A taxonomic revision of Tectaria (Tectariaceae) from New Caledonia

  • South China Botanical Garden, Chinese Academy of Sciences
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A taxonomic revision of Tectaria from New Caledonia is presented here based on a thorough study of herbarium specimens from New Caledonia and adjacent Pacific islands. Six species of Tectaria are presently recognized in New Caledonia. Tectaria kouniensis and T. pseudosinuata are reduced to T. dissecta and T. sinuata, respectively. The morphological limit between T. seemannii and T. sinuata and the confusion of specimen citations in previous accounts are clarified. Apart from T. dissecta, the other five species, i.e., T. lifuensis, T. moorei, T. seemannii, T. sinuata, and T. vieillardii, are mainly distributed in New Caledonia, with only T. seemannii and T. sinuata recently reported also from Vanuatu. A key to species, typification of accepted names and relevant synonyms, and brief comments on relationships of species are provided. This article is protected by copyright. All rights reserved.

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The fern genus Tectaria (Tectariaceae) is morphologically diverse and difficult in terms of recognizing species and species groups. To infer the systematic positions of some species and identity-unknown collections with special morphological characters, we conducted phylogenetic analyses based on sequences of five plastid regions (atpB, ndhF plus ndhF-trnL, rbcL, rps16-matK plus matK, and trnL-F). Three analysis methods (maximum parsimony, maximum likelihood, and Bayesian inference) were used to reconstruct the phylogeny of Tectaria. The most surprising result is that T. menyanthidis, T. ternata, and T. variabilis are revealed to represent a distinct lineage from Tectaria, which should be called Polydictyum and is supported as sister to Pteridrys. Other accessions of Tectaria are well resolved into four major clades, which is consistent with the results of previous studies. Of the four clades, Clade II (T. subtriphylla group) is unpredictable, with morphologically very diverse species clustered there, and is supposed to be a minor evolutionary line within Tectaria in the Old World. In addition, the position of the climbing genus Arthropteris and the utility of molecular data in recognizing species of Tectaria are briefly discussed. As a conclusion we formally reinstate the genus Polydictyum by providing diagnostic characters, key to species, nomenclature, and information of detailed distribution and habit for the currently known three species.
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The fern genus Tectaria has generally been placed in the family Tectariaceae or in subfamily Tectarioideae (placed in Dennstaedtiaceae, Dryopteridaceae or Polypodiaceae), both of which have been variously circumscribed in the past. Here we study for the first time the phylogenetic relationships of the associated genera Hypoderris (endemic to the Caribbean), Cionidium (endemic to New Caledonia) and Pseudotectaria (endemic to Madagascar and Comoros) using DNA sequence data. Based on a broad sampling of 72 species of eupolypods I (= Polypodiaceae sensu lato) and three plastid DNA regions (atpA, rbcL and the trnL-F intergenic spacer) we were able to place the three previously unsampled genera. Our results show that Cionidium, like Ctenitopsis, Fadyenia, Hemigramma and Quercifilix, is embedded in Tectaria, and the monophyly of Tectaria is therefore corroborated only if these segregate genera are included. Hypoderris is sister to Tectaria brauniana and together they are sister to Triplophyllum, which was found to be monophyletic. Despite their morphological similarity with Tectaria, the genera Pleocnemia and Pseudotectaria were placed in Dryopteridoideae. Polypodiaceae subfamily Tectarioideae (former family Tectariaceae) is hereby defined to include Arthropteris, Hypoderris, Pteridrys, Tectaria and Triplophyllum. Aenigmopteris may also belong here, but this genus remains unsampled. PDF available at:
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Aim To investigate areas of endemism in New Caledonia and their relationship with tectonic history. Location New Caledonia, south-west Pacific. Methods Panbiogeographical analysis. Results Biogeographical patterns within New Caledonia are described and illustrated with reference to eight terranes and ten centres of endemism. The basement terranes make up a centre of endemism for taxa including Amborella, the basal angiosperm. Three of the terranes that accreted to the basement in the Eocene (high-pressure metamorphic terrane, ultramafic nappe and Loyalty Ridge) have their own endemics. Main conclusions New Caledonia is not simply a fragment of Gondwana but, like New Zealand and New Guinea, is a complex mosaic of allochthonous terranes. The four New Caledonian basement terranes were all formed from island arc-derived and arc-associated material (including ophiolites) which accumulated in the pre-Pacific Ocean, not in Gondwana. They amalgamated and were accreted to Gondwana (eastern Australia) in the Late Jurassic/Early Cretaceous, but in the Late Cretaceous they separated from Australia with