American Fern Journal 97(3):127–135 (2007)

Hyalotrichopteris is Indeed
a Campyloneurum (Polypodiaceae)
HANS-PETER KREIER
Albrecht-von-HallerInstitute of Plant Sciences, Georg-August University Göttingen, Untere
Karspüle 3, 37073 Göttingen, Germany
ALEXANDER FCO. ROJAS ALVARADO
Jardan Botanico Lancaster, Universidad de Costa Rica, Cartago, Costa Rica
ALAN R. SMITH
University Herbarium, University of California, Berkeley, CA 94720-2465, USA
HARALD SCHNEIDER1
Albrecht-von-Haller Institute of Plant Sciences, Georg-August University Göttingen, Untere
Karspüle 3, 37073 Göttingen, Germany

ABSTRACT.—The relationships of the rare Mesoamerican fern Campyloneurum anetioides are

inferred by comparing sequences of trnLUAA-trnFGAA intergenic spacer of the plastid genome. In
the past, this taxon was either treated as the single member of the genus Hyalotrichopteris or as part
of the diverse Neotropical genus Campyloneurum. Analyses of the cpDNA give unambiguous
support to the taxonomic placement of this species within Campyloneurum. The closest relatives
within the genus Campyloneurum are currently unknown because limited taxon sampling and
variation of the cpDNA sequences do not allow to elucidate this question. However, we can
conclude that C. anetioides is unlikely the derivative of an early separation within Campylo-
neurum.

Relationships of Hyalotrichopteris anetioides (H. Christ) W. H. Wagner have

been controversial. The species was first described in Polypodium by Christ
(1909), and placed in its own genus, Hyalotricha (H. anetioides (H. Christ)
Copel.), by Copeland (1953). Hyalotricha Copel. is a later homonym of
Hyalotricha Dennis, and so the former has now been renamed Hyalotrichop-
teris W. H. Wagner (Wagner, 1978). Copeland (1953) indicated possible
relationships of Hyalotricha to grammitid ferns, Polypodiaceae s.s., Vittar-
iaceae, and to ‘‘Aspidiaceae’’ (Tectaria Cav. and Ctenitis (C. Chr.) C. Chr.), but
favored the last suggestion. On the basis of stomatal structure, van Cotthem
(1970) suggested an alliance of Hyalotricha to Vittariaceae. Crabbe et al. (1975)
placed Hyalotricha in the Grammitidaceae, near Loxogramme (Blume) C. Presl,
without further comment. Pichi Sermolli (1977) postulated closest affinities of
Hyalotricha to Goniophlebium C. Presl (5 Serpocaulon; Smith et al., 2006) and
a largely American group of polypod genera (his Polypodiaceae s.s.) that also
included Campyloneurum C. Presl. Wagner and Farrar (1976), in the most
careful examination of the morphology of Hyalotricha to date, accepted its

1
Corresponding Author: Current Address: Botany Department, Natural History Museum, SW7
5BD London, UK. E-mail: [email protected] or [email protected]
128 AMERICAN FERN JOURNAL: VOLUME 97 NUMBER 3 (2007)

