Corresponding author: Thaís Elias Almeida (
Academic editor: B. Leon
Almeida TE, Salino A, Dubuisson J-Y, Hennequin S (2017)
One of the most problematic circumscriptions is that of
One species that still contributes to this uncertainty is
The main goal of this study was to employ molecular and morphological data to investigate the relationships of
Fifty species from thirty-two genera (sensu
Total DNA was extracted from field-acquired silica gel-dried or fresh tissues, using the Qiagen DNeasy Plant mini kit (Qiagen Inc., Valencia, CA, USA). PCR amplifications were performed for two chloroplast genome regions:
Polymerase chain reactions were performed in a 20 μL solution containing 1.0 μL of non-diluted genomic DNA template, 2.0 μL of PCR buffer (Qiagen 10 × PCR Buffer), 1.0 μL of DMSO, 1.0 μL of BSA (4 mg/mL), 0.8 μL of dNTPS (10 mM), 0.32 μL (10 μM) of each primer, 0.12 units of
Sequence electropherograms were edited using the STADEN package software (
Datasets were analyzed using maximum likelihood (ML) and Bayesian inference (BI). Maximum likelihood was performed using IQ-TREE (
Selected models and parameter values for data partitions used in this study.
Region | Base frequencies | ||||||||
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Selected model | A | C | G | T | |||||
GTR+G | 0.3204 | 0.1924 | 0.1964 | 0.2908 | |||||
GTR+G | 0.3249 | 0.1545 | 0.1605 | 0.3601 | |||||
SYM+I+G | – | – | – | – | |||||
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0.7965 | 3.5099 | 0.1724 | 0.4720 | 3.0330 | 1 | 0 | 0 | 0.9640 | |
0.9825 | 2.7675 | 0.2147 | 0.7566 | 2.9398 | 1 | 0 | 0 | 0.5615 | |
1.9337 | 8.1010 | 1.1861 | 0.8284 | 11.3400 | 1 | 0.5474 | 0 | 0 |
Taxonomic conclusions were based on the study of specimens from the following herbaria: BHCB, BM, BR, G, GH, K, LPB, M, MO, NY, P, Q, QCA, QCNE, QPLS, US, USM, and USZ. Specimens with barcode are cited with herbarium acronym followed by barcode number. Abbreviation of genera and species followed IPNI (
The final combined dataset presented 2339 bp, 599 from
Inside the neotropical clade,
Majority rule consensus tree resultant from Bayesian inference analyses from combined
The polygrammoid topology recovered in our analyses agrees with previous results from several studies (
In our analyses,
Comparison of character states among
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Long-creeping, branched | Long- to short-creeping, sparingly branched | Short-creeping to erect, usually unbranched | Long- to short-creeping, unbranched | Long- to short-creeping, branched | Long- to short-creeping, branched | Long-creeping, branched |
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Peltate, non-clathrate | Peltate, clathrate | Basifixed, non-clathrate or clathrate, or absent | Basifixed to peltate, non-clathrate | Peltate, non-clathrate | Peltate, non-clathrate, to clathrate at margins | Peltate, non-clathrate |
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Monomorphic | Monomorphic | Monomorphic, or the distal fertile portion modified | Monomorphic | Monomorphic | Monomorphic to dimorphic | Monomorphic to dimorphic |
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Simple | Pinnatifid to pinnate, rarely simple | Simple to 3-pinnate | Pinnatisect to pinnate | Deeply pinnatifid to pinnate | Simple to pinnatifid, rarely pinnate-pinnatifid or more divided | Simple to lobate |
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Scales | Glabrous, trichomes, or scales (confined to costae and rachises) | Trichomes, and sometimes glands | Trichomes | Glabrous or with trichomes | Scales | Glabrous, trichomes |
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Free | Regularly anastomosing (goniophlebioid), areoles with one included veinlet | Usually free, sometimes anastomosing with or without included free veinlets | Free, rarely anastomosing, but never reticulate | Free to anastomosing, with one single included veinlet | Free to anastomosing, areoles with 1-3 free or netted included veins | Anastomosing, with simple included veinlets |
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Round to oblong, 1 row between costa and margins | Round, 1-10 rows between costa and margins | Round to elongate, 1 row between costa and margins, or confluent | Round to oblong, 1 row between costa and margins | Round to oblong, 1-5 rows between costa and margins | Round to oblong, or linear, rarely marginal and coalescing, in 1 row between costa and margins | Round to elongate, 1 row between costa and margins, confluent or forming several irregular rows between costa and margins |
