & M. Kessler ) Sundue Mycopteris longipilosa Sundue Mycopteris praeceps (Sundue & M. Kessler)

Xian & al. s.n. (UC)

China

JF514087

JF514019

no data

no data

JF514054

J. Cordeiro & O. Ribas 1398 (UC)

Brazil

AY459499

AY460666

KM106153

KM105999

no data

Jiménez 708 (GOET)

Bolivia

JF514065

JF513995

KM106154

KM106000

JF514029

Lehnert 182 (GOET)

Ecuador

JF514071

JF514002

KM106155

KM106001

JF514035

Sundue 1771 (NY)

Costa Rica

JF514076

JF514007

KM106156

KM106002

JF514040

Lehnert 1426 (GOET)

Ecuador

JF514070

JF514001

KM106157

KM106003

JF514034

Lehnert 596 (GOET)

Bolivia

JF514072

JF514003

KM106158

KM106004

JF514036

Lehnert 929 (UC)

Ecuador

JF514074

JF514005

KM106159

KM106005

JF514038

Vasco & Sundue 626 (NY)

Colombia

JF514084

JF514016

KM106160

KM106006

JF514051

Moran 7646 (NY)

Ecuador

GU476759

GU476922

KM106161

KM106007

GU476638

Sundue 1098 (NY)

Ecuador

GU476866

GU476866

GU387121,

GU387204

GU476705

Lehnert 1128 (UC)

Ecuador

GU476811

GU476848

KM106162

no data

GU476720

Jimenez 373 (UC)

Bolivia

GU476813

GU476840

KM106163

no data

GU476724

Sundue & Martin 1033 (NY) Jimenez 2173 (UC)

Ecuador

GU476814

GU476861

KM106164

KM106008

GU476726

Bolivia

GU476817

GU476839

KM106165

no data

GU476734

Sundue Mycopteris subtilis (Kunze ex Klotzsch) Sundue Mycopteris sp. aff. subtilis Mycopteris taxifolia (L.) Sundue Niphidium crassifolium (L.) Lellinger Niphidium crassifolium Notogrammitis angustifolia (Jacq.) Parris 1 Notogrammitis angustifolia 2 Notogrammitis billardierei (Willd.) Parris 1 Notogrammitis billardierei 2 Notogrammitis billardierei 3 Notogrammitis ciliata (Colenso) Parris 1 Notogrammitis ciliata 2 Notogrammitis ciliata 3 Notogrammitis crassior (Kirk) Parris Notogrammitis givenii (Parris) Parris Notogrammitis heterophylla (Labill.) Parris Notogrammitis patagonica (C. Chr.) Parris Notogrammitis pseudociliata (Parris) Parris Notogrammitis rawlingsii (Parris) Parris Notogrammitis rigida

Lehnert 1396 (UC)

Ecuador

GU376643

GU386984

GU387128

GU387210

GU387288

Sundue & Schuettpelz 1077 (NY) Labiak 4018 (NY)

Ecuador

no data

GU476865

KM106166

no data

GU476687

Brazil

GU476800

GU476914

KM106167

KM106009

GU476699

Schuettpelz 209 (DUKE)

Ecuador

EF463503

EF463254

no data

no data

no data

Kreier s.n. (GOET) Parris 12423 (AK)

cultivated New Zealand

no data JX499252

no data JQ904083

EU250358 no data

no data no data

EU250359 JQ904113

Lendemer 30006 (NY)

Australia

KM218824

KM218770

KM106168

KM106010

KM106074

Parris 12421 (AK)

New Zealand

AY459469

JQ904094

KM106169

KM106011

JQ904102

Brownsey P022138 (WELT) Sundue s.n. (VT)

Australia

JX499250,

JQ904096

no data

no data

JQ904103

Australia

no data

KM218755

KM106170

KM106012

KM106075

Parris 12360 (AK)

New Zealand

JX499247

JQ904078

no data

no data

JQ904104

Parris 12657 (AK) Jane s.n. (AK) Perrie 3742 (WELT)

New Zealand New Zealand New Zealand

JX499248 JX499249 JX499241

JQ904095 JQ904097 JQ904077

no data no data no data

no data no data no data

JQ904106 JQ904105 JQ904115

Perrie 3842 (WELT)

