Ranker 04

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Ranker & al. Phylogeny and evolution of grammitid ferns
Phylogeny and evolution of grammitid ferns (Grammitidaceae): a case of

rampant morphological homoplasy
Tom A. Ranker1, Alan R. Smith2, Barbara S. Parris3, Jennifer M. O. Geiger1,6, Christopher H.
Haufler4, Shannon C. K. Straub1,7, & Harald Schneider5
1 University
Museum and Department of Ecology and Evolutionary Biology, University of Colorado, Boulder,
Colorado 80309, U.S.A. ranker@colorado.edu (author for correspondence)
2 University Herbarium, University of California, Berkeley, California 94720, U.S.A. arsmith@uclink4.berkeley.edu
3 Fern Research Foundation, 21 James Kemp Place, Kerikeri, Bay of Islands, New Zealand.
bsparris@igrin.co.nz
4 Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, Kansas 66045, U.S.A.
vulgare@ku.edu
5 Albrecht-von-Haller-Institut fr Pflanzenwissenschaften, Abt. Systematische Botanik, Georg-AugustUniversitt, Gttingen, Untere Karsple 2, 37073 Gttingen, Germany. hschneid@duke.edu
6 Current address: Carroll College, Department of Natural Sciences, 1601 North Benton Ave., Helena,
Montana 59625, U.S.A. jgeiger@carroll.edu
7 Current address: Department of Plant Biology, 228 Plant Science Building, Cornell University, Ithaca, New
York 14853, U.S.A. ss463@cornell.edu
We conducted phylogenetic analyses of the fern family Grammitidaceae using sequences from two cpDNA
genes and from morphological characters. Data were obtained for 73 species from most recognized genera in
the family. The genera Adenophorus, Ceradenia, Calymmodon, Cochlidium, Enterosora, and Melpomene were
each strongly supported as being monophyletic. Other recognized genera that were not supported as monophyletic included Ctenopteris, Grammitis, Lellingeria, Micropolypodium, Prosaptia, and Terpsichore. Several
previously unrecognized clades were identified, some of which are characterized by distinctive morphological
features. Analyses of the distribution of morphological character states on our inferred phylogeny showed
extremely high levels of homoplastic evolution for many different characters. Homoplasy for morphological
characters was considerably greater than for molecular characters. Many of the characters that exhibited high
levels of convergent or parallel evolution across the phylogeny are features that have been commonly used to
circumscribe genera in this group (e.g., leaf blade dissection, various rhizome scale characters, and glandular
paraphyses). Conversely, some of the characters that exhibited relatively low levels of homoplasy have either
not been regarded as having taxonomic value or have been ignored (e.g., root insertion, rhizome scale sheen).
Our data support a New World origin of Grammitidaceae, with Old World taxa generally being more evolutionarily derived. Several clades are either primarily Neotropical or primarily Paleotropical but also have a few
members distributed in the opposite hemisphere. Thus, we postulate multiple, independent dispersal and colonization events in several lineages.
KEYWORDS: atpB, Grammitidaceae, homoplasy, molecular phylogeny, morphology, rbcL.
INTRODUCTION
Grammitids are a primarily tropical and subtropical
group of mostly epiphytic ferns (Fig. 1). The group comprises approximately 750 species and is generally characterized by green tetrahedral spores, sporangial stalks
consisting of only one row of cells, and leaf traces of single vascular strands (Parris, 1990, 1998b). This group of
species has been recognized at a variety of taxonomic
levels, most often as a distinct family (Newman, 1840;
Ching, 1940; Parris, 1990) or as a subfamily of the
Polypodiaceae (Lellinger, 1989; Tryon & Tryon, 1982).
Preliminary nucleotide sequence data from the rbcL gene
indicated that grammitids nest within Polypodiaceae

(Hasebe & al., 1995). That finding is strongly supported
by studies with more extensive taxonomic sampling
using evidence from three chloroplast DNA regions
(Schneider & al., 2004). In addition, the study of
Schneider & al. provides strong evidence for the monophyly of the whole grammitid group and for close relationships of grammitids and Neotropical taxa of
Polypodiaceae.
The circumscription and monophyly of the grammitid group have seldom been questioned, but there has
been little agreement on generic delimitation. For example, Parris (1990) recognized only four genera in the
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Fig. 1. Examples of endemic Hawaiian grammitids. A, Adenophorus hymenophylloides; B, Grammitis tenella; C, A.

tamariscinus; D, A. oahuensis. Photo credits: A, A. R. Smith; BD, T. A. Ranker.
family, whereas Copeland (1947) recognized 12, and

Parris (2003) recognized 18 genera and 14 species
groups. Smith (1993) recognized 10 genera of grammitids of primarily New World distribution, but only one
(Lomaphlebia J. Sm.) is endemic to the New World.
Murillo & Smith (2003) recently described a monotypic,
Neotropical endemic genus (Luisma). Smith (1993 and
references therein) used presence or absence of hydathodes (i.e., differentiated leaf epidermal cells at vein tips)
as one of the main characters separating groups of
Neotropical genera of grammitids. Anhydathodous genera are Ceradenia L. E. Bishop, Enterosora Baker, and
Zygophlebia L. E. Bishop, whereas all other Neotropical
416
genera have obvious adaxial hydathodes (with the exception of the austral, widespread species Grammitis poeppigiana, which lacks hydathodes, and G. patagonica,
which rarely has them). Paleotropical species also sometimes show this feature, but whether hydathodes are useful for circumscribing Old World species groups is still
uncertain. Additional evidence, both morphological and
molecular, is needed to confirm the value of this character (or any others) in distinguishing both Neotropical and
Paleotropical genera.
Toward this end, we conducted phylogenetic analyses of molecular and morphological data of a taxonomically and geographically broad sample of grammitids. In
53 (2) May 2004: 415428
this study, we pose the following systematic, phylogenetic, and biogeographic questions: Are the variously
recognized grammitid genera monophyletic? Are there
unrecognized monophyletic groups of species that merit
generic ranking? Do monophyletic groups segregate
along Old World-New World or other biogeographical
lines? Are there phylogenetically distinct lineages characterized by the presence or absence of hydathodes or
other distinctive morphological features? To address
these questions, we sampled sequences of two chloroplast genes, atpB and rbcL, compiled an extensive morphological dataset, and conducted phylogenetic analyses
of all datasets.
MATERIALS AND METHODS

Taxon sampling and DNA extraction.
Outgroup relationships of grammitids were estimated by
Schneider & al. (2004). Based on that analysis we chose
five polypodioids as outgroups for the present study.
Samples were either collected by one of us or were generously supplied by colleagues. We attempted to sample
ingroup species from as many recognized grammitid
genera as possible. Appendix 1 lists general locality, collector, collection number, herbarium of deposit for
voucher specimens, and GenBank accession numbers for
all taxa and DNA sequences studied (see internet version
of Taxon). More complete locality data and other information can be obtained from the following web site:
http://www.biology.duke.edu/pryerlab/ferndb/index.
html. Six species were represented by two accessions
each, which allowed us to assess intraspecific variation
of individual recognized species.
We extracted total cellular DNA using the CTAB
method of Doyle & Doyle (1987) modified by adding
3% PVP-40 and 5 mM ascorbic acid. Sample DNA concentrations were standardized to 10 mg/mL with the aid
of a minifluorometer.
PCR amplification and sequencing. We
PCR-amplified and sequenced two segments of the
cpDNA genome: a 1311 basepair (bp) fragment of the
rbcL gene, and a 1266 bp fragment of the atpB gene.
Methods followed those of Ranker & al. (2003).
Sequences of rbcL were obtained for all 79 samples of
grammitids and the five polypodioid outgroups.
Sequences of atpB were obtained for 77 samples of
grammitids and the five outgroup taxa.
Phylogenetic analyses of molecular data.
Sequence fragments were edited by visual inspection of
electropherograms in Sequencher (Gene Codes Corp.)
and consensus sequences were aligned manually in
PAUP* 4.0b10 (Swofford, 1998) with no inferred gaps.
We first conducted maximum parsimony (MP) phy-
logenetic analyses as implemented in PAUP* 4.0b10

(Swofford, 1998) on each set of DNA sequences separately. All characters were unordered and equally weighted. Invariant characters were omitted from all analyses.
For each single-gene dataset, we employed heuristic
searches with 1000 random addition sequence replicates
with MulTrees activated, TBR branch swapping, and
ACCTRAN character-state optimization. Bootstrap
analyses of single-gene datasets were conducted with
100 replicates, each with 10 random addition sequence
replicates. The topologies of the strict consensus trees
resulting from the two single-gene analyses were compared with Kishino-Hasegawa (KH) and ShimodairaHasegawa (SH) tests as implemented in PAUP* (Bull &
al., 1993; Cunningham, 1997; Goldman & al., 2000;
Schneider & al., 2004). Because the topologies did not
differ statistically for either test (KH, P = 0.791; SH, P =
0.393), we conducted analyses of a combined dataset. On
the combined dataset, we performed heuristic searches,
as above, but with 5000 random addition sequence replicates. We also conducted a decay analysis of branch support (Bremer, 1988; Donoghue & al., 1992) on the combined dataset with AutoDecay 4.0.1 (Eriksson, 1998). We
conducted a heuristic bootstrap analysis of the combined
dataset in PAUP* with 100 replicates, each with 10 random addition sequence replicates. We conducted another
heuristic analysis of the combined dataset, as above, but
with a transition:transversion (ti:tv) weighting scheme
that took into account the parsimony informativeness of
transitions vs. transversions in each gene separately. We
estimated ti:tv ratios by comparing the length of MP trees
with transitions omitted vs. with all variable sites included (see Martin & Naylor, 1997). For rbcL we found a
ti:tv ratio of 4:1, and for atpB we found a ratio of 8:1.
MODELTEST version 3.06 (Posada & Crandall,
1998) was employed to find the model of DNA substitution that best fit the data. Both hierarchical likelihood
ratio tests and the Akaike Information Criterion method
found that the TVM+G model was the best fit, and the
following settings were inserted in the PAUP* combined
dataset file: Lset Base = (0.2609 0.2262 0.1815) Nst = 6
Rmat = (1.4824 8.0219 0.7070 1.0769 8.0219) Rates =
gamma Shape = 3.1038 Pinvar = 0; with those settings,
we conducted a heuristic maximum likelihood (ML)
phylogenetic analysis in PAUP* without a molecular
clock constraint and with MulTrees activated and TBR
branch swapping. We also conducted a ML bootstrap
analysis with the same settings and with 100 repetitions
and 10 random addition sequence replicates each.
Clade credibility values were estimated for the combined dataset by calculating the posterior probability for
each node using Bayesian inference with a MarkovChain Monte Carlo (MCMC) sampling method as implemented in MrBayes 2.1 (Huelsenbeck & Bollback, 2001;
417
53 (2) May 2004: 415428
Huelsenbeck & Ronquist, 2001) and employing the

TVM+G model found by MODELTEST. A tree was sampled every 100 generations for 1,500,000 generations,
and each analysis was run with four simultaneous
MCMC chains. A majority-rule consensus tree was computed in PAUP* on the last 12,000 sampled trees, excluding the 3,000 trees found in the burn-in period, which
was estimated following the method of Schneider & al.
(2004). Posterior probability values above P = 95% were
considered to be statistically significant (Rannala &
Yang, 1996; Larget & Simon, 1999; Huelsenbeck &
Ronquist, 2001; Lewis, 2001).
Phylogenetic analyses of morphological
data. A subset of the study group comprising 69
species of grammitids (i.e., excluding four species of
Adenophorus, because a more detailed analysis of the
whole genus has been published elsewhere (Ranker &
al., 2003) and the five polypodioid outgroup species
were scored for 87 morphological and chromosomal
characters (Table 1). The data matrix is available in
Appendix 2 (internet version of Taxon). We conducted
maximum parsimony phylogenetic analyses in PAUP*
4.0b10 (Swofford, 1998) of the morphological dataset.
All characters were unordered and equally weighted. We
performed a heuristic search with 100 random addition
sequence replicates and with MulTrees activated, TBR
branch swapping, and ACCTRAN character-state optimization.
Phylogenetic analyses of combined molecular and morphological data. We also conducted a
heuristic analysis with the same conditions listed in the
previous section and a bootstrap analysis each with 1000
replicates of a dataset combining the two molecular
datasets and the morphological dataset.
Our list of morphological characters and character
states was composed from information in various monographic works, floristic accounts, and character lists for
genera of Polypodiaceae (e.g., Evans, 1969; Lellinger,
1972; Roos, 1985; Hovenkamp, 1986; Hensen, 1990;
Rdl-Linder, 1990, 1994a, 1994b; Bosman, 1991;
Hovenkamp & Franken, 1993; Zink 1993) and
Grammitidaceae (e.g., Copeland, 1952, 1956; Bishop,
1974, 1989; Parris, 1983, 1990). Studies of character
analysis in Polypodiaceae (e.g., Sen & Hennipman,
1981; Hetterscheid & Hennipman, 1984; Baayen &
Hennipman, 1987a, b), synoptical treatments of genera
of Grammitidaceae (e.g., Bishop, 1977, 1978, 1988;
Smith, 1992, 1993; Smith & Moran, 1992; Smith & al.,
1991) and more comprehensive works on ferns (e.g.,
Tryon & Tryon, 1982; Kubitzki, 1990) were also used as
sources of information about characters and character
states. In addition, we examined herbarium species of
each taxon in Appendix 1 in several herbaria, particularly AK and UC. Coding of the character states does not
imply a transformation series, i.e., the coding is neutral.