generic distinctness, and adduced strong evidence for its inclusion in

subfamily Polypodioideae of the Polypodiaceae (rather than with the
grammitid ferns, or Vittariaceae, or ‘‘Aspidiaceae’’); however, Wagner and
Farrar did not speculate on the generic level affinities of Hyalotricha, nor did
they mention Campyloneurum as a possible relative. The later genus includes
about 50 species occurring throughout South America, Mesoamerica, the
Caribbean, to Florida in the North (Tryon and Tryon, 1982a, b; León, 1992,
1995).
Most recently, American authors have regarded Hyalotrichopteris anetioides
(5 Campyloneurum anetioides (H. Christ) R. M. Tryon & A. F. Tryon) either as
a member of Campyloneurum (Tryon and Tryon, 1982a, b; León, 1992, 1995),
or have accepted its generic distinctness as Hyalotrichopteris (Wagner, 1978;
Lellinger, 1988, 1989). Lellinger (1988, 1989) considered Hyalotrichopteris
a satellite of Campyloneurum, based on characters believed to be rare in the
latter genus, namely, the small (not exceeding 10 cm) spathulate leaves,
a single excurrent free veinlet in each areole, marginal free veinlets, and
multicellular branched hairs Hyalotrichopteris (Lellinger, 1988). In her
revision of Campyloneurum, León (1992, 1995) noted that these supposed
unique features in Hyalotrichopteris are shared by species of Campyloneurum.
Relatively small leaves (but still considerably larger than in Hyalotrichopteris)
are found in C. chrysopodum (Klotzsch) Fée and C. falcoideum (Kuhn ex
Hieron.) M. Meyer ex Lellinger, whereas branched multicellular hairs occur
also in C. aphanophlebium (Kunze) T. Moore and C. repens (Aubl.) C. Presl.
León considered the presence of Campyloneurum-like venation in Hyalo-
trichopteris anetioides as a critical character supporting close relationships of
this species to Campyloneurum. Excurrent free veinlets along the blade
margins occur in several species of Campyloneurum such as C. brevifolium
(Lodd. ex Link) Link, as well as in juvenile fronds of larger species, and
excurrent veinlets within non-costal areoles occur in C. aphanophlebium and
C. falcoideum. Based on the shared morphological features, León (1992)
considered C. aphanophlebium as the putative closest extant species to
Hyalotrichopteris/Campyloneurum anetioides.
The introduction of DNA sequence data as a further marker to discover
relationships has greatly improved our understanding of the phylogeny of
polygrammoid ferns, as well as other groups of ferns (Schneider et al., 2004a,
b). These studies have resolved many uncertainties about the interpretation of
relationships and the underlying morphological evidence. As an example, an
extensive study on the grammitids found strong support for several genera that
were recently proposed based on morphological evidence, which appears to be
remarkable considering the high degree of homoplasy in these ferns (Ranker et
al., 2004). Other genera such as Microsorum Link and Polypodium L. were
found to be polyphyletic as currently defined (Schneider et al. 2004a, b, 2006a,
b; Smith et al., 2006). Using nucleotide sequence variation of the chloroplast
genome (cpDNA), relationships of several enigmatic polypod genera have
recently been discovered, the Himalayan genus Gymnogrammitis Griff.
(Schneider et al., 2002), Malesian Thylacopteris Kunze ex J.Sm. (Schneider
KREIER ET AL.: HYALOTRICHOPTERIS IS A CAMPYLONEURUM 129

et al., 2004a), and southern South American Synammia C. Presl (Schneider et

al., 2006b). cpDNA data (Janssen and Schneider, 2005) have also confirmed the
generic concept of Aglaomorpha Schott proposed by Roos (1985), who utilized
exclusively morphological evidence.
Existing studies on polypod ferns (Polypodiaceae) have included one or
more samples of Campyloneurum, and almost all other genera in the family,
but sequence data has been unavailable for Hyalotrichopteris anetioides. In
this study, we explore the relationships of this narrowly distributed Costa
Rican and Panamanian species using cpDNA data.