New Zealand

JX499242

JQ904079

no data

no data

JQ904110

Parris 12419 (AK)

New Zealand

AY459462

AY460629

no data

no data

JQ904100

Perrie 4760 (WELT)

New Zealand

JX499246

JQ904088

no data

no data

JQ904075

Perrie 4088 (WELT)

New Zealand

JX499245

JQ904080

no data

no data

JQ904117

Young s.n. (AK 296949)

New Zealand

JX499240

JQ904085

no data

no data

JQ904119

Perrie 4857 (WELT)

New Zealand

no data

JQ904082

no data

no data

JQ904121

(Hombron) Parris Oreogrammitis adspersa (Blume) Parris Oreogrammitis congener (Blume) Parris 1 Oreogrammitis congener 2 Oreogrammitis forbesiana (W. H. Wagner) Parris Oreogrammitis graniticola Parris ined. Oreogrammitis hookeri (Brack.) Parris Oreogrammitis locellata (Baker) Parris Oreogrammitis reinwardtioides (Copel.) Parris Oreogrammitis reinwardtioides Oreogrammitis uapensis (E. D. Br.) Parris Oreogrammitis ultramaficola Parris ined. Oreogrammitis wurunuran (Parris) Parris 1 Oreogrammitis wurunuran 2 Polypodium glycyrrhiza D. C. Eaton Prosaptia alata (Blume) Christ Prosaptia celebica (Blume) Tagawa & K. Iwats. Prosaptia contigua (G. Forst.) C. Presl 1 Prosaptia contigua 2 Prosaptia davalliacea (F. Muell. & Baker) Copel.

Sundue 2233 (VT)

Sabah, Malaysia

no data

KM218808

KM106171

KM106013

KM106076

Parris 12782 (AK)

Peninsular Malaysia

KM218845

KM218762

KM106172

KM106014

no data

T. A. Ranker 2126 & S. T. Klimas (BORH, SAN, SNP, COLO) Ranker 1321 (COLO)

Sabah, Malaysia

EF178634

EF178617

no data

no data

EF178650

Hawaii, USA

AY459472

AY460640

no data

no data

EF178651

Sundue 2212 (VT)

Sabah, Malaysia

KM218846

KM218757

no data

KM106015

KM106078

Ranker 1116 (COLO)

Hawaii, USA

AY459473

AY460642

no data

no data

EF178655

M.S. Clemens 12380 (UC)

Papua New Guinea

no data

KM218763

KM106173

no data

KM106079

Sundue 2202 (VT)

Sabah, Malaysia

KM218847

no data

no data

KM106016

KM106080

Ranker 2228 (COLO)

Sabah, Malaysia

no data

KM218756

KM106174

no data

EF178661

K. R. Wood 10825 (US)

Marquesas Islands

no data

KM218787

no data

KM105995

KM106070

Ranker 2153 (COLO)

Sabah, Malaysia

KM218841

KM218754

KM106151

no data

KM106071

Kessler 14307 (VT)

Australia

KM218842

KM218760

KM106175

KM106017

no data

Kessler 14251 (VT

Australia

KM218843

KM218761

KM106176

KM106018

no data

Haufler & Mesler s.n. (KANU) T. Flynn 6004 (UC)

United States

AY459518

U21146

no data

no data

GU476671

Kosrae, Micronesia

AY459493

AY460660

no data

KM106019

KM106081

Parris 12835 (K)

Philippines

KM218849

KM218786

KM106178

KM106021

no data

Sundue 2219 (VT)

Sabah, Malaysia

KM218850

KM218765

no data

KM106022

KM106083

Sundue 2740 (VT) Sundue 2210 (VT)