To explore the evolution of morphological traits, we
employed the strict consensus tree resulting from the
heuristic MP analysis of the combined rbcL-atpB. We
used MacClade 4.0 PPC (Maddison & Maddison, 2000)
to optimize character-state changes onto the strict consensus tree resulting from that MP analysis of the combined molecular dataset. [NB: using topologies of either
the Bayesian inference strict consensus or the ML tree
did not alter our interpretations of morphological evolution; results not shown.] We employed the all most parsimonious states at each node resolving option.
418
RESULTS
Molecular data and phylogenetic analyses.
For the 79 grammitid rbcL sequences obtained, 923
bp were invariant, 388 bp were variable, and 234 bp were

parsimony-informative. For the 77 ingroup sequences of
atpB obtained, 928 bp were invariant, 338 bp were variable, and 243 bp were parsimony-informative.
The MP analysis of the rbcL dataset found 8,021
equally parsimonious trees. A length (L) of 1152 steps,
consistency index (CI) of 0.45, and retention index (RI)
of 0.67 characterized each tree. The MP analysis of the
atpB dataset found 53,721 equally parsimonious trees
with L = 955, CI = 0.36, and RI = 0.69. The strict consensus trees from both analyses (not shown) were highly,
but not completely, dichotomously resolved. Only three
of the 12 genera (Adenophorus, Enterosora, and
Melpomene) from which we sampled two or more
species were resolved as monophyletic in both singlegene analyses; all other such genera were resolved as
monophyletic by only one analysis or by neither.
Bootstrap (BS) values supporting the monophyly of
Adenophorus, Enterosora, and Melpomene for rbcL were
77, 94, and 100, respectively, and for atpB were 66, 58,
and 100, respectively.
The MP analysis of the combined rbcL-atpB dataset
found 72 equally parsimonious trees (Fig. 2) with L =
2231, CI = 0.43, and RI = 0.68. Bootstrap (BS) support
varied across branches of the strict consensus tree, was
greater or equal to 95% at 34 of the 76 ingroup nodes,
and was greater than 70% at 55 ingroup nodes. The strict
consensus tree from the Bayesian analysis of the combined molecular dataset produced a nearly identical
topology to that from the MP analysis (Fig. 2). At 66 of
the 76 ingroup nodes, clade credibility (= posterior probability, PP) values were greater than 95%. Although the
strict consensus tree from our heuristic MP analysis
resolved Clades V and VI as sister taxa (see Fig. 2), there
was little BS or PP support for that relationship. We conducted a SH test (as implemented in PAUP*) to compare
53 (2) May 2004: 415428
Table 1. Morphological characters and character states used in the phylogenetic analysis of grammitid ferns. NA = Not Applicable.
HABIT: 1 growth habit: (0) terrestrial, (1) epigaeus (in moss layers on ground), (2) epilithic, (3) epiphytic, (4) rheophytic and/or rooted aquatic.
ROOTS: 2 root density: (0) = 5 roots/mm of rhizome length, as viewed from one side, (1) > 5 roots/mm; 3 hair density: (0) = 25 hairs/mm of
root length, as viewed from one side, (1) > 25 hairs/mm; 4 proliferous, producing new plantlets: (0) no, (1) yes; 5 insertion [NA for taxa with
radial rhizomes]: (0) dorsally and ventrally, (1) ventral side only.
RHIZOMES: 6 color: (0) brown (or green in vivo), (1) glaucous or whitish; 7 symmetry: (0) radial, (1) dorsiventral; 8 shape in cross-section:
(0) terete, (1) dorsiventrally flattened; 9 diameter: (0) < 2 mm, (1) 25 mm; 10 branching: (0) branched, (1) commonly unbranched; 11 vascular strands: (0) 1, (1) 2-9, (2) 10-20, (3) >20; 12 sclerenchyma/collenchyma sheaths around vascular bundles: (0) absent, (1) present; 13 isolated sclerenchyma strands in cortex and pith: (0) absent, (1) present; 14 sclerenchyma/collenchyma sheath in the cortex: (0) absent, (1)
present.
RHIZOME INDUMENT: 15 indumentum: (0) hairs only, (1) mainly scales; 16 scale color [en masse]: (0) orangish, (1) light brown/tan, (2)
red-brown to brown, (3) blackish (or gray to dark gray); 17 scale iridescence: (0) no, (1) yes; 18 scale sheen: (0) dull, (1) subglossy, (2) glossy;
19 scale cell turgidity (at midscale): (0) not turgid, (1) subturgid, (2) turgid; 20 scale attachment: (0) basifixed, (1) pseudopeltately attached,
(2) peltately attached; 21 scale base shape: (0) non-auriculate, (1) auriculate; 22 scale apex: (0) obtuse, (1) acute, (2) acuminate, (3) filiform; 23
scale shape: (0) round, (1) ovate [length = 2 width], (2) lanceolate, (3) linear [length 12 width]; 24 cell shape at mid-scale in scale center
[often but not always correlated with character 25]: (0) isodiametric, (1) length 12 width, (2) length > 2 width; 25 scale orientation: (0) tightly appressed, (1) slightly spreading, (2) strongly spreading; 26 scale clathrateness: (0) uniformly colored, (2) subclathrate, (3) clathrate throughout; 27 scale margins, mid-scale: (0) entire, (1) toothed, (2) setulose, hairs eglandular, unicellular, (3) glandular, glands unbranched, (4) setulose,
hairs eglandular, multicellular; 28 scale margins, apex: (0) entire, (2) setulose, hairs eglandular, unicellular, (3) glandular, glands unbranched, (4)
eglandular, hairs branched; 29 scale surfaces: (0) glabrous, (1) setulose, (2) glandular.
LEAVES: 30 blade termination: (0) determinate, (1) indeterminate, often with seasonal constrictions; 31 fertile-sterile leaf differentiation: (0)
(nearly) monomorphic, (1) hemidimorphic - leaf tip fertile (internal dimorphism); 32 blade dissection, at least at base: (0) simple (undivided),
(1) pinnatifid, (2) pinnatisect, (3) 1-pinnate, pinnae adnate, (4) pinnate-pinnatifid or more divided; 33 blade base: (0) tapered, (1) truncate; 34
blade apex: (0) gradually reduced w/confluent divisions, (1) conform to the pinnae; 35 blade tissue: (0) not spongiose, (1) spongiose;
36 sclerenchyma: (0) invisible on all parts of blade, (1) faintly visible, (2) prominently visible; 37 black blade margin: (0) absent, (1) present
38 hydathodes: (0) absent, (1) present; 39 lime dots from hydathodes: (0) absent, (1) present; 40 leaf articulation: (0) absent, (1) present; 41
pinna articulation: (0) absent, (1) present; 42 adaxial outline of stipe: (0) terete, (1) flattened, (2) sulcate with lateral ridges or flanges; 43 sclerenchyma coloration on stipe: (0) not dark-pigmented, (1) dark-pigmented; 44 stipe stele number: (0) monostele, (1) distele, (2) polystele; 45
phyllopodia: (0) absent, (1) present; 46 phyllopodia/stipe separation: (0) 02 rhizome width apart, (1) 25 rhizome width apart, (2) 510
rhizome width apart, (3) 1020 rhizome width apart; 47 blade with black clavate fungal fruiting bodies (Acrospermum maxonii): (0) absent,
(1) present; 48 blade with aromatic aroma (somewhat sweet and spicy) when dried: (0) absent, (1) present.
VENATION: 49 vein orders: (0) two, (1) three, (2) four or more; 50 secondary vein form: [NA if 2o vein are unbranched] (0) dichotomous,
(1) anisotomous/non-dichotomous; 51 vein fusion in sterile blades: (0) non-anastomosing, (1) anastomosing; 52 rows of vein areoles between
costa and margin: (0) 1-2, (1) 3-4, (2) 5-8; 53 vein prominence adaxially: (0) not prominent, (1) slightly prominent, (2) prominent; 54 vein
prominence abaxially [raised and/or darkened]: (0) not prominent, (1) slightly prominent, (2) prominent.
BLADE INDUMENT: 55 hairs: (0) absent, (1) present; 56 hair types: (0) uniseriate, unicellular, (1) uniseriate, multicellular, (2) branched;
57 branched hairs [soral paraphyses excluded]: (0) branches sessile-glandular (e;g;, Ceradenia); (1) branches hairlike, acicular, (2) branches capitate-glandular, (3) branches catenate; 58 hairs on costae and rachis adaxially: (0) absent, (1) acicular, (2) glandular, (3) branched; 59 hair color:
(0) hyaline or light yellowish or pale reddish, (1) dark red brown; 60 blade scales: (0) absent, (1) present; 61 blade scale color pattern: (0) uniformly colored, (1) sharply bicolored, (2) clathrate.
SORI AND INDUSIA: 62 soral receptacle: (0) (nearly) flat, (1) convex; 63 sporangial stalk width at mid-stalk: (0) 2-3 cells, (1) 1 cell; 64
sorus number per leaf: (0) >2, (1) 2 (coenosori); 65 sorus distribution: (0) throughout blade or only on proximal part, (1) confined to distal portion of blade; 66 mature sorus outline: (0) round or slightly oblong, (1) elongate, 210 longer than wide, (2) > 10 longer than wide; 67 sorus
depth: (0) surficial, (1) sunken with sloping sides, (2) sunken with vertical sides; 68 sori embossed on adaxial surface: (0) no, (1) slightly, (2)
yes, decidedly; 69 soral pit rims: (0) entirely from abaxial surface, (1) in part by blade margin; 70 soral position [applicable only when sori are
uniseriate between axis and margin]: (0) nearest to penultimate axis, (1) midway between axis and margin, (2) nearer to margin than to penultimate axis; 71 sorus presence in marginal areoles (or on their bordering veins): (0) absent, (1) sometimes present, (2) always present; 72 sorus
presence in costal areoles (or bordering veins): (0) absent, (1) sometimes present, (2) always present; 73 paraphyses from receptacle or disproportionately from vicinity of sorus: (0) absent, (1) present; 74 hairlike paraphyses [excluding paraphyses from sporangial walls or capsules]: (0) eglandular, (1) glandular; 75 episporangial paraphyses: (0) absent, (2) hairlike; 76 position of episporangial paraphyses: (0) from
stalk, (1) from side walls of capsule, (2) from annulus; 77 localization of episporangial paraphyses at distal end: (1) scattered, (2) aggregated
[as in OW Grammitis]; 78 blade margin infolded over sori: (0) no, (1) yes.
SPORES: 79 color: (0) whitish, (1) yellowish, (2) brownish, (3) greenish (chlorophyllous); 80 spore number: (0) 64, (1) 32, (2) 16; 81 spore
laesura: (0) linear, (1) triradiate; 82 number of cells in newly shed spores: (0) one, (1) two, (2) three to four; 83 perispore (epispore) surface:
(0) (nearly) smooth or plain, (1) obviously patterned or sculptured; 84 exospore (exine) surface: (0) (nearly smooth or plain, (1) obviously patterned or sculptured.
GAMETOPHYTES: 85 form: (0) cordate-thalloid, (1) unknown, (2) elongate-thalloid to filamentous; 86 rhizoid developed in the first cell
division: (0) no, (1) yes.
CHROMOSOMES: 87 base number: (0) 37, (1) 35, (2) 33, (3) 32.
419
100
54 100
85
d7
<50
85
67
100
53 (2) May 2004: 415428
<50
100
91
100
d2
d2
d3
57
95
<50
98
d3
100
100
100
100
d13
<50
99
d1
<50
91
98
100
d1
95
100
78
100
d2
d2
d1
d1
d1
d19
53
97
63
98
<50
96
71
100
98
100
d7
86
d4 100
d2
d3 100
100
d2
d8
100
95 100
100 d21
92
100
d6
100
91
100
d5 100
d5
100
100
d33
d15
65
99
57
99
100
100
d2
d1
<50
72
92
d22 100
d1
d7
81
100
ML
94
<50 100 d2
65 62 d3
91
96
100
91
100
88
100
d4
d1
d6
99
52 100
99
d4
100
100
d13
d2
100
100
100
100
d16
d9
66
99
100
100
99
100
d1
d10
d11
100
100
91
100
d22
75
100 100
100 d2
84
100 d14
d3
100
100
100
d3 100
99
100
d9
d11
d7
100
100
d4
100
d15 100
96
100
61
81
100
100
d13
d1
95
100
d13
79
100
98
100
100
100
d33
96
100
d5
d3
d8
99
100
57
94
d6
d1
54
94
d1
d7
100
100
51
d16
100
100
93
100
d10
84 65
100
d4
63
100
d2
d7
100
100
100
100
d18
Grammitis hookeri
Grammitis forbesiana
Grammitis knutsfordiana
Xiphopteris conjunctisora
Themelium tenuisectum
Grammitis parva
Grammitis hirtelloides
Prosaptia pubipes
Prosaptia palauensis
Prosaptia contigua
Ctenopteris rhodocarpa
Prosaptia obliquata
Prosaptia alata
Ctenopteris nutans
Grammitis ciliata
Grammitis pseudociliata
Grammitis billiardieri
Grammitis poeppigiana AU
Grammitis poeppigiana CH
Ctenopteris heterophylla
Grammitis deplanchei
Ctenopteris lasiostipes
Ctenopteris aff. repandula
Ctenopteris repandula
Scleroglossum sulcatum
Calymmodon luerssenianus
Calymmodon gracilis
Micropolypodium taenifolium
Micropolypodium zurquinum
Terpsichore achilleifolia
Terpsichore longisetosa
Micropolypodium hyalinum
Chrysogrammitis musgraviana
Lellingeria apiculata
Lellingeria subsessilis
Lellingeria hirsuta
Lellingeria limula
Melpomene moniliformis
Melpomene pseudonutans
Melpomene flabelliformis
Terpsichore subscabra
Terpsichore sp.
Terpsichore alsopteris
Terpsichore semihirsuta
Terpsichore subtilis
Terpsichore pichinchae
Terpsichore anfractuosa
Ceradenia spixiana
Ceradenia kalbreyeri
Ceradenia aulaeifolia
Ceradenia pilipes
Ceradenia jungermannioides
Enterosora percrassa
Enterosora trifurcata
Terpsichore cultrata
Terpsichore senilis
Terpsichore lanigera
Lellingeria schenckei
Lellingeria pseudomitchellae
Adenophorus haalilioanus
Adenophorus pinnatifidus
Adenophorus tamariscinus
Adenophorus hymenophylloides
Adenophorus tripinnatifidus
Adenophorus periens
Grammitis tenella
Cochlidium punctatum
Cochlidium seminudum
Cochlidium rostratum
Grammitis bryophila
Grammitis melanoloma
Terpsichore eggersii
Terpsichore hanekeana
Terpsichore lehmanniana
Ia
Ib
Ic
II
III
IVa
IVb
Va
Vb
VI
VII
VIII
IX
Polypodium glycyrrhiza
Pecluma alfredii763R
Campyloneurum augustifolium
Microgramma squamulosa
Microgramma percussa
Fig. 2. Majority-rule consensus tree from the Bayesian analysis summarizing analyses of the rbcL + atpB dataset. Upper
number above each branch is a bootstrap value; lower number is a posterior probability (%). Decay indices are below
branches. ML indicates the position of Chrysogrammitis musgraviana supported by the maximum likelihood analysis.
Roman numerals to the right indicate clades as discussed in the text.
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53 (2) May 2004: 415428
the relative likelihood of two tree topologies: VVI unresolved (as in Fig. 2) vs. V sister to VI. The results found
that the likelihoods of the two trees were not significantly different (P = 0.697). The ML heuristic analysis produced a tree with a nearly identical topology to that from
the Bayesian analysis, with the exception of the placement of Chrysogrammitis musgraviana (see Fig. 2).
Recognized genera from which we sampled two or more
species and that were supported as being monophyletic
in the parsimony bootstrap (BS), Bayesian inference
(PP), and ML bootstrap analyses (MLBS) (decay index
values, d, also shown) were Adenophorus (BS 98; PP
100; MLBS 94; d5), Ceradenia (BS 84; PP 100; MLBS
87; d3), Calymmodon (BS 100; PP 100; MLBS 100;
d15), Cochlidium (BS 85; PP 100; MLBS 84; d4),
Enterosora (BS 99; PP 100; MLBS 99; d7), and
Melpomene (BS 100; PP 100; MLBS 100; d13). Other
recognized genera from which we sampled two or more
species and which were not supported as monophyletic
included Ctenopteris, Grammitis, Lellingeria, Micropolypodium, Prosaptia, and Terpsichore.
The MP heuristic analysis of the combined dataset
employing ti:tv differential weighting resulted in 24 most
parsimonious trees with L = 4402, CI = 0.53, and RI =
0.70 (not shown). Although the CI and RI values were
greater using the weighted dataset than using the
unweighted, a SH test comparing the topologies of the
strict consensus trees from the unweighted vs. the
weighted analysis showed that they did not differ statistically (P = 0.263).
Morphological data and phylogenetic analyses. Of the 87 variable morphological characters for
the whole dataset, 66 were variable and 58 were parsimony-informative among the ingroup species. Heuristic
MP analysis of the morphological dataset alone found
1,681 equally parsimonious trees, with L = 413, CI =
0.28, and RI = 0.65. The strict consensus tree showed
very little dichotomous resolution of relationships (not
shown).
Analysis of combined morphological and
molecular datasets. Heuristic MP analysis of the
combined morphological and molecular datasets found
112 equally parsimonious trees with L = 2445, CI = 0.33,
and RI = 0.64. The topology of the strict consensus tree
(not shown), from which we had a more limited sampling, was similar to that based only on molecular data
(Fig. 2). Bootstrap support was variable across clades in
the strict consensus tree from the combined analysis. The
same clades as numbered in Fig. 2 were recovered in the
combined analysis of morphological and molecular data,
although relationships of species within clades were
sometimes different. The primary differences in the
arrangement of clades in the combined morphology
+molecule analysis relative to the rbcL-atpB analysis
were: (1) Clade Ic was sister to Ib, rather than to Ia + Ib;