the opening of the Tasman Sea and break-up of Gondwana. An Eocene collision of the basement terranes with an island arc to the north-east – possibly the Loyalty Ridge – is of special biogeographical interest in connection with New Caledonia–central Pacific affinities. The Loyalty–Three Kings Ridge has had a separate history from that of the Norfolk Ridge/New Caledonia, although both now run in parallel between Vanuatu and New Zealand. The South Loyalty Basin opened between Grande Terre and the Loyalty Ridge in the Cretaceous and attained a width of 750 km. However, it was almost completely destroyed by subduction in the Eocene which brought the Loyalty Ridge and Grande Terre together again, after 30 Myr of separation. The tectonic history is reflected in the strong biogeographical differences between Grande Terre and the Loyalty Islands. Many Loyalty Islands taxa are widespread in the Pacific but do not occur on Grande Terre, and many Grande Terre/Australian groups are not on the Loyalty Islands. The Loyalty Islands are young (2 Myr old) but they are merely the currently emergent parts of the Loyalty Ridge whose ancestor arcs have a history of volcanism dating back to the Cretaceous. Old taxa endemic to the young Loyalty Ridge islands persist over geological time as a dynamic metapopulation surviving in situ on the individually ephemeral islands and atolls found around subduction zones. The current Loyalty Islands, like the Grande Terre terranes, have inherited their biota from previous islands. On Grande Terre, the ultramafic terrane was emplaced on Grande Terre in the Eocene (about the same time as the collision with the island arc). The very diverse endemic flora on the ultramafics may have been inherited by the obducting nappe from prior base-rich habitat in the region, including the mafic Poya terrane and the limestones typical of arc and intraplate volcanic islands.
The pantropical genus Tectaria Cav. (Tectariaceae) is one of the largest fern genera. It has been estimated to contain ca. 210 mostly tropical species in Africa, the Americas, Asia, and the Indian Ocean and Pacific Ocean islands. Tectaria had perhaps been the most confusing fern genus in terms of its circumscription and phylogeny. Recent studies have recircumscribed Tectaria and resolved the relationships within the genus. However, no efforts have been made to propose an infrageneric classification of the genus based on molecular and morphological evidence. In the present study, we synthesize chloroplast and nuclear DNA evidence, morphology, and/or distribution information and divide Tectaria into four subgenera: Tectaria subg. Ctenitopsis (Ching ex Tardieu & C. Chr.) Li Bing Zhang & Liang Zhang (comb. & stat. nov.), Tectaria subg. Phlebiogonium (Fée) Li Bing Zhang & Liang Zhang (comb. & stat. nov.), Tectaria subg. Tectaria, and Tectaria subg. Tectaridium (Copel.) Li Bing Zhang & Liang Zhang (comb. & stat. nov.). The latter three are further divided into seven, two, and two sections, respectively. Sixteen generic names are synonymized to individual infrageneric taxa, while four generic names are only synonymized to Tectaria because of inadequate data. A key to the infrageneric taxa is given. A nomenclatural account of each infrageneric taxon is provided.
Tectaria (Tectariaceae) is one of the most confusing fern genus in terms of its circumscription and phylogeny. Since its original description, a number of genera had been moved into or related with this genus, while others had been segregated from it. Tectaria is also among the largest fern genera, comprising 150–210 mostly tropical species. Previous molecular studies have been far from comprehensive (sampling no more than 76 accessions from 52 species), limited in geographic scope (mainly restricted to Asia), and based exclusively on plastid markers. In the present study, DNA sequences of eight plastid and one nuclear marker of 360 accessions representing ca. 130 species of Tectaria, ca. 36 species of six non-Tectaria genera in Tectariaceae, 12 species of Davalliaceae, Oleandraceae, and Polypodiaceae, and 13 species of Lomariopsidaceae were used to infer a phylogeny with maximum likelihood, Bayesian inference, and maximum parsimony approaches. Our major results include: (1) the most recently proposed circumscription of Tectaria is strongly supported as monophyletic; (2) the genera Lenda, Microbrochis, Phlebiogonium, and Sagenia, sampled here for the first time, are resolved as part of Tectaria; (3) four superclades representing early splits in Tectaria are identified, with the Old World species being sister to the New World species; (4) 12 well-supported major clades in Tectaria are revealed, differing from one another in molecular, morphological, and geographical features; (5) evolution of 13 morphological characters is inferred in a phylogenetic context and morphological synapomorphies of various clades are identified; and in particular (6) free venation in Tectaria is inferred to be repeatedly derived from anastomosing venation, an evolutionary phenomenon not documented previously in vascular plants in a phylogenetic context based on both plastid and nuclear evidence.
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