Australia Sabah, Malaysia

KM218851 KM218852

KM218767 KM218798

KM106180 KM106179

KM106025 KM106023

no data KM106084

Prosaptia maidenii (Watts) Parris Prosaptia obliquata (Blume) Mett. Prosaptia palauensis Hosokawa Prosaptia pinnatifida C. Presl Prosaptia venulosa (Blume) M. G. Price Radiogrammitis cheesemanii (Parris) Parris Radiogrammitis exilis Parris ined. Radiogrammitis exilis Radiogrammitis holttumii (Copel.) Parris Radiogrammitis kinabaluensis (Copel.) Parris Radiogrammitis multifolia (Copel.) Parris Radiogrammitis parva (Brause) Parris Radiogrammitis setigera (Blume) Parris Radiogrammitis setigera Scleroglossum pusillum (Blume) Alderw. Scleroglossum sulcatum (Kuhn) Alderw. Scleroglossum wooroonooran (F. M. Bailey) C. Chr. 1 Scleroglossum wooroonooran 2 Serpocaulon fraxinifolium (Jacq.) A. R. Sm. Serpocaulon triseriale (Sw.) A. R. Sm. Stenogrammitis hartii (Jenman) Labiak

Kessler 14243 (VT)

Australia

no data

KM218766

no data

KM106024

no data

Karger 166 (Z)

Philippines

KM218854

KM218768

KM106181

KM106026

KM106085

R. J. Rondeau 93-011& Rodda (UC) Karger 608 (Z)

Palau, Micronesia

no data

AY460662

no data

KM106027

no data

Philippines

KM218848

KM218764

KM106177

KM106020

KM106082

Karger 859 (Z)

Philippines

KM218853

KM218769

KM106182

no data

KM106086

John Game 85-53 (UC)

Rarotonga

KM218861

no data

KM106183

no data

KM106087

Sundue 2225 (VT)

Sabah, Malaysia

KM218857

no data

KM106201

no data

KM106103

Sundue 2223 (VT) Sundue 2208 (VT)

Sabah, Malaysia Sabah, Malaysia

no data KM218856

no data KM218771

no data KM106184

KM106045 KM106028

no data EF178653

Sundue 2211 (VT)

Sabah, Malaysia

KM218855

KM218773

KM106185

no data

KM106088

M. Tagawa, K. Iwatsuki, Thailand & N. Fukuoka T4797 (US) Ranker 1763a (COLO, UC) Papua New Guinea

no data

KM218772

KM106146

no data

KM106066

AY459476

AY460644

no data

no data

no data

Fei-Wei Li 982 (TAIF)

Philippines

no data

no data

no data

KM106029

no data

Karger 813 (Z) Parris 12779 (AK)

Philippines Peninsular Malaysia

no data no data

KM218759 KM218812

KM106186 KM106187

no data KM106030

KM106089 KM106090

Bowden-Kerby 24182b (UC, GUAM) Kessler 14289 (VT)

Pohnpei, Micronesia

AY459498

AY460665

KM106188

no data

JF514049

Australia

KM218835

KM218809

KM106189

KM106031

KM106091

Kessler 14304 (VT)

Australia

KM218836

KM218810

KM106190

KM106032

KM106092

Jimenez 3302 (UC)

Bolivia

no data

EF551071

EF551096

no data

EF551134

Smith 2902 (UC)

Brazil

no data

EF551078

EF551118

no data

EF551157

Lellinger 455 (US)

Dominican Republic

GU376583

GU386985

GU387057

GU387143

GU387225

Stenogrammitis hellwigii (Mickel & Beitel) Labiak Stenogrammitis jamesonii (Hook.) Labiak Stenogrammitis limula (Christ) Labiak Stenogrammitis myosuroides (Sw.) Labiak Stenogrammitis oosora (Baker) Labiak Stenogrammitis prionodes (Mickel & Beitel) Labiak Stenogrammitis pumila (Labiak) Labiak Stenogrammitis saffordii (Maxon) Labiak Stenogrammitis wittigiana (Fee & Glaziou ex Fee) Labiak Terpsichore asplenifolia (L.) A. R. Sm. Terpsichore chrysleri (Proctor ex Copel.) A. R. Sm. Terpsichore eggersii (Baker) A. R. Sm. Terpsichore hanekeana (Proctor) A.R.Sm. Terpsichore lehmanniana (Hieron.) A. R. Sm. Themelium conjunctisorum (Baker) Parris Themelium decrescens (Christ) Parris Themelium halconense (Copel.) Parris Tomophyllum macrum (Copel.) Parris Tomophyllum macrum Tomophyllum repandulum (Mett.)