(2) the relationships of Clades IIV were unresolved as a
polytomy; and (3) the placement of Ctenopteris
lasiostipes was unresolved in that polytomy, rather than
as a member of Clade II.
DISCUSSION
Taxonomic implications: putatively monophyletic genera. Several recognized genera were
strongly supported as being monophyletic, including

Adenophorus, Ceradenia, Calymmodon, Cochlidium,
Enterosora, and Melpomene. However, sampling in several of these genera is still insufficiently broad, or does
not yet include species of more problematic affinity. Our
analysis of the distribution of morphological character
states on the molecular-based phylogenetic hypotheses
identified synapomorphies for each genus.
Adenophorus is a genus of 10 species endemic to the
Hawaiian Islands and is discussed in greater detail in
Ranker & al. (2003).
Ceradenia is primarily a Neotropical genus of about
60 spp. (Bishop, 1988; Parris, 2002), with two spp. also
found in the Azores and Liberia (Smith, 1995) and
another six in Africa including Madagascar (Parris,
2002). The only known morphological feature distinguishing this genus from other groups is a peculiar and
unique type of glandular paraphysis found in all species.
Each paraphysis is multicellular and uniseriate, with 25
distal cells producing sessile, white, wax-like globose or
pyriform glands. These paraphyses correspond to the
group 9 paraphyses of Parris (1997a) and the type II4a of Baayen & Hennipman (1987a; 1987b). In our
morphological dataset, species were scored as either having or lacking glandular paraphyses, and the character
appeared to show considerable homoplasy in our mapping of the character states onto the molecular-based
phylogeny (CI for the character = 0.25). Otherwise, the
genus is polymorphic for many different features, including leaf blade dissection and various rhizome scale characters.
Calymmodon is a genus of about 30 species primarily from Southeast Asia and the Indo-Pacific region
(Copeland, 1927; Parris, 1998b). Species in this group
are unique among grammitids in having the basiscopic
margins of each pinna infolded over a single sorus (see
fig. 148 H, I of Parris, 1998b).
Cochlidium includes 16 species of primarily
Neotropical distribution, with one species (C. serrulatum) also ranging to Africa and islands of the Indian
Ocean (Bishop, 1995). Species of Cochlidium are unique
in having hemi-dimorphic leaves (i.e., blade tip fertile).
Gametic chromosome numbers of n = 37 have been
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53 (2) May 2004: 415428
reported for most groups of grammitids; however, n = 33

has been reported for Cochlidium rostratum (Wagner,
1980) and for Lellingeria hartii (Walker, 1985). Further,
the Cochlidium clade is polymorphic for gametic chromosome number, with n = 35 having been reported for C.
serrulatum (Walker, 1985). Another feature of the genus
is a high frequency of coenosori, formed from an apparent developmental fusion of independent sori. Coenosori
apparently evolved independently in Scleroglossum.
Enterosora contains eight Neotropical species and
two African species (Bishop & Smith, 1992; Parris,
2002). The only morphological feature that seems to be
unique to this group is the common occurrence of anastomosing veins in sterile leaves. Leaf blades also are usually prominently spongiose in texture, although this
feature occurs independently in a less developed form in
Cochlidium rostratum, in species of Ceradenia, and
probably in other grammitids. A noticeably dull sheen on
rhizome scales is also found in this group, a feature separating it from its putative sister clade, Ceradenia, but
that state is also found in species of the CochlidiumGrammitis bryophila clade.
The only other named genus that was supported by
our data as being monophyletic was Melpomene. Most of
the approximately 20 species are entirely Neotropical,
but at least one (M. flabelliformis) occurs also in Africa,
Madagascar, and La Runion (Smith & Moran, 1992).
Although species in this group share a suite of character
states such as clathrate rhizome scales, basally cordate
rhizome scales, and rhizome scales with apical papillae,
we are not aware of a single morphological trait that is
unique to this group. The only single potentially unique
feature that we could identify is an undescribed chemical
or group of chemicals that produces a sweet-smelling
aroma in dried leaves (Smith & Moran, 1992).
Taxonomic implications: previously unrecognized groups. Recognized genera from which we
sampled two or more species and which were not supported as monophyletic included Ctenopteris,
Grammitis, Lellingeria, Micropolypodium, Prosaptia,
and Terpsichore. Other than the monophyletic genera
discussed above, several strongly supported clades were
found that comprise various combinations of species
from two or more genera. Each clade is also defined at
various levels by combinations of morphological characters, most of which have been independently derived
multiple times on the phylogeny. We have numerically
labeled clades in Fig. 2 for the sake of discussion.
Clades IIII consist of species that are primarily Old
World in distribution, with a few species that are essentially circumaustral (e.g., Grammitis poeppigiana).
Clade I (Fig. 1) was supported with BS 100 and PP
100 and a decay index of 13. All species in this clade are
from a broad region of the SW Pacific/E Asia or the
Hawaiian Islands. Clade I includes three smaller, mostly

well supported clades. The single morphological feature
that appears to be a synapomorphy for Clade I is the presence of an obvious petiolar abscission layer (leaf articulation), found in Ctenopteris nutans (Ic) and Clade Ib.
We must infer, however, that this feature was lost in the
ancestor of Clade Ia. Clade Ia (BS 65, PP 100, d3)
includes some species of the highly polyphyletic
Grammitis, Xiphopteris, and Themelium. Themelium spp.
have highly dissected leaves (Parris, 1997b), much more
so than other species in this clade. Highly dissected
leaves have evolved independently in Hawaiian
Adenophorus, as well as in the two Terpsichore species
in Clade IV. Clade Ib consists primarily of species of
Prosaptia, plus Ctenopteris rhodocarpa. Perhaps the
most prominent morphological features of Prosaptia
species are that sori are typically embedded either in pits
in the leaf lamina or in marginal pouches and the leaves
have distinctive articulation. Although shallow soral pits
are present in species typically treated in Ctenopteris and
several other unrelated grammitid genera, among the
species of Ctenopteris in our analysis only C.
rhodocarpa and C. nutans share the definitive leaf articulation with Prosaptia.
Clade II (Fig. 2; BS 81, PP 100, d2) consists of a
group of austral species ranging across Australia, New
Caledonia, New Zealand, and southern South America.
Species in this clade have been treated in Grammitis and
Ctenopteris. This is a difficult group to delimit morphologically, and we can find no single trait that uniquely
defines the clade. There is rampant polymorphism and
homoplastic evolution across species for many characters. A more detailed study of this group is warranted.
Clade III (Fig. 2; BS 91, PP 100, d5) consists of three
highly distinct and separate lineages that, though cladistically close, are each well supported. Decay indices for
the branches supporting the three clades of Ctenopteris
spp., Scleroglossum sulcatum, and Calymmodon spp.
were d21, d33, and d15, respectively. Ctenopteris is a
large group of about 100 primarily Old World species
(Parris, 1998b). The six species included in our molecular dataset were hypothesized to belong to at least four
different phylogenetic lineages (Fig. 2). This group clearly needs further study, along with species of Prosaptia
and Grammitis, to identify lineages and relationships.
Scleroglossum is a small group of only about eight
species (Parris, 1998b) from the SE Asian/SW Pacific
region. Further sampling is necessary to determine if this
is a monophyletic genus. As discussed above,
Calymmodon is strongly supported as a monophyletic
genus in our study, but further sampling is required to
test this hypothesis.
Clades IVX consist predominantly of New World
species, but with a few oceanic island groups and a few
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53 (2) May 2004: 415428
isolated Old World species that appear to be outliers of

largely New World groups.
Clade IV includes species treated in the genera
Micropolypodium, Terpsichore, and Chrysogrammitis.
Species of the first two genera are primarily New World
in distribution, whereas the only two described species of
Chrysogrammitis are from SE Asia through Malesia to
Melanesia (Parris, 1998a). Clade IVa (Fig. 2) includes all
species in Clade IV excluding Chrysogrammitis musgraviana and was strongly supported (BS 100, PP 100, d22).
Micropolypodium consists of about 30 species, primarily
of New World distribution, with a few species occurring
in E Asia and the SW Pacific region (Smith, 1992). All
three species of Micropolypodium sampled for the present study were placed in Clade IVa, but the genus is rendered paraphyletic by the inclusion of two species of
Terpsichore. In describing Terpsichore (ca. 50 spp.),
Smith (1993) recognized five informal, morphologically
coherent groups. The two species of Terpsichore in our
clade IVa were placed by Smith (1993) in his group 5,
along with one other species, T. gradata (Baker) A. R.
Sm. All three of these species are aberrant from other
species previously included in Terpsichore by their light
orangish to light tan rhizome scales, similar to those
found in many species of Micropolypodium and unlike
the much darker scales in other groups of Terpsichore. A
notable feature of Micropolypodium species is that sori
are at the base of pinnules, with the margins of mature
sori touching or overlapping the leaf midrib (see fig. 1 of
Smith, 1992). Species of Smiths group 5 of Terpsichore
also share this feature (see fig. 1E of Smith, 1993); soral
placement in other species of Terpsichore is generally
further away from the midrib (see figs. 1AD of Smith,
1993). It now seems likely that Terpsichore group 5 of
Smith (1993) is misplaced, belonging instead near to or
part of the clade containing New World species of
Micropolypodium; this is yet another example in the
grammitids of blade dissection being a false indicator of
affinities.
The placement of Chrysogrammitis musgraviana is
one of the most poorly supported aspects of our analyses.
Bootstrap support of the sister-taxon relationship of C.
musgraviana and Clade IVa was less than 50%, Bayesian
posterior probability for that branch was only 72%, and
the decay index was only d1. Additionally, maximum
likelihood analysis placed C. musgraviana as sister to a
large clade containing all of Clades IIVa (see dashed
line in Fig. 2) and, in the strict consensus tree resulting
from the MP heuristic search, the placement of C. musgraviana was unresolved as one branch of a trichotomy.
Because of the similarity of the glandular paraphyses and
other laminar hairs of Chrysogrammitis to species of the
Hawaiian Adenophorus, Parris (1997a, 1998a) suggested
a possible close relationship between these two genera.
Although our results poorly resolve the sister-taxon relationship of Chrysogrammitis, they clearly do not support
a close affinity of this genus with Adenophorus. We constructed a tree topology identical to that in Fig. 2 except
that C. musgraviana was forced as the sister taxon to the
Adenophorus clade. The resulting tree was 45 steps
longer than the original and its log-likelihood was significantly less (KH test, p = 0.000). It is notable that species
of Chrysogrammitis lack hydathodes in contrast to other
members of Clade IV. Additional data are needed to
resolve the exact relationships of Chrysogrammitis.
We have delineated Clade V (Fig. 2; BS 65, PP 99,
d1) to include the monophyletic Melpomene and several
other clades that include species of the polyphyletic genera Lellingeria and Terpsichore. All species of this clade
have pronounced hydathodes and all are Neotropical,
with the exception of a few outlying (and as yet unsampled) species in Africa and oceanic islands. As with
Clade II, this also is a difficult group to delimit morphologically because of high levels of polymorphism and
homoplastic evolution of many characters. Both
Melpomene and Lellingeria species in this clade (Clade
Va) have clathrate rhizome scales. Such scales, however,
appear to have arisen independently in several grammitid
lineages. A detailed study of the development and/or
mature morphology of clathrate scales across diverse
grammitid lineages might reveal subtle differences
across clades. An unexpected potential synapomorphy
for the members of Clade V is that they share the state
ventral root insertion on rhizomes (vs. dorsal and ventral, or radial, insertion; Fig. 4D). That character state,
however, is shared with taxa in Clades VI and X. The
relationship between Clades V and VI was unresolved in
most of our analyses. Thus, ventral root insertion could
have arisen separately in each clade, or in a common
ancestor, if Clades V and VI are really sister taxa.
An interesting feature that appears to have arisen
uniquely in the two basal-most lineages within Clade V
is an apparent propensity to being infected by the endophytic fungus Acrospermum maxonii Farlow (Smith,
1993) in some species of Terpsichore (Fig. 4E). Smith
(1993) observed that specimens placed in his informal
groups 2 and 4 of Terpsichore are commonly infected by
this fungus, which is manifested by the presence of
black, club-shaped fruiting bodies on leaf rachises (see
fig. 1 of Smith, 1993), costae, and sometimes sori. Smith
& Moran (1992) noted that only one species of
Melpomene (M. anfractuosa; transferred to Terpsichore
by Len & Smith, 2003) was regularly infected with this
same fungus, an occurrence which essentially predicted the placement of M. anfractuosa among species of
Terpsichore in our phylogenetic analysis (Fig. 2).
Clade VI (Fig. 2; BS 100, PP 100, d9) comprises the
monophyletic, putatively sister genera Ceradenia and
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53 (2) May 2004: 415428
Grammitis hookeri
Grammitis knudsfordiana
Grammitis parva
Prosaptia pubipes
Prosaptia contigua
Prosaptia obliquata
Prosaptia alata
Ctenopteris nutans
Grammitis billardieri
Grammitis ciliata
Grammitis poeppigiana
Calymmodon gracilis
Terpsichore longisetosum
Lellingeria hirsuta
Lellingeria limula
Terpsichore sp.
Ceradenia spixiana
Ceradenia pilipes
Terpsichore senilis
Grammitis tenella
Grammitis bryophila
Campyloneuron angustifolium
Pecluma alfredii
radial
dorsiventral
polymorphic
equivocal
orangish
absent
hairlike
absent
light brown to tan

red-brown to brown
blackish or gray
scalelike
polymorphic
polymorphic
polymorphic
equivocal
present
equivocal
equivocal
simple
pinnatifid
pinnatisect
1-pinnate, pinnae adnate
pinnate-pinnatifid or more divided
polymorphic
equivocal
Fig. 3. Examples of characters exhibiting high levels of homoplasy mapped onto the strict consensus tree resulting
from the heuristic maximum parsimony analysis of the combined rbcL-atpB dataset. A = rhizome symmetry; B = rhizome scale color; C = paraphyses; D = hydathodes; E = leaf dissection.
Enterosora. These are two of the three primarily New