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Peltate, pedicellate, scales | Absent or short 2-3-celled glands | Present or absent | Present, glandular trichomes | Absent or if present, filamentous or branched | Absent, or round peltate, | Trichomes or sessile scales, or absent |
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Bilateral, monolete, verrucate | Bilateral, monolete, verrucate, generally tuberculate, occasionally winged | Tetrahedral-globose, trilete | Bilateral, |
Bilateral, monolete | Bilateral, monolete, shallowly to prominently verrucate | Bilateral, monolete, tuberculate |
1
2
3
4
Sanín (2014,
Grammitid ferns, the lineage sister to the
Our results do not support the inclusion of
Ecuador. Andes de Quito,
Plants epiphytic or epipetric, rarely terrestrial. Rhizomes long-creeping, branched, 0.6–0.9 mm wide, cylindrical, with four vascular bundles; short, perpendicular roots about 5–20 mm long, these regularly spaced, covered with brownish root hairs; rhizome scales 4.3–7.1 mm long, peltate, not clathrate, linear-lanceolate, with elongate cells, the margins entire from the base to the middle and toothed beyond the middle, with 1- or 2-celled marginal teeth, scales concolorous, stramineous and usually darker at the attachment point. Fronds remote, 2.2–4.5 mm apart, articulate, monomorphic. Stipes nearly absent to 55 mm long, 0.4–0.7 mm in diameter, covered with sparse peltate, lanceolate, sessile, non-clathrate, concolorous, stramineous scales, these 1.4–3.5 mm long, darker at the attachment point; phyllopodia darker than stipes. Laminae light green, 5.0–17.0 × 1.0–2.5 cm, simple, chartaceous, linear-lanceate to lanceolate, bases acuminate to attenuate, decurrent in the distal third of the stipe, apices acute to obtuse, laminar surfaces squamose on both sides, scales lanceolate, peltate, sessile, non-clathrate, concolorous, translucent, stramineous, darker at attachment point, slightly erose at bases and entire at apices, scales present also on abaxial and adaxial sides of costae and veins, and on the laminar margins, 1.3–2.6 mm long. Veins free, immersed, obscure, 1–2 furcate, not reaching laminar margins, midribs and lateral veins immersed on both sides of the laminae, not evident. Sori superficial, rounded to oblong, 1.6–2.6 × 1.9–5.0 mm, terminal to subterminal on veins, receptacles elongate, sporangia long-stalked, paraphyses present, scale-like, similar those of the laminar surfaces, peltate, pedicellate, with pedicels as long as those of the sporangia, paraphyses completely covering immature sori. Spores yellow, with verrucate surfaces.
Restricted to central and southern Andes, with known collections from Ecuador, Peru, Bolivia, and Argentina (Fig.
Least Concern (LC -
The generic epithet refers to the most distinctive character of the species, the free venation (Fig.
Morphologically,
Distribution of
This work was supported by FAPEMIG (Fundação de Amparo à Pesquisa do Estado de Minas Gerais through grant CRA-APQ 01599-10, CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico) through grants 478723/2010-5 and 563568/2010, and for productivity grants, all attributed to AS, and the ATM “Biodiversité actuelle et fossile” of the Muséum national d’Histoire naturelle (2012) attributed to SH. CNPq also provided research scholarships to TEA (555226/2010-7 and 202160/2011-4). We thank the Muséum national d’Histoire naturelle de Paris, France (MNHN), the Service de Systématique Moléculaire (SSM - MNHN) and Céline Bonillo for the support and help. L.L. Giacomin for helping with fieldwork and for carefully reviewing the manuscript. A. Field, A. Smith, B. León, D. Sanín, E. Smidt, J. Prado, J.A.N. Batista, and V.A.O. Dittrich for critical reviews and comments. T. Martínez for guiding us in Bolivia; to the curators of all visited herbaria; Claudia Martín for the pictures, and J.T. Mickel for allowing the use of the drawings.
Collection information for voucher specimens and GenBank accession numbers for sequences used in this study. Locality and voucher specimen (herbarium) are given to sequences newly generated in this study. *sequences submitted to GenBank
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