Mickel & Leonard 4636 (NY) Sundue 1149 (NY)

Mexico

GU376584

GU386990

GU387058

GU387144

GU387226

Ecuador

GU376596

GU386999

GU387074

GU387160

GU387237

Sundue 1736 (NY)

Costa Rica

GU476765

GU476903

GU387066

GU387152

GU476651

Liogier 4 (NY)

Guatemala

GU376599

GU386988

GU387077

GU387163

GU387240

Cable 77 (K)

Cameroon

GU376600

GU386972

GU387078

GU387164

GU387241

Mickel & Beitel 6567 (NY) Mexico

GU376605

GU386991

GU387086

GU387170

GU387249

Labiak 4015 (UPCB)

Brazil

GU376608

GU387022

GU387089

GU387173

GU387252

Ranker 1892 (BISH, COLO) Labiak 4441 (UPCB)

Hawaii, USA

EF178645

EF178628

KM106191

no data

EF178662

Brazil

GU376625

GU387029

GU387106

GU387189

GU387270

Moraga & Rojas 506 (MO) I. Jiménez 1369 (UC, LPB)

Costa Rica

JF514059

KM218802

KM106192

no data

no data

Bolivia

KM218859

KM218813

no data

KM106033

no data

S. Hill 29109 (UC)

Dominican Republic

AF469785

AF468209

AY362694

no data

AF469798

Ranker 1610 (COLO)

Puerto Rico

AY459503

AY460670

no data

no data

no data

K. A. Wilson 2589 (UC)

Ecuador

AY459506

AY460673

no data

no data

no data

Ranker 1758 & Trapp (COLO, UC)

Papua New Guinea

AY459514

AY460680

no data

no data

no data

Parris 12836 (AK)

Philippines

no data

KM218758

KM106193

KM106034

KM106093

Sundue 2226 (VT)

Sabah, Malaysia

KM218844

no data

KM106194

KM106035

KM106094

Karger 165 (Z)

Philippines

KM218830

no data

no data

no data

no data

Karger 532 (Z) Ranker 1767 & Trapp (COLO, UC)

Philippines Papua New Guinea

no data AY459466

KM218780 AY460633

no data KM106036

no data KM106036

no data no data

Parris Tomophyllum secundum (Ridl.) Parris Tomophyllum walleri (Maiden & Betche) Parris Xiphopterella hieronymusii (C. Chr.) Parris Xiphopterella sparsipilosa (Holttum) Parris Zyglophlebia devoluta (Baker) Parris 1 Zygophlebia devoluta 2 Zygophlebia mathewsii Zygophlebia mathewsii (Kunze ex Mett.) L. E. Bishop Zygophlebia sectifrons (Kunze ex Mett.) L. E. Bishop Zygophlebia sp.

Ranker 2193 (COLO)

Sabah, Malaysia

KM218832

KM218781

KM106195

KM106037

KM106095

Kessler 14250 (VT)

Australia

KM218831

KM218782

KM106196

KM106038

KM106096

Parris 12781 (AK)

Peninsular Malaysia

no data

KM218783

KM106197

KM106039

KM106097

Parris 12790 (AK)

Peninsular Malaysia

no data

KM218784

KM106198

KM106040

KM106098

Kluge 7940 (Z)

Madagascar

KM218826

KM218792

no data

KM106041

KM106099

Kluge 7937 (Z) Sundue 1254 & Vasco (NY) Sundue 1119 (NY)

Madagascar Colombia

KM218827 GU476895

KM218793 GU476895

KM106199 no data

KM106042 no data

KM106100 GU476743;

Ecuador

no data

GU476868

GU476868

KM106043

no data

Sundue 1757 (NY)

Costa Rica

no data

GU476904

GU476905

KM106044

KM106101

Massawe & Phillipson 467 (MO)

Madagascar

no data

KM218796

KM106200

no data

KM106102

Figure 1

Figure 2 bottom

Figure 2 top

• • • • •

We examined historical biogeography of grammitid ferns The grammitid clade arose among Neotropical polypod ancestors about 31.4 Ma Afro-Malagasy species were more closely related to Neotropical taxa A large clade distributed in the Asia-Malesia-Pacific region originated ca 23.4 Ma The circumaustral genus Notogrammitis originated in Australia or New Zealand