World genera whose members lack obvious hydathodes
(Fig. 2D). We were unable to obtain molecular data from
the third New World anhydathodous genus, Zygophlebia,
a small genus of ca. 14 species (Bishop, 1989; Parris,
2002), but expect it will also fall within Clade VI, on the
basis of several additional morphological characters.
Clade VII (Fig. 2; BS 95, PP 100, d5) includes two
well-separated clades, one with species of Terpsichore,
the other with species of Lellingeria. Taxa in this clade
share apical and marginal rhizome scale hairs that are
424
unicellular, setulose, and eglandular. Similar hairs appear

to have arisen independently in various grammitid lineages, but do not occur in clades adjacent to Clade VII.
Smith (1993) placed the species of Terpsichore in Clade
VII in his informal group 3. They share the presence of
setose, episporangial paraphyses that are scattered on the
capsule wall adjacent to and for much of the length of the
sporangial annulus. Similar paraphyses are also found in
the Terpsichore species in Clade X. Although setose,
episporangial paraphyses are found in some species in
Clade Ia, the paraphyses in that clade are clustered near
53 (2) May 2004: 415428
Grammitis hookeri
Grammitis knudsfordiana
Grammitis parva
Prosaptia pubipes
Prosaptia contigua
Prosaptia obliquata
Prosaptia alata
Ctenopteris nutans
Grammitis billardieri
Grammitis ciliata
Calymmodon gracilis
Terpsichore longisetosum
Lellingeria hirsuta
Lellingeria limula
Terpsichore sp.
Ceradenia spixiana
Ceradenia pilipes
Terpsichore senilis
Grammitis tenella
Grammitis bryophila
Campyloneuron angustifolium
Pecluma alfredii
absent
dull
absent
present
subglossy
glossy
present
dorsally & ventrally

ventrally only
equivocal
absent
present
equivocal
polymorphic
equivocal
Fig. 4. Examples of characters exhibiting low levels of homoplasy mapped onto the strict consensus tree resulting from
the heuristic maximum parsimony analysis of the combined rbcL-atpB dataset. A = aromatic blade; B = rhizome scale
sheen; C = leaf articulation; D = root insertion; E = black clavate fungi.
the distal end of the annulus. Terpsichore species in

Clade VII appear to be unique among all grammitids in
being able to produce both monolete and trilete spores,
sometimes even on the same plant (Smith, 1993); by contrast, only trilete spores have been observed in other
grammitids. Terpsichore species in this clade also share
the unique feature of having leaf blade hairs that are
paired or clustered (e.g., see fig. 2B of Smith, 1993). The
two species of Lellingeria in this clade are representative
of a small group of about four species that are aberrant
from all other species of Lellingeria in having numerous
long, tawny hairs that bear an inconspicuous lateral cell
at the base (see fig. 1F of Smith & al., 1991). From the
molecular sequence data reported herein, it appears that
these species are taxonomically misplaced.
Clade VIII (Fig. 2; BS 96, PP 100, d7) includes the
genus Adenophorus and Grammitis tenella. All species
in this clade are endemic to the Hawaiian Islands. Ranker
& al. (2003) conducted a detailed molecular phylogenetic analysis of this group and we will not discuss it further
here.
Clade IX (Fig. 2; BS 100, PP 100, d10) is a strongly
supported group including the monophyletic
Cochlidium, which is sister to a pair of Grammitis
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53 (2) May 2004: 415428
species. These species of Grammitis both have distinctive sclerified, black leaf margins characteristic of the
type of the genus [G. marginella (Sw.) Sw.]. This feature
is unique to a group of about 22 taxa, comprising both
New and Old World species that have been treated as
Grammitis sect. Grammitis (Bishop, 1977) and as
Grammitis subg. Melanoloma (Copeland, 1952). All taxa
in Clade IX have simple blades, a feature that has arisen
multiple times in grammitid phylogeny (Fig. 3E). Other
character states shared by all members of Clade IX but
that arise convergently in multiple lineages across the
grammitid phylogeny include: radial rhizome symmetry,
the lack of adaxial hairs on leaf rachises and costae, sori
confined to distal portion of blade, and the absence of
soral paraphyses.
Clade X (Fig. 2; BS 100, PP 100, d23) is sister to all
other grammitids in our molecular phylogenetic analyses
and includes the species of Terpsichore treated by Smith
(1993) in his informal Group 1. As with several other
clades, we could find no single character state that
uniquely defines this group, with synapomorphic traits
for this clade arising independently elsewhere on the
tree. Members of the group do have scattered episporangial paraphyses that arise from sidewalls of sporangia
adjacent to the annulus, but similar paraphyses appear to
have evolved independently in the Terpsichore species of
Clade VII.
Inferred evolution of morphological characters. The most obvious and, arguably, the most striking feature of our inferred patterns of morphological evolution in grammitid ferns is the extremely high levels of
homoplastic evolution for many different characters.
Overall, homoplasy for morphological characters was
considerably greater than for molecular characters. For
example, in the hypothetical phylogeny (Fig. 2) the CI
values for the morphological and molecular datasets
were 0.24 and 0.47, respectively, and the RI values were
0.48 and 0.64, respectively. Many of the characters that
exhibited high levels of convergent or parallel evolution
across the phylogeny are features that have been commonly used in fern genus-level classifications (Fig. 3;
e.g., leaf blade dissection, various rhizome scale characters, and glandular paraphyses). Conversely, some of the
characters that exhibited relatively low levels of homoplasy have either not been regarded as having taxonomic
value or simply have not been discussed (Fig. 4; e.g., root
insertion, rhizome scale sheen). Of course, some characters that have been used to help define some genera were
found here to be of high taxonomic value and to exhibit
little to no homoplasy (e.g., the black sclerified leaf margins in Grammitis sect. Grammitis, the infolded basiscopic pinnae blade margins in Calymmodon, and the
multicellular, wax-like paraphyses of Ceradenia). The
presence or absence of obvious hydathodes, although
broadly useful among Neotropical taxa to distinguish

large clades, appears to be less useful among
Paleotropical lineages because of higher levels of homoplasy (Fig. 3D). This may be a reflection, however, of our
less intense sampling of Old World species.
Historical biogeographical implications. In
their molecular phylogeny of Polypodiaceae and several
representative grammitids, Schneider & al. (2004) found
that the monophyletic grammitid clade was most closely
related to several New World lineages of polypodioids.
Although their data and analyses could not identify the
exact extant sister clade of grammitids, they clearly support a hypothesis that the grammitid lineage arose in the
New World. In our analyses, the basalmost lineages,
comprising Clades IV to X, contain primarily New World
species. Conversely, the more derived clades (IIII) contain mostly Old World or Pacific species. Thus, we
hypothesize that there was one dispersal and colonizing
event involving the hypothetical common ancestor of
Clades IIII from the Neotropics to the Paleotropics or
Oceania (Fig. 2). Among the taxa in our study, we can
also hypothesize two separate dispersal events to the
Hawaiian Islands: one from the SW Pacific giving rise to
the Hawaiian Grammitis species in Clade Ia and the
other possibly from the Neotropics giving rise to the
Adenophorus clade (VIII). A third dispersal event
undoubtedly brought Lellingeria saffordii or its ancestor
to the Hawaiian Islands, however, we did not sample this
species. As noted in our discussion, there are several
groups that are either primarily Neotropical or primarily
Paleotropical but that also have a few members distributed in the opposite hemisphere. Thus, hypotheses of
multiple, independent dispersal and colonization events
in various lineages are warranted. Data from many more
taxa, especially Paleotropical, are needed to test these
hypotheses more rigorously.
General conclusions. Our results suggest to us
that some named genera of Grammitidaceae are clearly
monophyletic and should continue to be recognized taxonomically, whereas some are clearly para- or polyphyletic. Many of the previously unrecognized clades are
so strongly supported by our data that we feel that some
taxonomic revisions are warranted. It is also clear, however, that many more taxa should be sampled, especially
in the Old World, to represent adequately the multitude
of diverse lineages in this large group of ferns. We prefer
to await results of additional studies before making revisionary changes.
426
ACKNOWLEDGEMENTS
This research was supported by a collaborative grant from the
National Science Foundation to TAR (DEB-9726607), ARS
53 (2) May 2004: 415428
(DEB-9807053), and CHH (DEB-9807054). We thank the following colleagues for graciously collecting material: Wen-Liang
Chiou, Cynthia Dassler, Don Farrar, John Game, Chrissen
Gemmill, Steven Hill, Don Hodel, Michael Kessler, Blanca Len,
Lynn Raulerson, Alexander Rojas, Rene Rondeau, Alexandre
Salino, Patricia Snchez-Baracaldo, Carl Taylor, Henk van der
Werff, and Ken Wilson.
LITERATURE CITED
Baayen, R. P. & Hennipman, E. 1987a. The paraphyses of the
Polypodiaceae (Filicales). 1. General part. Beit. Biol.
Pflanzen 62: 251316.
Baayen, R. P. & Hennipman, E. 1987b. The paraphyses of the
Polypodiaceae (Filicales). 2. Taxonomic part. Beit. Biol.
Pflanzen 62: 317347.
Bishop, L. E. 1974. Revision of the genus Adenophorus
(Grammitidaceae). Brittonia 26: 217240.
Bishop, L. E. 1977. The American species of Grammitis sect.
Grammitis. Amer. Fern J. 67: 101106.
Bishop, L. E. 1978. Revision of the genus Cochlidium
(Grammitidaceae). Amer. Fern J. 68: 7694.
Bishop, L. E. 1988. Ceradenia, a new genus of
Grammitidaceae. Amer. Fern J. 78: 15.
Bishop, L. E. 1989. Zygophlebia, a new genus of
Grammitidaceae. Amer. Fern J. 79: 103118.
Bishop, L. E. 1995. Cochlidium. Pp. 371372 in: Davidse, G.,
Sousa, M. S. & Knapp, S. (eds.), Flora Mesoamericana,
vol. 1. Universidad Nacional Autnoma de Mxico,
Mxico, D.F.
Bishop, L. E. & Smith, A. R. 1992. Revision of the fern genus
Enterosora (Grammitidaceae) in the New World. Syst.
Bot. 17: 345362.
Bosman, M. T. M. 1991. A monograph of the fern genus
Microsorum (Polypodiaceae). Leiden Bot. Ser. 14: 1161.
Bremer, K. 1988. The limits of amino acid sequence data in
angiosperm phylogenetic reconstruction. Evolution 42:
795803.
Bull, J. J., Huelsenbeck, J. P., Cunningham, C. W.,
Swofford, D. L. & Wadell, P. J. 1993. Partitioning and
combining data in phylogenetic analysis. Syst. Biol. 42:
38397.
Ching, R. C. 1940. On natural classification of the family
Polypodiaceae. Sunyatsenia 5: 201268.
Copeland, E. B. 1927. The genus Calymmodon. Philipp. J. Sci.
34: 259269.
Copeland, E. B. 1947. Genera Filicum. Chronica Botanica
Co., Waltham, Mass.
Copeland, E. B. 1952. Grammitis. Philipp. J. Sci. 80: 93276.
Copeland, E. B. 1956. Ctenopteris in America. Philipp. J. Sci.
84: 381475.
Cunningham, C. W. 1997. Can three incongruence tests predict when data should be combined? Molec. Biol. Evol. 14:
733740.
Donoghue, M. J., Olmstead, R. G., Smith, J. F. & Palmer, J.
D. 1992. Phylogenetic relationships of Dipsacales based
on rbcL sequences. Ann. Missouri Bot. Gard. 79: 333345.
Doyle, J. J. & Doyle, J. L. 1987. A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochem.
Bull. 19: 1115.