MATERIALS AND METHODS

A sample of the rare Hyalotrichopteris anetioides was collected by one of us
(A.F.R.A.) in Costa Rica. All samples representing other species of Campylo-
neurum or other genera included were collected either in Botanical Gardens or
provided by colleagues. Some sequences were obtained from Genbank (see
Table 1 for accession numbers). The sampling strategy follows the phyloge-
netic hypotheses outlined in previous studies (Schneider et al., 2004a, b,
2006a, b). Besides H. anetioides, we collected sequences for eight species of
Campyloneurum, one species of Niphidium (sister genus of Campyloneurum)
and seven species of Microgramma, the sister to the clade comprising
Campyloneurum and Niphidium. Microgramma was assigned as the outgroup
clade. Table 1 gives information on vouchers and Genbank accession numbers.
Sequences of the trnLUAA-trnFGAA intergenic spacer (trnL-F IGS) region were
obtained using primers and protocols described in previous studies (Haufler et
al., 2003; Schneider et al., 2004a, b; Janssen and Schneider, 2005). Sequencing
was carried out on a MegaBACE 1000 capillary sequencer using DYEnamic ET
Primer DNA Sequencing Reagent (Amersham Biosciences, UK). All sequences
were assembled and manually aligned using TreV (Staden Package, http://
sourceforge.net/projects/staden). The final alignment was adjusted manually
in MacClade 4.0 (Maddison and Maddison, 2000). Ambiguously aligned
regions were excluded from all analyses. The few scattered indels did not
include any phylogenetic information concerning the relationships of H.
anatioides.
Maximum parsimony (MP) and maximum likelihood (ML) analyses were
performed with PAUP* version 4.0b10 (Swofford, 2000). Model and
parameters were selected using the hierarchical likelihood ratio test and the
Akaike information criterion as implemented in Modeltest (Posada and
Crandall, 1998). Selected model and parameters were implemented in PAUP
before conducting ML analysis. MP and ML analyses were calculated using the
heuristic mode with 1000 (MP) and 100 (ML) random-addition-sequence
replicates, TBR branch swapping, and MULTREES on. Bootstrap values were
calculated for both (MP and ML) with 1000 bootstrap replicates, each with 10
random-addition-sequence replicates, TBR branch swapping and MULTREES
on. ML analyses were also performed using PHYML (Guindon and Gascuel,
2003; Guindon et al., 2005). In addition, we explored the robustness of the
130 AMERICAN FERN JOURNAL: VOLUME 97 NUMBER 3 (2007)

TABLE 1. Taxon sampling: For each taxon the following information is given: voucher information
including herbarium of deposition, accession number in the fern DNA database (http://
www.pryerlab.net), and Genbank accession numbers

Campyloneurum C. Presl
Campyloneurum anetioides (H. Christ) R. M. Tryon & A. F. Tryon: Costa Rica; Rojas 6281 (CR);
EF104510 – Campyloneurum angustifolium (Sw.) Fée: Costa Rica; Chisaki & Carter 1004 (UC
1618523); AY083647 – Campyloneurum aphanophlebium (Kunze) T. Moore: Bolivia; Acebey 772
(UC 1735546); EF104511 – Campyloneurum asplundii (C. Chr.) Ching: Mexico; Kessler s.n.
(GOET); EF104512 – Campyloneurum brevifolium (Lodd. ex Link) Link: cult. Göttingen, Old Bot.
Garden; Kreier s.n. (GOET); EF104513 – Campyloneurum chlorolepis Alston: Venezuela; Smith
1159 (UC 1487590); AY083648 – Campyloneurum phyllitidis (L.) C. Presl: cult. Berlin, Bot. Garden
(Acc. 017-59-74-63); Schuettpelz 612 (GOET); EF104514 – Campyloneurum sphenodes (Kunze ex
Klotzsch) Fée: Costa Rica; Horich s.n. (UC 1617915); AY083649 – Campyloneurum xalapense Fée:
(1) cult. Göttingen, Old Bot. Garden; Kreier s.n. (GOET); EF104515 & (2) Mexico; Lautner L02-41
(GOET); EF104516.
Niphidium J. Sm.
Niphidium nidulare (Rosenst.) Lellinger: Costa Rica; Ranker 1831 (COLO); EF104519.
Microgramma C. Presl
Microgramma bifrons (Hook.) Lellinger: Peru; van der Werff 18062 (MO); DQ642224 –
Microgramma latevagans (Maxon & C. Chr.) Lellinger: Bolivia; Jimenez 1285 (LPB); EF104517 –
Microgramma mauritiana (Willd.) Tardieu:m (1) cult. Zürich Bot. Garden; Kreier s.n. (GOET);
DQ642225 & (2) cult. Zürich Bot. Garden; Kreier s.n. (GOET); DQ642226 – Microgramma nitida (J.
Sm.) A. R. Sm.: Mexico; Krömer 2678 (GOET); EF104518 – Microgramma squamulosa (Kaulf.) de la
Sota: cult. Zürich Bot. Garden; Kreier s.n. (GOET); DQ642228 – Microgramma tecta (Kaulf.) Alston:
cult. RBG Edinburgh (Acc. 19875234); Schneider s.n. (E); DQ642230 – Microgramma vacciniifolia
(Langsd. & Fisch.) Copel.: cult. Charles Alford; Kreier s.n. (GOET); DQ642231