Eriksson, T. 1998. AutoDecay ver. 4.0.1 (program distributed
by the author). Department of Botany, Stockholm Univ.,
Stockholm.
Evans, A. M. 1969. Interspecific relationships in the
Polypodium pectinatum-plumula complex. Ann. Missouri
Bot. Gard. 55: 193293.
Goldman, N., Anderson, J. P. & Rodrigo, A. G. 2000.
Likelihood-based tests of topologies in phylogenetics.
Syst. Biol. 49: 652670.
Hasebe, M., Wolf, P. G., Pryer, K. M., Ueda, K., Ito, M.,
Sano, R., Gastony, G. J., Yokoyama, J., Manhart, J. R.,
Murakami, N., Crane, E. H., Haufler, C. H. & Hauk,
W. D. 1995. Fern phylogeny based on rbcL nucleotide
sequences. Amer. Fern J. 85: 134181.
Hensen, R. V. 1990. Revision of the Polypodium loriceumcomplex (Filicales, Polypodiaceae). Nova Hedwigia 50:
279336.
Hetterscheid, W. L. A. & Hennipman, E. 1984. Venation patterns, soral characteristics, and shape of the fronds of the
microsorioid Polypodiaceae. Bot. Jahrb. Syst. 105: 11-47.
Hovenkamp, P. H. 1986. A monograph of the fern genus
Pyrrosia (Polypodiaceae). Leiden Bot. Ser. 2: 1331.
Hovenkamp, P. H. & Franken, N. A. P. 1993. An account of
the fern genus Belvisia Mirbel (Polypodiaceae). Blumea
37: 511527.
Huelsenbeck, J. P. & Bollback, J. P. 2001. Application of the
likelihood function in phylogenetic analysis. Pp. 415443
in: Balding, D. J., Bishop, M. & Cannings, C. (eds.),
Handbook of Statistical Genetics. John Wiley & Sons,
Ltd., Chichester.
Huelsenbeck, J. P. & Ronquist, F. 2001. MrBayes: Bayesian
inference of phylogenetic trees. Bioinformatics 17:
754755.
Kubitzki, K. (ed.). 1990. The Families and Genera of Vascular
Plants, vol. 1. Springer-Verlag, Berlin.
Larget, B. & Simon, L. 1999. Markov chain Monte Carlo
algorithms in the Bayesian analysis of phylogenetic trees.
Molec. Biol. Evol. 16: 750759.
Lellinger, D. B. 1972. A revision of the fern genus Niphidium.
Amer. Fern J. 62: 101120.
Lellinger, D. B. 1989. The ferns and fern-allies of Costa Rica,
Panama, and the Choc. (Part 1: Psilotaceae through
Dicksoniaceae). Pteridologia 2A: 1364.
Len, B. & Smith, A. R. 2003. New species and new combinations of Grammitidaceae from Peru. Amer. Fern J. 93:
8189.
Lewis, P. O. 2001. Phylogenetic systematics turns over a new
leaf. Trends Ecol. Evol. 16: 3037.
Maddison, D. R. & Maddison, W. P. 2000. MacClade 4.0.
Sinauer Associates, Sunderland, Massachusetts.
Martin, A. P. & Naylor, G. J. P. 1997. Independent origins of
filter-feeding in megamouth and basking sharks (Order
Lamniformes) inferred from phylogenetic analysis of
cytochrome b gene sequences. Pp. 3950 in: Yano, K.,
Morrissey, J. F., Yabumoto, Y. & Nakaya, K. (eds.),
Biology of Megamouth Shark. Tokai Univ. Press, Tokyo.
Murillo, M. T. & Smith, A. R. 2003. Luisma, a new genus of
Grammitidaceae (Pteridophyta) from Colombia. Novon
13: 313317.
Newman, E. 1840. A History of British Ferns and Allied
Plants. J. Van Voorst, London.
427
53 (2) May 2004: 415428
Parris, B. S. 1983. A taxonomic revision of the genus

Grammitis Swartz (Grammitidaceae: Filicales) in New
Guinea. Blumea 29: 13222.
Parris, B. S. 1990. Grammitidaceae. Pp. 153157 in: Kubitzki,
K. (ed.), The Families and Genera of Vascular Plants, vol.
1. Springer-Verlag, Berlin.
Parris, B. S. 1997a. Receptacular paraphyses in Asian,
Australasian and Pacific Islands taxa of Grammitidaceae
(Filicales). Pp. 8190 in: Johns, R. J. (ed.), Holttum
Memorial Volume. Royal Botanic Gardens, Kew.
Parris, B. S. 1997b. Themelium, a new genus of
Grammitidaceae (Filicales). Kew Bull. 52: 737741.
Parris, B. S. 1998a. Chrysogrammitis, a new genus of
Grammitidaceae (Filicales). Kew Bull. 53: 909918.
Parris, B. S. 1998b. Grammitidaceae. Pp. 450468 in:
McCarthy, P. M. (ed.), Flora of Australia, vol. 48.
ABRS/CSIRO Australia, Melbourne.
Parris, B. S. 2002. New species and new combinations in
African Grammitidaceae (Filicales). Kew Bull. 57:
423434.
Parris, B. S. 2003. Distribution of Grammitidaceae inside and
outside Malesia. Telopea 10: 451466.
Posada, D. & Crandall, K. A. 1998. Modeltest: testing the
model of DNA substitution. Bioinformatics 14: 817818.
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. Phylogenet. Evol.
26: 337347.
Rannala, B. & Yang, Z. 1996. Probability distribution of
molecular evolutionary trees: a new method of phylogenetic inference. J. Molec. Evol. 43: 304311.
Rdl-Linder, G. 1990. A monograph of the fern genus
Goniophlebium (Polypodiaceae). Blumea 34: 277423.
Rdl-Linder, G. 1994a. A monograph of the fern genus
Thylacopteris (Polypodiaceae). Blumea 39: 351364.
Rdl-Linder, G. 1994b. Revision of the fern genus
Polypodiopteris (Polypodiaceae). Blumea 39: 365371.
Roos, M. C. 1985. Phylogenetic Systematics of the
Drynarioideae (Polypodiaceae). Univ. Press, Utrecht.
Schneider, H., Smith, A. R., Cranfill, R., Hildebrand, T.,
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. Phylogenet. Evol. 31:
10411063.
Sen, U. & Hennipman, E. 1981. Structure and ontogeny of
stomata in Polypodiaceae. Blumea 27:175201.
Smith, A. R. 1992. A review of the fern genus
Micropolypodium (Grammitidaceae). Novon 2: 419425.
Smith, A. R. 1993. Terpsichore, a new genus of
Grammitidaceae (Pteridophyta). Novon 3: 478489.
Smith, A. R. 1995. Grammitidaceae. Pp. 366393 in: Davidse,
G., Sousa, M. S. & Knapp, S. (eds.), Flora
Mesoamericana, vol. 1. Universidad Nacional Autnoma
de Mxico, Mxico, D.F.
Smith, A. R. & Moran, R. C. 1992. Melpomene, a new genus
of Grammitidaceae (Pteridophyta). Novon 2: 426432.
Smith, A. R., Moran, R. C. & Bishop, L. E. 1991.
Lellingeria, a new genus of Grammitidaceae. Amer. Fern
J. 81: 7688.
Swofford, D. L. 1998. PAUP*. Phylogenetic Analysis Using
Parsimony (*and Other Methods). Version 4. Sinauer

Associates, Sunderland, Massachusetts.
Tryon, R. M. & Tryon, A. F. 1982. Ferns and Allied Plants,
with Special Reference to Tropical America. SpringerVerlag, New York.
Wagner, F. S. 1980. New basic chromosome numbers for genera of neotropical ferns. Amer. J. Bot. 67: 733738.
Walker, T. G. 1985. Cytotaxonomic studies of the ferns of
Trinidad 2. The cytology and taxonomic implications.
Bull. Brit. Mus. (Nat. Hist.), Bot. 13:149249.
Zink, M. J. 1993. Systematics of the Fern Genus Lepisorus (J.
Smith) Ching (Polypodiaceae-Lepisoreae). ADAG
Administration & Druck AG, Zurich.
428
53 (2) May 2004: Appendices
Appendix 1. Species list, collection and voucher information, and GenBank accession numbers.
Species
Adenophorus haalilioanus (Brack.) K.A. Wilson
Adenophorus hymenophylloides (Kaulf.)
Hook. & Grev.
Adenophorus periens L.E. Bishop
Adenophorus pinnatifidus Gaudich
Adenophorus tamariscinus (Kaulf.) Hook. & Grev.
Adenophorus tripinnatifidus Gaudich.
Calymmodon gracilis (Fe) Copel.
Calymmodon luerssenianus (Domin) Copel.
Campyloneurum augustifolium (Sw.) Fe
Ceradenia aulaeifolia L. E. Bishop
Ceradenia jungermannioides (Klotzsch) L.E. Bishop
Ceradenia kalbreyeri (Baker) L.E. Bishop
Ceradenia pilipes (Hook.) L.E. Bishop
Ceradenia spixiana (Mart. ex Mett.) L.E. Bishop
Chrysogrammitis musgraviana (Baker) Parris
Cochlidium punctatum (Raddi) L.E. Bishop
Cochlidium rostratum (Hook.) Maxon ex C. Chr.
Cochlidium seminudum (Willd.) Maxon
Ctenopteris heterophylla (Labill.) Tindale
Ctenopteris heterophylla (Labill.) Tindale
Ctenopteris lasiostipes (Mett.) Brownlie
Ctenopteris nutans (Blume) J. Sm.
Ctenopteris repandula (Mett.) C. Chr. & Tardieu
Ctenopteris aff. repandula (Mett.) C. Chr. & Tardieu
Ctenopteris rhodocarpa Copel.
Enterosora percrassa (Baker) L.E. Bishop
Enterosora trifurcata (L.) L.E. Bishop
Grammitis billardierei Willd.
Grammitis bryophila (Maxon) F. Seym.
Grammitis ciliata Colenso
Grammitis deplanchei (Baker) Copel.
Grammitis forbesiana W. H. Wagner
Grammitis hirtelloides (Copel.) Copel.
Grammitis hookeri (Brack.) Copel.
Grammitis knutsfordiana (Baker) Copel.
Grammitis melanoloma (Cordem.) Tardieu
Grammitis parva (Brause) Copel.
Grammitis poeppigiana (Mett.) Pic. Serm.
Grammitis poeppigiana (Mett.) Pic. Serm.
Grammitis pseudociliata Parris
Grammitis tenella Kaulf.
Lellingeria apiculata (Kunze ex Klotzsch)
A.R. Sm. & R.C. Moran
Lellingeria hirsuta (Baker ex Hemsl.)
Lellingeria limula (H. Christ)
Lellingeria pseudomitchellae (Lellinger)
Lellingeria schenckei (Hieron.)
Lellingeria subsessilis (Baker)
Melpomene flabelliformis (Poir.)
Melpomene moniliformis (Lag. ex Sw.)
Locality
Hawaii, U.S.A.
Hawaii, U.S.A.
Collector, number, and herbarium

Ranker 1561 (COLO)
Ranker 1203 (COLO)
GenBank acc. no.

rbcL
atpB
AF468200 AF469775
AF468204 AF469780
Hawaii, U.S.A.
Hawaii, U.S.A.
Hawaii, U.S.A.
Hawaii, U.S.A.
Taiwan
Papua New Guinea
Costa Rica
Costa Rica
Costa Rica
Costa Rica
Costa Rica
Brazil
Sabah
Brazil
Panama
Dominican Republic
New Zealand
New Zealand
New Caledonia
Papua New Guinea
Papua New Guinea
Papua New Guinea
Papua New Guinea
Costa Rica
Puerto Rico
New Zealand
Costa Rica
New Zealand
New Caledonia
Hawaii, U.S.A.
Fiji
Hawaii, U.S.A.
Fiji
La Runion
Papua New Guinea
Chile
Australia
New Zealand
Hawaii, U.S.A.
Brazil
Ranker 1114 (COLO)

Ranker 1559 (COLO)
Ranker 1031 (COLO)
Ranker 1102 (COLO)
Chiou 97-09-12-01 (COLO, TAIF, UC)
Ranker 1771 & Trapp (COLO, UC)
Chisaki & Carter 1004 (UC)
A. Rojas & al. 3232 (CR, INB, MO, UC)
A. R. Smith 2576 (UC)
A. Rojas & al. 3323 (voucher lost)
A. Rojas & al. 3233 (INB)
Salino 3008 (UC)
Kessler 12570 (UC)
Salino 3127 (BHCB, UC)
I. Valdespino & J. Aranda 180 (UC)
S. R. Hill 29102A no voucher1
Parris 12419 (AK)
Hodel 1448 (UC)
Moraga & Rojas 508 (UC)
Ranker 1608 (COLO)
Parris 12421 (AK)
A. Rojas & al. 3240 (UC)
Hodel 1450 (UC)
Ranker 1321 (COLO)
J. Game 95-62 (UC)
Ranker 1116 (COLO)
J. Game 95-81 (UC)
Ranker 1504 & Adsersen (COLO)
Ranker 1763a (COLO, UC)
C. Taylor 6072 (UC)
Weber 13772 (COLO)
Parris 12420 (AK)
Ranker 1352 (COLO)
Salino 3009 (UC)
AF468199
AF468201
AF468206
AF468207
AY362341
AY460618
AF470344
AY460619
AY460620
AY460621
AY460622
AY460623
AY460624
AY460625
AY460626
AY460627
AY460628
AY460629
AY460630
AY460631
AY460633
AY460632
AY460634
AY460635
AY460636
AY460637
AF468208
AY460638
AY460639
AY460640
AY460641
AY460642
AY362342
AY460643
AY460644
AY460646
AY460647
AY460645
AF468198
AY362343
Costa Rica
AY460648 AY459481
Costa Rica
A. Rojas & E. Fletes 3145 (UC)
AY460649 AY459482
Costa Rica
AY460650 AY459523
Costa Rica
A. Rojas 3005 (MO)
AY460652 AY459484
Brazil
AY460651 AY459483
Costa Rica
A. Rojas 3022 & M. Mata (UC)
AY460653 AY459485
Colombia
P. Sanchez-B. 183 (UC)
AY460656 AY459488
Costa Rica
M. Moraga & A. Rojas 446 (INB)
AY460654 AY459486
AF469774
AF469777
AF469782
AF469783
AY459451
AY459452
AY459515
AY459453
AY459454
AY459455
AY459456
AY459457
AY459458
AY459520
AY459459
AY459460
AY459461
AY459462
AY459463
AY459464
AY459466
AY459465
AY459467
AY459468
AY459521
AY459469
AF469784
AY459470
AY459471
AY459472
AY459522
AY459473
AY459474
AY459475
AY459476
AY459479
AY459478
AY459477
AF469773
AY459480
1 see Hill 27921, ILLS, ATRC and VT; C. seminudum collected from vicinity of Hill 29102A
53 (2) May 2004: xxxx
Appendix 1 (continued.)
Locality
Costa Rica
Collector, number, and herbarium

GenBank acc. no.

rbcL
atpB
AY460648 AY459481
Costa Rica
A. Rojas & E. Fletes 3145 (UC)
AY460649 AY459482
Costa Rica
AY460650 AY459523
Costa Rica
A. Rojas 3005 (MO)
AY460652 AY459484
Brazil
AY460651 AY459483
Costa Rica
AY460653 AY459485
Colombia
P. Sanchez-B. 183 (UC)
AY460656 AY459488
Costa Rica
AY460654 AY459486
Ecuador
K. A. Wilson 2806 (UC)
AY460657 AY459489
A. Smith 1357 (UC)

NYBG 954/95A (UC)
AY362574 AY459516
AY362579 AY459517
Micropolypodium hyalinum (Maxon) A.R. Sm.

Micropolypodium taenifolium (Jenman) A.R. Sm.
Micropolypodium zurquinum (Copel.) A.R. Sm.
Pecluma alfredii (Rosenst.) M.G. Price
Polypodium glycyrrhiza D.C. Eaton
Prosaptia alata (Blume) H. Christ
Venezuela
provenance unknown
(cultivated source)
Costa Rica
Costa Rica
Costa Rica
Mexico
California, USA
Kosrae
AY362344
AY460658
AY460659
AY096206
U21146
AY460660
AY459490
AY459491
AY459492
AY459519
AY459518
AY459493
Prosaptia contigua (G. Forst.) C. Presl

Prosaptia obliquata (Blume) Mett.
Prosaptia palauensis Hosok.
Prosaptia pubipes Copel.
Scleroglossum sulcatum (Kuhn) Alderw.
Scleroglossum sulcatum (Kuhn) Alderw.
Taiwan
Taiwan
Palau
Fiji
Pohnpei
Pohnpei
AY362345
AY460661
AY460662
AY460663
AY460664
AY460665
AY459494
AY459495
no data
AY459496
AY459497
AY459498
Terpsichore achilleifolia (Kaulf.) A.R. Sm.

Terpsichore alsopteris (C. V. Morton) A.R. Sm.
Terpsichore anfractuosa (Kunze ex Klotzsch)
B. Len & A.R. Sm.
Terpsichore cultrata (Bory ex Willd.) A. R. Sm.
Terpsichore eggersii (Baker ex Hook.) A.R. Sm.
Terpsichore hanekeana (Proctor) A.R. Sm.
Terpsichore lanigera (Desv.) A.R. Sm.
Terpsichore lanigera (Desv.) A.R. Sm.
Terpsichore lehmanniana (Hieron.) A.R. Sm.
Terpsichore longisetosa (Hook.) A.R. Sm.
Terpsichore pichinchae (Sodiro) A.R. Sm.
Terpsichore semihirsuta (Klotzsch) A. R. Sm.
Terpsichore senilis (Fe) A.R. Sm.
Terpsichore sp.
Terpsichore subscabra (Fe) A.R. Sm.
Terpsichore subtilis (Kunze ex Klotzsch) A.R. Sm.
Themelium tenuisectum (Blume) Parris
Xiphopteris conjunctisora (Baker) Copel.
Brazil
Ecuador
Costa Rica

A. Rojas 3007 (UC)
Kimnach 95 (UC)
Haufler & Mesler s.n. (KANU)
T. Flynn 6004
(UC, AD, BISH, PTBG, US)
Chiou 97-09-12-05 (COLO, UC, TAIF)
Chiou 97-09-12-04 (COLO, UC, TAIF)
R. J. Rondeau 93-011& Rodda (UC)
J. Game 95-65 (UC)
T. Flynn 6287 (PTBG, US, UC)
Bowden-Kerby 24182b
(UC, GUAM)
J. Cordeiro & O. Ribas 1398 (UC)
K. A. Wilson 2609a (UC)
Colombia
Dominican Republic
Puerto Rico
Costa Rica
Peru
Ecuador
Costa Rica
Colombia
Peru
Costa Rica
Peru
Costa Rica
Costa Rica
Taiwan
Papua New Guinea
C. Dassler 94-7-19-1 (ILLS)