results using the neighbor net algorithm as implemented in Splitstree 4 (Huson

and Bryant, 2006), using different settings to evaluate the robustness of the
analyses (e.g., different distance measures such as LogDets and number of
dimensions calculated).
Analyses were performed in two steps. In the first step, we compared the
trnL-F sequence of Hyalotrichopteris anetioides with all currently available
sequences of this cpDNA region available in GenBank and with additional
available data. This comparison was made using Blastn (Altschul et al., 1990)
and by performing maximum parsimony analyses and maximum likelihood
analyses with all available data. In the second step, we inferred the
relationships of this species with a data set including only Campyloneurum,
Microgramma, and Niphidium species. The later dataset was inferred
employing MP, ML and neighbor net analyses.

RESULTS
A sequence of 342 base pairs was generated for H. anetioides, which was in
the range of length variation (ca. 340 to 370 bp) currently known for the trnL-F
IGS in Campyloneuron and relatives. The data set consisted of 230 included
characters of which 37 were parsimonious informative and 35 were variable
but non-parsimonious informative. The number of informative sites dropped
KREIER ET AL.: HYALOTRICHOPTERIS IS A CAMPYLONEURUM 131

to 16 considering only Campyloneurum and H. anetioides. Both Blastn and the

maximum parsimony analyses using all available trnL-F IGS sequences of
Polypodiaceae indicated Hyalotrichopteris anetioides as a member of
Campyloneurum (results not shown). Phylogenetic reconstructions using
maximum parsimony, maximum likelihood, and network approaches found
this species embedded in Campyloneurum (Figs. 1, 2). The maximum
parsimony analysis resulted in 45 most parsimonious trees with a length of
96 steps (consistency index [CI] 5 0.7288, homoplasy index [HI] 5 0.2712,
retention index [RI] 5 0.8730, and rescaled consistency index [RC] 5 0.7275).
Campyloneurum was monophyletic in all 45 most parsimonious trees, and
Hyalotrichopteris anetioides was sister to Campyloneurum angustifolium in
89% of 45 most parsimonious trees. A single unambiguous character state
change separated H. anetioides from C. angustifolium and in turn one
unambiguous character state change characterized this clade. The Campylo-
neurum clade had a bootstrap value of 98% (Fig. 1) and the clade was
characterized by six unambiguous character state changes. Only two well
supported clades were found in all most parsimonious trees: a clade
comprising the two collections of C. xalapense and a clade consisting of C.
asplundii and C. chlorolepis). Maximum likelihood analysis found a single
tree (Fig. 1) with -ln 5 834.16001, using the TVM model with freqA 5 0.3331,
freqC 5 0.1636, freqG 5 0.1842, freqT 5 0.3190, R(a) 5 1.9602, R(b) 5 3.3907,
R(c) 5 0.0435, R(d) 5 1.3489, R(e) 5 3.3987, R(f) 5 1.000. Based on the
hierarchical likelihood ratio test, this model and parameters were best,
whereas the Akaike information criterion preferred a slightly different model
TVM + I. Analyses with the alternative model found the same topology.
The ML tree was fully resolved, but bootstrap values .75% were found only
for clades present in the strict consensus tree of the most parsimonious trees
obtained in the maximum parsimony analysis. Hyalotrichopteris anetioides
was found to be sister to C. angustifolium (Fig. 1), but alternative relationships
were found in the Splitgraph analysis (Fig. 2).