S. Hill 29109 (UC)
Ranker 1610 (COLO)
B. Len 3647 (USM, UC)
K. A. Wilson 2589 (UC)
C. Dassler 94-7-13-1 (ILLS)
Chiou 97-09-12-02 (COLO, TAIF, UC)
Species
Lellingeria hirsuta (Baker ex Hemsl.)
Lellingeria limula (H. Christ)
Lellingeria pseudomitchellae (Lellinger)
Lellingeria schenckei (Hieron.)
Lellingeria subsessilis (Baker)
Melpomene flabelliformis (Poir.)
Melpomene moniliformis (Lag. ex Sw.)
Melpomene pseudonutans (H. Christ & Rosenst.)
Microgramma percussa (Cav.) de la Sota
Microgramma squamulosa (Kaulf.) de la Sota
AY460666 AY459499
AY460667 AY459500
AY460668 AY459501
AY460669
AF468209
AY460670
AY460671
AY460672
AY460673
AY460674
AY460675
AY460676
AY096208
AY460679
AY460677
AY460678
AY362346
AY460680
AY459502
AF469785
AY459503
AY459504
AY459505
AY459506
AY459507
AY459508
AY459509
AY459510
AY459513
AY459511
AY459512
no data
AY459514
Appendix 2. Morphological character states for species studied, listed in order from 187 as in Table 1. Character-state
codes in parentheses represent polymorphisms for single characters. Annotations after duplicate accessions of single
species refer to particular specimens as in Appendix 1. Character numbers are as in Table 1.
1
30
59
Adenophorus
(234)
pinnatifidus
0
(01)
Adenophorus
3
hymenophylloides 0
?
Calymmodon
(234)
gracilis
0
0
Calymmodon
3
luerssenianus
0
0
Ceradenia
3
aulaeifolia
1
1
Ceradenia
3
jungermannioides 0
1
Ceradenia kalbreyeri 3
0
1
Ceradenia pilipes
3
0
1
Ceradenia spixiana (23)
0
1
Chrysogrammitis
3
musgraviana
0
0
Cochlidium
3
punctatum
0
?
Cochlidium
3
rostratum
0
?
Cochlidium
3
seminudum
0
?
Ctenopteris
(0234)
heterophylla
0
(Smith)
0
Ctenopteris
(0234)
heterophylla
0
(Parris)
0
Ctenopteris
3
lasiostipes
0
1
Ctenopteris nutans
3
0
1
Ctenopteris
(234)
repandula
0
0
Ctenopteris
(23)
aff. repandula
0
(01)
Ctenopteris
3
rhodocarpa
0
(01)
Enterosora percassa 3
0
1
Enterosora trifurcata 3
0
1
Grammitis
(023)
billardierei
0
0
Grammitis
(13)
bryophila
0
?
2
31
60
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
1
0
1
1
0
1
1
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
0
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
3
32
61
0
2
?
0
5
?
0
2
?
0
2
?
0
3
?
0
0
?
0
2
?
0
2
?
0
2
?
0
2
?
0
0
?
0
0
?
0
0
?
0
5
?
0
5
?
0
3
?
0
3
?
0
2
?
0
2
?
0
2
?
0
0
?
0
1
?
0
0
?
0
0
?
4
33
62
1
?
0
0
0
0
0
0
0
0
0
0
0
0
0
0
?
0
0
1
0
0
0
0
0
1
0
0
0
0
0
?
0
0
?
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
?
0
0
0
0
0
?
0
0
?
0
5
34
63
0
?
1
0
0
1
?
0
1
?
0
1
1
0
1
?
?
1
1
0
1
?
0
1
1
0
1
0
0
1
?
?
1
?
?
1
?
?
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
?
0
1
0
0
1
1
?
1
1
0
1
0
?
1
?
?
1
6
35
64
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
2
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
7
8
9 10 11
36 37 38 39 40
65 66 67 68 69
1
0
0
1
0
2
0
0
?
0
1
0
0
0
?
1
0
0
1
0
2
0
0
?
0
1
0
0
0
?
0
0
0
1
0
1
0
1
0
0
1
1
0
0
?
0
0
0
1
0
2
0
1
0
0
1 (01) 0
0
?
1
0
1
1
0
2
0
0
?
0
0
0
0
0
?
0
0
0
1
0
1
0
0
?
0
0
0
0
0
?
1
0
1
1
0
2
0
0
?
0
0
0
0
0
?
0
0
1
1
0
1
0
0
?
0
1
0
0
0
?
1
0
1
1
0
1
0
0
?
0
0
0
0
0
?
1
0
0
1
0
1
0
0
?
0
1
0
0
0
?
0
0
0
1
0
0
0
1
0
0
1
1
0
0
?
0
0
0
1
0
0
0
0
?
0
1
2
2
0
?
0
0
0
1
0
0
0 (01) 0
0
1
2
0
0
?
1
0
0
1
0
2
0
1
0
0
1
0
0
0
?
1
0
0
1
0
2
0
1
0
0
1
0
0
0
?
1
0
0
1
0
2
0
1
0
0
1
0
0
0
?
1
0
0
1
0
2
0
0
?
1
1
0
0
0
?
0
0
0
1
0
1
0
1
0
?
1
0
0
0
?
0
0
?
1
?
1
0
1
0
0
0
0
0 (01) ?
1
0
1
1
0
2
0
0
?
1
1
0
0
0
?
1
0 (01) 1
0
0
0
0
?
0
0
0
2
0
?
1
0
1
1
0
0
0
0
?
0
0 (01) 1
0
?
1
0
0
1
0
2
0
0
?
0
1
1
1
1
?
0
0
0
1
0
2
1
0
0
0
1 (01) 0
0
?
Character number
12 13 14 15
41 42 43 44
70 71 72 73
0
0
0
1
0
0
1
0
1
?
?
1
0
0
0
1
1
0
1
0
1
?
?
1
0
0
0
1
0
0
1
0
1
?
?
0
0
0
0
1
0
0
1
0
1
?
?
0
0
0
0
1
0
0
1
0
1
?
?
1
0
0
0
0
?
0
1
0
1
?
?
1
0
0
0
1
0
0
1
0
1
?
?
1
0
0
0
0
0
0 (01) 0
1
?
?
1
0
0
0
1
0
0
1
0
1
?
?
1
0
0
0
1
0
0
1
0
0
?
?
1
0
0
0
1
?
0
0
0
0
?
?
0
0
0
0
1
?
0
0
0
0
?
?
0
0
0
0
1
0
0
0
0
0
?
?
0
0
0
0
1
0
0
1
0
1
?
?
0
0
0
0
1
0
0
1
0
1
?
?
0
0
0
0
1
0
0
1
0
1
?
?
1
0
0
0
1
0
0
1
0
1
?
?
1
0
0
0
1
0
0
1
0
1
?
?
0
?
?
?
1
0
0
0
0
1
?
?
0
0
0
0
1
0
0
1
0
1
?
?
0
0
0
0
1
0
0
1
0
?
?
?
1
0
0
0
1
0
0
1
0
1
?
? (01)
0
0
0
1
?
0
1
0
0
?
?
0
0
0
0
1
?
0
1
0
1
?
?
0
16
45
74
2
0
1
2
0
1
1
0
?
1
0
?
2
0
1
?
0
1
2
1
1
2
0
1
2
1
1
2
0
1
1
0
?
(12)
0
?
1
0
?
2
0
?
2
0
?
3
0
0
2
1
0
1
0
?
1
0
?
3
1
?
(01)
1
0
(01)
1
0
1
0
?
2
0
?
17
46
75
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
?
0
0
0
(01)
0
0
0
0
0
(01)
0
0
0
0
0
0
0
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
18 19 20
47 48 49
76 77 78
1
2
0
0
0
1
?
?
0
2
2
0
0
0
1
?
?
0
1
0
0
0
0
1
?
?
1
1
0
0
0
0
0
?
?
1
2
1
0
0
0
1
?
?
0
?
?
?
0
0
1
?
?
0
2
2
0
0
0
1
?
?
0
2
1
0
0
0
1
?
?
0
2
1
0
0
0
1
?
?
0
1
0
0
0
0
2
?
?
0
0
0
?
0
0
0
?
?
0
0
0 (12)
0
0
0
?
?
0
0
0
?
0
0
0
?
?
0
1
0
0
0
0
1
?
?
0
1
0
0
0
0
1
?
?
0
1
0
0
0
0
1
?
?
0
1
2
0
0
0
1
?
?
0
1
0
0
0
0
1
?
?
0
1 (01) 0
0
0
1
?
?
0
1
2
0
0
0
1
?
?
0
0
0 (12)
0
0
1
?
?
0
0
0
0
0
0
1
?
?
0
1
0
0
0
0
1
?
?
0
0
2
0
0
0
0
?
?
0
21 22 23
50 51 52
79 80 81
0
2
2
1
0
?
3
0
1
0
2
1
1
0
?
3
0
1
0
1
1
?
0
?
3
0
1
0
1
1
?
0
?
3
0
1
0
2
3
1
0
?
3
0
1
?
?
?
1
0
?
3
0
1
0
2 (23)
1
0
?
3
0
1
0
2
2
1
0
?
3
0
1
0
2 (23)
1
0
?
3
0
1
0 (01) 1
1
0
?
3
0
1
(01) (12) 2
1
0
?
3
1
1
(01) (12) 2
1
0
?
3
0
1
? (12) 2
1
0
?
3
?
1
0
?
2
1
0
?
3
?
1
0
?
2
1
0
?
3
?
1
0
2
2
1
0
?
3
0
1
0
1
2
1
0
?
3
0
1
0
1
2
1
0
?
3
0
1
0 (012) 2
1
0
?
3
0
1
0
1
2
1
0
?
3
0
1
0 (01) 2
1
1
0
3
0
1
0
0
2
1 (01) 0
3
?
1
0
2
2
0
0
?
3
0
1
0 (23) 3
1
0
?
3
?
1
24
53
82
2
2
0
(12)
2
0
2
2
0
?
2
0
2
0
?
?
0
0
(12)
0
1
2
0
?
2
0
1
0
2
0
2
0
1
2
0
0
2
0
?
2
2
1
2
2
1
2
2
0
2
2
0
2
2
1
(12)
0
0
2
2
0
(12)
0
1
(12)
0
?
2
1
0
2
0
0
25
54
83
?
0
0
0
0
0
0
0
0
0
0
0
1
?
0
?
0
0
1
0
0
1
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
(12)
0
0
0
0
0
1
0
0
26
55
84
0
1
1
0
1
1
0
1
1
0
1
1
0
1
1
?
1
1
0
1
1
0
1
1
0
1
1
0
1
1
0
0
1
0
0
1
0
1
1
0
1
1
0
1
1
3
1
1
3
1
1
0
1
1
0
1
0
2
1
1
0
1
1
0
1
1
0
1
1
0
0
1
27 28
56 57
85 86
0
3
(12) (23)
1
1
0
3
1
?
1
1
0
2
(02) 1
1
1
0 (02)
2
1
1
1
3
3
2
3
1
?
?
?
1
?
1
1
(02) 0
1 (23)
1
1
3
3
2
0
1
?
(23) (23)
(12) (23)
1
1
3
3
(12) 2
1
1
1
0
?
?
1
1
1
0
?
?
1
1
0
0
1
?
1
?
0
0
(12) (13)
1
1
0
0
(12) (13)
1
1
0
0
(012) 1
1
1
2
2
?
1
1
1
0 (02)
(012) 1
1
1
0
2
(02) 1
?
?
2
2
(012) 1
1
1
3
3
1 (23)
1
1
3
3
1 (23)
1
1
0
0
(012) 1
1
1
0
0
?
?
1
1
29
58
87
0
(13)
0
0
2
0
0
?
?
0
3
?
0
1
?
?
1
0
0
1
0
0
2
?
0
1
0
0
2
?
0
0
?
0
0
4
0
0
?
0
(23)
?
0
(23)
?
0
1
?
0
1
?
0
1
?
0
(13)
?
0
0
?
2
1
0
0
1
0
0
0
?
0
0
?
1
30
59
Grammitis ciliata (023)
0
0
Grammitis
3
deplanchei
0
?
Grammitis
3
forbesiana
0
1
Grammitis
(23)
hirtelloides
0
1
Grammitis hookeri (23)
0
1
Grammitis
(123)
knutsfordiana
0
1
Grammitis
3
melanoloma
0
0
Grammitis parva
3
0
1
Grammitis
3
poeppigiana (Chile) 0
?
Grammitis
2
poeppigiana
0
(Australia)
0
Grammitis
3
pseudociliata
0
(01)
Grammitis tenella (123)
0
0
Lellingeria
(23)
apiculata (Brazil)
0
1
Lellingeria
(23)
apiculata
0
(Costa Rica)
1
Lellingeria hirsuta
3
0
1
Lellingeria limula
3
0
(01)
Lellingeria
3
pseudomitchellae
0
0
Lellingeria schenckei 2
0
0
Lellingeria
3
subsessilis
0
0
Melpomene
(023)
flabelliformis
0
1
Melpomene
0
moniliformis
0
(Costa Rica)
1
Melpomene
0
moniliformis
0
(Mexico)
1
Melpomene
3
pseudonutans
0
1
Micropolypodium
3
hyalinum
0
1
Micropolypodium
3
taenifolium
0
1
2
31
60
0
0
0
1
0
0
0
0
0
1
0
0
1
0
0
0
0
0
1
0
0
1
0
0
1
0
0
1
0
0
0
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
3
4
5
32 33 34
61 62 63
0
0
0
0
?
?
?
0
1
0
0
0
0
?
?
?
0
1
0
0
0
0
?
?
?
0
1
0
0
?
0
?
?
?
0
1
0
0
0
0
?
?
?
0
1
0
0
0
0
?
?
?
0
1
0
0
?
0
?
?
?
0
1
0
0
?
0
?
?
?
0
1
0
0
?
0
?
?
?
0
1
0
0
0
0
?
?
?
0
1
0
0
0
0
?
?
?
0
1
0
0
0
0
?
?
? (01) 1
0
0
1
2
1
0
?
0
1
0
0
1
2
1
0
?
0
1
0
0
1
2
1
0
?
0
1
0
0
?
2
0
0
?
0
1
0
0
?
1
0
0
?
0
1
0
0
?
1
0
0
?
0
1
0
0
?
2
0
0
?
0
1
0
0
1
2
0
0
?
0
1
0
0
1
2
0
0
?
0
1
0
0
1
2
0
0
?
0
1
0
0
1
2
0
0
?
0
1
0
0
?
2
0
0
?
0
1
0
0
?
2
0
0
?
0
1
6
7
8
9 10
35 36 37 38 39
64 65 66 67 68
0
1
0
0
1
0
1
0
0
?
0
0
0
0
0
0
1
0
1
1
0
0
0
0
?
0
1
0 (01) (01)
0
1
0
1
1
0
1
0
1
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
?
0
1
0
0
0
0
1
0
0
1
?
1
0
1
0
0
1
0
0
0
0
1
0 (01) 1
0
1
0
1
0
0
0
0
0
0
0
0
0
0
1
0
1
1
0
?
0
1
0
0
0
0
0
0
0
1
0
1
0
0
?
0
1
0
0
0
0
0
0
0
1
0
0
0
1
0
0 (01) 0
0
0
0
1
0
0
1
0
0
0 (01) 0
0
1
0
0
0
0
1
0
0
1
0
1
0
1
0
0
0 (01) 0
0
0
1
0
0
1
0
1
0
0
?
0
1
0
0
0
0
1
0
1
1
0
2
0
1
0
0
1
0
0
0
0
1
0
1
1
0
2
0
1
0
0
1
0
0
0
0
1
0
1
1
0
2
0
1
0
0
0
0
1
0
0
0
0
0
1
0
2
0
1
0
0
1
1 (01) 0
0
0
0
0
1
0
2
0
1
0
0
1
0
0
0
0
0
0
0
1
0
1
0
1
0
0
1
0
0
0
0
0
0
0
1
0
2
0
1
0
0
0
0
1
0
0
1
0 (01) 0
0
2
0
1
0
0
1
0
0
0
0
1
0 (01) (01)
0
2
0
1
0
0
1
0
0
0
0
1
0 (01) (01)
0
2
0
1
0
0
1
0
0
0
0
1
0 (01) 1
0
2
0
1
0
0
1
0
0
0
0
0
0
0
1
0
1
0
1
0
0
1
0
0
0
0
0
0
0
1
0
2
0
1
0
0
1
0
0
0
11
40
69
?
0
?
0
0
?
?
0
?
0
0
?
0
0
?
?
0
?
0
0
?
0
0
?
0
0
?
0
0
?
?
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
Character number
12 13 14 15
41 42 43 44
70 71 72 73
?
?
?
1
0
0
1
0
0
?
?
1
0
0
0
1
?
0
0
0
2
?
?
0
?
?
?
1
0
0 (01) 0
0
?
?
0
0
0
0
1
?
0
1
0
0
?
?
0
0
0
0
1
?
0
1
0
0
?
?
0
?
?
?
1
0
0 (01) 0
0
?
?
1
0
0
0
1
?
0 (01) 0
0
?
?
0
0
0
0
0
?
0
1
0
0
?
?
1
0
0
0
1
?
0
0
0
1
?
?
0
0
0
0
1
?
0
0
0
0
?
?
0
?
?
?
1
0
0
1
0
0
?
?
1
0
0
0
1
?
0
1
0
1
?
?
1
0
0
0
1
0
0
1
0
1
?
?
0
0
0
0
1
0
0
1
0
1
?
?
0
0
0
0
1
0
0
1
0
1
?
?
0
0
0
0
1
0
0
1
0
0
?
?
0
0
0
0
0
0
0
1
0
0
?
?
1
0
0
0
1
0
0
1
0
0
?
?
0
0
0
0
1
0
0
1
0
1
?
?
0
0
0
0
1
0
0
1
0
1
?
?
1
0
0
0
1
0
0
1
0
1
?
?
0
0
0
0
1
0
0
1
0
1
?
?
0
0
0
0
1
0
0
1
0
1
?
?
1
0
0
0
1
0
0
1
0
0
?
?
1
0
0
0
1
0
0
1
0
0
?
?
1
16 17
45 46
74 75
1
0
0
0
1
0
2
0
0
0
?
2
1
0
0
0
?
1
3
1
0
0
?
2
1
0
1
0
?
2
1 (01)
0
0
1
1
2
0
0
0
?
0
?
?
0
0
0
2
(01) 0
0
0
?
0
1
0
?
0
?
0
1
0
0
0
1
0
2
0
0 (123)
1
0
3
0
0
0
?
0
3
0
0
0
?
0
3
0
0
0
?
0
3
1
0
0
?
0
?
?
0
0
0
0
3
0
0
0
?
0
2
0
0
0
?
0
2
1
0
1
0
0
2
1
0
1
?
0
2
1
0
1
?
0
2
1
0
2
0
0
(01) 0
0
0
0
0
2
0
0
0
0
0
18
47
76
1
0
?
1
0
2
1
0
1
1
0
?
1
0
1
1
0
1
2
0
?
?
0
1
1
0
?
1
0
?
1
0
?
?
0
?
1
0
?
1
0
?
2
0
?
1
0
?
?
0
?
1
0
?
2
0
?
2
0
?
2
0
?
2
0
?
2
0
?
2
0
?
2
0
?
19 20 21
48 49 50
77 78 79
0
0
0
0
0
?
?
0
3
2
0
0
0
1
0
1
0
3
0
0
0
0
0
?
2
0
3
0
0
0
0
1
1
?
0
3
0
0
0
0
1
1
2
0
3
0 (01) 0
0
0
?
2
0
3
2
0
0
0
1
0
?
0
3
?
?
?
0
1
0
2
0
3
0
0
0
0
0
1
?
0
3
0
0
0
0
1
0
?
0
3
0 (01) 0
0
0
?
?
0
3
2
0
0
0
1
?
?
0
3
0
0
0
0
1
1
?
0
3
0
0
0
0
1
1
?
0
3
1
0
0
0
1
1
?
0
3
0
0
0
0
0
?
?
0
3
?
?
?
0
0
?
?
0
3
0
0
0
0
1
1
?
0
3
0
0
0
0
1
1
?
0
3
0
0 (01)
1
1
1
?
0
3
0
1 (01)
1
1
1
?
0
3
0
1 (01)
1
1
1
?
0
3
0
1 (01)
1
1
1
?
0
3
1
0
0
0
1
1
?
0
3
2
0
0
0
1
1
?
0
3
22
51
80
1
0
0
3
0
0
(12)
0
0
(01)
0
0
(12)
0
0
1
0
0
3
0
0
?
0
0
1
0
0
2
0
0
1
0
0
3
0
0
2
0
0
2
0
0
2
0
?
0
0
0
?
0
0
1
0
0
1
0
0
1
0
2
1
0
?
1
0
?
1
0
0
1
0
0
1
0
0
23 24 25
52 53 54
81 82 83
2
2
1
?
0
0
1
0
0
2 (12) ?
?
2
0
1
0
0
2 (012) 1
?
0
0
1
0
0
2 (12) ?
?
1
0
1
0
0
2 (12) ?
?
1
0
1
0
0
2 (12) 1
? (12) 0
1
0
0
2
2
?
?
2
0
1
0
0
?
?
?
?
2
0
1
0
0
2 (12) 1
?
0
0
1
1
0
2
2
0
?
1
0
1
1
0
2 (12) 1
?
0
0
1
0
0
(23) 2
?
?
2
0
1
0
0
2
2
1
?
0
0
1
1
0
2
2
1
?
0
0
1
1
0
2
2
1
?
0
0
1
?
0
1
0 (01)
?
1
0
1
0
0
?
?
?
?
0
0
1
1
0
3
2
1
?
0
0
1
1
0
2
2
1
?
0
0
1
1
0
2 (12) 1
?
0
0
1
0
0
2 (12) 1
?
0
0
1
?
0
2 (12) 1
?
0
0
1
?
0
2 (12) 1
?
0
0
1
1
0
2
2
1
?
0
0
1
1
0
2
2
1
?
0
0
1
2
0
26
55
84
0
1
0
0
1
1
0
1
0
3
1
1
0
1
1
0
1
0
0
1
1
?
1
1
0
0
1
0
1
1
0
1
0
0
1
1
3
1
1
3
1
1
3
1
1
3
1
1
?
1
1
3
1
1
3
1
1
3
1
1
3
1
1
3
1
1
3
1
1
0
1
1
0
1
1
27 28
56 57
85 86
0
0
(02) 1
?
?
2
2
(12) 1
1
1
0
0
0
?
?
?
0
0
(01) ?
1
1
0
0
(01) ?
1
1
0
0
0
?
?
?
0
3
(12) 3
1
1
?
?
(01) ?
1
1
0
0
?
?
1
1
0
0
(12) 3
1
1
0
0
(02) 1
?
?
0
3
(12) 3
1
1
2
2
(012) 1
1
1
2
2
(012) 1
1
1
2
2
0
?
1
1
0
0
(012) 1
1
1
?
?
(12) 1
1
1
2
2
(012) 1
1
1
2
2
(012) 1
1
1
0 (34)
1 (23)
1
1
0
3
1
?
1
?
0
3
1
?
1
?
0
3
1
?
1
1
(23) 3
1 (23)
1
1
2
2
1 (23)
1
1
29
58
87
0
1
?
0
0
?
0
1
?
0
1
?
0
1
?
0
1
?
0
0
?
?
1
?
0
0
?
0
0
0
0
1
?
0
(23)
2
1
1
?
1
1
?
0
1
?
0
(13)
5
?
1
?
0
0
?
1
0
?
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
1
0
1
30
59
Micropolypodium
3
zurquinum
0
1
Prosaptia alata
3
0
1
Prosaptia contigua (023)
0
(01)
Prosaptia
(0234)
obliquata
0
(01)
Prosaptia paluaensis 3
0
(01)
Prosaptia pubipes
3
0
1
Scleroglossum
(23)
sulcatum (Flynn)
0
0
Scleroglossum
(23)
sulcatum
0
(Bowden-Kirby)
0
Terpsichore
3
achilleifolia
0
1
Terpsichore
3
alsopteris
0
1
Terpsichore
3
anfractuosa
0
1
Terpsichore cultrata 3
0
0
Terpsichore eggersii 3
0
1
Terpsichore
3
hanekeana
0
1
Terpsichore
3
lanigera
1
(Costa Rica)
(01)
Terpsichore
3
lanigera (Peru)
1
(01)
Terpsichore
3
lehmanniana
1
(01)
Terpsichore
(023)
longisetosa
0
1
Terpsichore
3
pichinchae
0
1
Terpsichore
3
semihirsuta
0
1
Terpsichore senilis
3
1
0
Terpsichore
3
subscabra
1
0
Terpsichore subtilis 3
0
(01)
Terpsichore sp.
3
0
1
Themelium
(13)
tenuisectum
0
(01)
2
3
4
31 32 33
60 61 62
1
0
0
0
2
0
0
?
0
0
0
0
0
1
0
0
?
0
1
0
0
0
2
0
0
?
0
1
0
0
0
3
0
0
?
0
0
0
0
0
2
0
0
?
0
0
0
0
0
2
0
0
?
0
1
0
0
0
0
?
0
?
0
1
0
0
0
0
?
0
?
0
1
0
0
0
2
0
0
?
0
1
0
0
0
2
0
0
?
0
1
0
1
0
2
0
0
?
0
1
0
0
0 (23) 0
0
?
0
1
0
0
0
2 (01)
0
?
0
1
?
0
0
2
1
0
?
0
1
0
0
0 (23) 0
0
?
0
1
0
0
0 (23) 0
0
?
0
1
0
0
0
2
1
0
?
0
1
0
0
0
5
1
0
?
0
1
0
0
0
2
0
0
?
0
1
1
0
0
2
0
0
?
0
1
0
0
0
3
0
0
?
0
1
0
0
0
3
0
0
?
0
1
0
0
0
2
0
0
?
0
1
0
0
0
2
?
0
?
0
1
0
0
0
5
0
0
?
0
5
34
63
?
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
?
?
1
?
?
1
?
0
1
?
0
1
?
0
1
?
0
1
1
0
1
?
0
1
0
0
1
0
0
1
1
0
1
?
0
1
?
0
1
?
0
1
?
0
1
?
0
1
?
0
1
?
0
1
0
0
1
6
7
8
9 10
35 36 37 38 39
64 65 66 67 68
0
0
0
0
1
0
1
0
1
0
0
1
0
0
0
0
1
0 (01) 1
0 (01) 0
0
?
0
0
0
2
1
0
1
0
1
1
0
2
0
0
?
0
1
1
0
1
0
1
0
0
1
0
1
0
0
?
0
1
0
2
1
0
1
0
0
1
0
1
0
0
?
0
0
0
2
1
0
1
0
0
1
0
1
0
0
?
0
0
1
2
1
0
0
0
0
1
0
0
0
0
?
1
1
2
2
0
0
0
0
0
1
0
0
0
0
?
1
1
2
2
0
0 (01) 0
0
1
0
1
0
1
0
0
0
0
0
0
0
0
0
1
1
0
2
0
1
1
0
0
0
0
0
0
0
0
0
1
0
2
0
1
0
0
1
0
0
0
0
0
0
0
1
0
2
0
1
0
0
0
0
0
0
0
1
0
0
1
0
2
0
1
0
0
0
0
0
0
0
0
0
0
1
0
2
0
1
?
0
0
0
0
0
0
1
0 (01) 1
0
2
0
1
0
0
0
0
0
0
0
1
0 (01) 1
0
2
0
1
0
0
0
0
0
0
0
1
0
1
1
0
1
0
1
0
0
0
0
0
0
0
0
0
1
1
0
2
0
1
0
0
0
0
0
0
0
0
0
0
1
0
2
0
1
0
0
0
0
0
0
0
0
0
1
1
0
2
0
1
1
0
0
0
0
0
0
0
0
0
1
0
2
0
1
0
0
0
0
0
0
0
0
0
0
1
0
2
0
1
0
0
0
0
0
0
0
0
0
0
1
0
2
0
1
1
0
0
0
0
0
0
?
0
1
1
0
2
0
1
0
0
0
0
0
0
0
1
0
0
1
0
2
0
1
0
0
1
0
0
0
11
40
69
0
0
?
?
1
1
0
1
1
0
1
0
?
1
1
?
1
1
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0
0
?
Character number
12 13 14 15
41 42 43 44
70 71 72 73
0
0
0
1
0
0
1
0
0
?
?
1
?
?
?
1
0
0 (01) 0
2
?
?
0
0
0
?
1
0
0
1
0
2
?
?
0
0
0
0
1
0
0
1
0
1
?
?
0
?
?
?
1
0
0 (01) 0
2
?
?
0
?
?
?
1
0
0
1
0
2
?
?
0
0
0
0
1
?
0
0
0
0
?
?
0
0
0
0
1
?
0
0
0
0
?
?
0
0
0
0
1
0
0
1
0
0
?
?
0
0
0
0
1
0
0
1
0
2
?
?
0
0
0
0
1
0
0
1
0
0
?
?
1
0
0
0
1
0
0
1
0
1
?
?
0
0
0
0
1
0
0
1
0
1
?
?
1
0
0
0
1
0
0
1
0
1
?
?
0
0
0
0
1
0
0
1
0
1
?
?
0
0
0
0
1
0
0
1
0
1
?
?
0
0
0
0
1
0
0
1
0
1
?
?
0
0
0
0
1
0
0
1
0
0
?
?
1
0
0
0
1
0
0
1
0
1
?
?
0
0
0
0
1
0
0
1
0
2
?
?
0
0
0
0
1
0
0
1
0
1
?
?
0
0
0
0
1
0
0
1
0
1
?
?
1
0
0
0
1
0
0
1
0
0
?
?
1
0
0
0
1
0
0
1
0
1
?
?
?
0
0
0
1
0
0
1
0
0
?
?
0
16
45
74
(01)
0
0
2
1
?
3
1
?
3
1
?
2
1
?
2
1
?
2
0
?
2
0
?
0
0
?
2
0
?
3
0
0
2
0
?
2
0
0
2
0
?
2
0
?
2
0
?
2
0
?
0
0
0
3
0
?
2
0
?
2
0
?
2
0
1
3
0
0
?
0
?
2
0
?
17
46
75
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
0
0
2
0
0
2
0
0
2
0
0
2
0
0
(02)
0
0
0
0
0
0
0
0
0
0
0
2
0
0
0
0
0
0
0
0
0
1
0
0
18
47
76
1
0
?
1
0
?
1
0
?
1
0
?
(12)
0
?
1
0
?
1
0
?
1
0
?
1
0
?
2
1
?
2
1
?
2
0
1
2
0
1
?
0
1
2
0
1
2
0
1
(01)
0
1
1
0
?
2
1
?
2
1
?
?
0
1
1
0
?
1
1
?
?
0
?
1
0
?
19 20 21 22 23
48 49 50 51 52
77 78 79 80 81
2
0
0
1
2
0
0
?
0
?
?
0
3
?
1
0
0
0
1
2
0
1
1
0
?
?
0
3
0
1
0
0
0 (12) 2
0
1
1
0
?
?
0
3
0
1
0
0
0
2
2
0
1
1
0
?
?
0
3
0
1
0
0
0
2
2
0
1
1
0
?
?
0
3
0
1
0 (01) 0
1
2
0
1
1
0
?
?
0
3
0
1
2
0
0
1
2
0
1
0
0
?
?
0
3
0
1
2
0
0
1
2
0
1
0
0
?
?
0
3
0
1
1
0
0
1 (23)
0
1
1
0
?
?
0
3
0
1
2
0
0
1
2
0
1
1
0
?
?
0
3
?
1
0
0
0
1 (12)
0
1
1
0
?
?
0
3
0
1
1
0
0
1
2
0
1
1
0
?
?
0
3
? (01)
1
0
0
1
2
0
1
1
0
?
1
0
3
0
1
?
0
0
1
2
0
1
1
0
?
1
0
3
?
1
2
0
0
1
2
0
1
1
0
?
1
0
3
0 (01)
2
0
0
1
2
0
1
1
0
?
1
0
3
0 (01)
0
0
0
1
1
0
1
1
0
?
1
0
3
?
1
0
0
0 (12) (12)
0 (12) 1
0
?
?
0
3
0
1
2
0
0
1
2
0
1
1
0
?
?
0
3
?
1
2
0
0 (12) 2
0
1
1
0
?
?
0
3
?
1
1
0
0
1
2
0
1
1
0
?
1
0
3
0
0
0
0
0
1
1
0
1
1
0
?
?
0
3
0
1
0
0
0
1
2
0
1
1
0
?
?
0
3
0
1
?
0
0
?
2
0
1
1
0
?
?
0
3
?
1
0
0
0
1
2
0
1
1
0
?
?
0
3
0
1
24
53
82
2
0
?
2
(01)
0
2
2
0
2
2
0
(12)
1
0
(12)
(01)
0
2
1
0
2
1
0
2
0
1
2
2
?
0
0
1
(01)
1
1
2
0
0
2
0
?
1
(01)
1
1
(01)
1
0
1
?
(12)
0
1
2
1
?
2
2
?
1
0
?
0
0
1
0
0
1
1
0
?
1
2
0
25
54
83
1
0
0
1
0
0
?
0
0
?
0
0
1
0
0
1
0
0
?
0
0
?
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
1
0
(01)
0
0
1
0
0
1
?
0
?
0
0
26 27 28
55 56 57
84 85 86
0 (03) 0
1
1 (23)
1
1
1
2
2
2
1 (02) 1
0
?
?
3
2
2
1 (012) 1
1
1
1
3
2
2
1 (012) 1
1
1
1
2
2
2
1 (02) 1
0
?
?
(23) 2
2
1 (02) 1
0
?
?
0
0
0
1 (12) 3
1
1
1
0
0
0
1 (12) 3
1
1
1
0
0 (03)
1
1 (23)
1
1
1
0
2
2
1
1
?
1
1
1
3
0
0
1
1
?
1
1
1
0
2
2
1 (12) 1
1
1
1
0 (27) 2
1
1 (23)
1
1
1
0
2
2
1
1
?
1
1
1
0
2
2
1 (12) 1
1
1
1
0
2
2
1 (12) 1
1
1
1
0
2
2
1
1
?
1
1
1
0
3
3
1
1 (23)
1
1
1
0
2
2
1
1
?
1
1
1
0 (27) 2
1
1
?
1
1
1
0
2
2
1
2
1
1
1
?
0
3
3
1
1 (23)
1
1
1
0
2
2
1
1
?
1
1
1
?
2
2
1
1
?
1
1
?
0
0
0
1 (01) ?
1
1
1
29
58
87
0
1
0
1
(13)
?
1
(13)
0
1
(13)
0
0
(13)
?
0
1
?
0
3
?
0
3
?
0
1
0
0
1
0
0
1
0
1
1
0
1
1
0
?
1
0
0
1
0
0
1
0
1
1
0
(02)
1
0
0
1
0
0
1
0
?
1
?
2
2
0
0
1
0
?
1
?
0
(12)
?
Xiphopteris
conjunctisora
Campyloneuron
angustifolium
Microgramma
percussa
Microgramma
squamulosa
Pecluma alfredii
Polypodium
glycyrrhiza
1
2
30 31
59 60
(23) 1
0
0
1
0
3
0
0
0
?
1
3
0
0
0
?
1
3
0
0
0
?
1
3
1
0
0
0
1
(02) 1
0
0
?
1
3
32
61
0
3
?
0
0
2
0
0
2
0
0
2
0
2
0
0
2
0
4
33
62
0
0
0
0
?
1
0
?
1
0
0
1
0
1
1
0
1
1
5
34
63
0
0
1
1
?
0
1
?
0
1
?
0
0
1
0
1
1
0
6
35
64
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
7
36
65
1
2
1
1
0
0
1
0
0
1
0
0
1
0
1
1
0
0
8
9 10
37 38 39
66 67 68
0
1
1
0
1
0
0
0
0
0 (12) 1
0
1
0
0
0
0
1
1
0
0
1
0
0
0
1
1
1
0
0
1
0
0
0
0
0
1
1
0
1
1
0
0
0
0
2
0
0
1
0
0
0
0
11
40
69
0
0
?
1
1
?
1
1
0
1
1
?
1
1
?
1
1
?
Character number
12 13 14 15
41 42 43 44
70 71 72 73
0
0
0
1
0
0
1
0
0
?
?
0
1
1
0
1
?
1
1
2
1
2
2
0
1
0
1
1
?
1
1
2
1
0
0
2
1
0
0
1
?
1
1
2
1
0
0
0
0
0
0
1
0
0
1
0
?
?
?
1
1
0
0
1
0
3
0
2
0
0
1
0
16
45
74
2
0
?
3
1
?
2
1
?
2
1
?
2
1
?
0
1
?
17
46
75
1
0
0
0
0
0
0
3
0
0
3
0
0
0
1
0
1
0
18
47
76
1
0
?
0
0
?
0
0
?
0
0
?
0
0
1
0
0
?
19
48
77
0
0
?
0
0
?
0
0
?
0
0
?
0
0
0
0
0
?
20 21 22
49 50 51
78 79 80
1
0
2
1
1
0
0
3
0
1
1
2
2
1
1
0
0
0
1
1
2
2
1
1
0
1
0
1
1
2
2
1
1
0
1
0
0
0
3
2
1
0
0 (01) 0
0
0
1
2
1 (01)
0
1
0
23
52
81
?
?
1
1
0
0
2
1
0
2
1
0
2
?
0
2
?
0
24
53
82
2
?
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
25
54
83
0
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
26 27 28
55 56 57
84 85 86
2
0
0
1 (01) ?
1
1
1
0
0
0
0
?
?
0
0
0
0
1
1
0
?
?
0
0
0
0
1
1
0
?
?
0
0
0
0
1
1
1
1
0
0
0
0
0
0
0
0
?
?
0
0
0
29
58
87
0
1
?
0
?
0
0
?
0
0
?
0
0
?
0
0
?
0

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53 (2) May 2004: 415428

Ranker & al. Phylogeny and evolution of grammitid ferns

Phylogeny and evolution of grammitid ferns (Grammitidaceae): a case of

KEYWORDS: atpB, Grammitidaceae, homoplasy, molecular phylogeny, morphology, rbcL.

indicated that grammitids nest within Polypodiaceae

Ranker & al. Phylogeny and evolution of grammitid ferns

53 (2) May 2004: 415428

Fig. 1. Examples of endemic Hawaiian grammitids. A, Adenophorus hymenophylloides; B, Grammitis tenella; C, A.

family, whereas Copeland (1947) recognized 12, and

53 (2) May 2004: 415428

MATERIALS AND METHODS

Ranker & al. Phylogeny and evolution of grammitid ferns

logenetic analyses as implemented in PAUP* 4.0b10

Ranker & al. Phylogeny and evolution of grammitid ferns

53 (2) May 2004: 415428

Huelsenbeck & Ronquist, 2001) and employing the

imply a transformation series, i.e., the coding is neutral.

bp were invariant, 388 bp were variable, and 234 bp were

53 (2) May 2004: 415428

Ranker & al. Phylogeny and evolution of grammitid ferns

Ranker & al. Phylogeny and evolution of grammitid ferns

53 (2) May 2004: 415428

53 (2) May 2004: 415428

Ranker & al. Phylogeny and evolution of grammitid ferns

were: (1) Clade Ic was sister to Ib, rather than to Ia + Ib;

strongly supported as being monophyletic, including

Ranker & al. Phylogeny and evolution of grammitid ferns

53 (2) May 2004: 415428

reported for most groups of grammitids; however, n = 33

Hawaiian Islands. Clade I includes three smaller, mostly

53 (2) May 2004: 415428

isolated Old World species that appear to be outliers of

Ranker & al. Phylogeny and evolution of grammitid ferns

Ranker & al. Phylogeny and evolution of grammitid ferns

53 (2) May 2004: 415428

light brown to tan

Enterosora. These are two of the three primarily New

unicellular, setulose, and eglandular. Similar hairs appear

53 (2) May 2004: 415428

Ranker & al. Phylogeny and evolution of grammitid ferns

dorsally & ventrally

the distal end of the annulus. Terpsichore species in

Ranker & al. Phylogeny and evolution of grammitid ferns

53 (2) May 2004: 415428

broadly useful among Neotropical taxa to distinguish

53 (2) May 2004: 415428

Ranker & al. Phylogeny and evolution of grammitid ferns

Bull. 19: 1115.

Ranker & al. Phylogeny and evolution of grammitid ferns

53 (2) May 2004: 415428

Parris, B. S. 1983. A taxonomic revision of the genus

Parsimony (*and Other Methods). Version 4. Sinauer

53 (2) May 2004: Appendices

Ranker & al. Phylogeny and evolution of grammitid ferns

Collector, number, and herbarium

GenBank acc. no.

Ranker 1114 (COLO)

A. Rojas & al. 3298 (UC)

A. Rojas & E. Fletes 3145 (UC)

Moraga & Rojas 501 (UC)

A. Rojas 3005 (MO)

Salino 4538 (BHCB, UC)

A. Rojas 3022 & M. Mata (UC)

P. Sanchez-B. 183 (UC)