DISCUSSION

Status of Hyalotrichopteris anetioides.—Our data suggest that Hyalotrichop-

teris anetioides nests within Campyloneurum. To separate it generically
renders Campyloneurum paraphyletic. Hyalotrichopteris anetioides shares
with species of Campyloneurum the principal characters defining the genus,
namely areolate venation with costal areoles containing one excurrent free
veinlet and non-costal areoles containing (1–)2–5 excurrent free veinlets (León,
1992). Characters used by Lellinger (1988, 1989) to separate Hyalotrichopteris
are either not restricted to H. anetioides, such as the branched multicellular
hairs (which also occur in Campyloneurum aphanophlebium), or are unreli-
able, such as frond size and habitat. The unusually large number of marginal
excurrent free veinlets, which is uncommon in Campyloneurum in general,
may correlate with the reduction in leaf size.
132 AMERICAN FERN JOURNAL: VOLUME 97 NUMBER 3 (2007)

FIG. 1. Phylogram generated in a maximum likelihood analysis of the trnL-F IGS data set.
Numbers above branches correspond to ML bootstrap values, and numbers below branches
correspond to MP bootstrap values. BS values $ 75% are given. Thickened lines indicate branches
present in the strict consensus of 45 most parsimonious trees. Abbreviations: Cam 5
Campyloneurum, Hya 5 Hyalotrichopteris, Mgr 5 Microgramma, Nip 5 Niphidium.
KREIER ET AL.: HYALOTRICHOPTERIS IS A CAMPYLONEURUM 133

FIG. 2. Splitstree generated with the Neighbornet algorithm and uncorrected P-distances. Settings:
weight modified as least squares, and maximum dimensions set to four. Statistics; nsSplits 5 45,
total weight 5 0.40393516. Abbreviations as in Fig. 1.

Relationships within Campyloneurum.—León (1992) tentatively divided

Campyloneurum, a genus of ca. 50 species, into ten groups, of which six are
represented in our study. She placed C. anetioides in the C. aphanophlebium
group, comprising only these two species. This group is characterized by
having specialized branched, multicellular hairs. Further shared character-
istics are the presence of undivided, primary, non-costal areoles, free veinlets
along the margin, and medial sori.
Our results (Fig. 1) suggest that there is some affinity between Campylo-
neurum anetioides and C. aphanophlebium, but other species such as C.
angustifolium may have closer relationships. Our current taxon sampling is
insufficient to explore the relationships among species of the genus
Campyloneurum as demonstrated in the low resolution in our maximum
parsimony analyses and the many alternative topologies in neighbor net
analyses. We cannot draw any conclusions about the monophyly of some
proposed groups within the genus Campyloneurum and/or the species
delimitation of some putative closely related species in the C. angustifolium
group. These questions require a much denser taxonomic sampling. This
restriction also applies to the position of taxa at the earliest split within the
134 AMERICAN FERN JOURNAL: VOLUME 97 NUMBER 3 (2007)

genus, although C. phyllitidis is not only the sister to the remaining genus but
also shows similarities in its morphology with the sister genus Niphidium
such as prominent costa and veins on the abaxial leaf surface, coriaceous
blades, and erect fronds. Our data set contains only 36 parsimony-informative
sites from a total of 230 included characters. This low percentage (about 16%)
of informative sites may indicate that Campyloneurum has diverged and
radiated relatively recently. Similar low percentage of informative and/or
variable sites were reported in other studies employing the trnL-F IGS region to
elucidate relationships between closely related species of Polypodiaceae
(Haufler et al., 2003; Janssen and Schneider, 2005; Schneider et al., 2006a, b;
Smith et al., 2006) and Aspleniaceae (Schneider et al., 2005). In the
Aspleniaceae study, divergence times were estimated for a lineage of
asplenioid ferns (Schneider et al., 2005) and the results indicated a di-
versification in the Miocene corresponding to a level of sequence divergence as
observed in Campyloneurum. The exact timing of this genus diversification is
currently unknown but the distribution of the genus suggests putative
correlations with geological and climatic changes in the upper Tertiary
shaping the extant plant diversity of South and Central America. Future
studies require not only denser sampling of species within the genus and but
also sampling of more variable markers.

ACKNOWLEDGMENTS
This investigation was supported by the German Science Foundation (DFG) grant to HS
(SCHN758/2-1 and SCHN758/2-2) as part of the Schwerpunkt program (SPP 1127) ‘‘Radiations –
origin of biological diversity’’. We thank also two anonymous reviewers for the kind suggestions
that helped to improve this publication.

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