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Taxonomic studies of Pteridophyta VII. A revision of the genus Stegnogramma emend

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... endemic to the New World (Watkins and Farrar 2005;PPG I 2016). Stegnogramma is characterized by having lowermost pinnae that are not reduced or only a little reduced, distal pinnae that are adnate to the rachis, exindusiate sori extending along the veins, non-glandular hairs on the sporangial capsules, echinate spores, and a chromosome number of n 5 36 (Iwatsuki 1963;Hirabayashi 1969;Holttum 1982;Tsai and Shieh 1985;Kurizono 1987;Smith 1990). The genus has sometimes been segregated into distinct genera, i.e. ...
... Ching 1963;Shing et al. 1999;Lin et al. 2013), or sections (e.g. Iwatsuki 1963), mainly based on venation characters. Among them, the Dictyocline group (sensu genus Dictyocline in Lin et al. 2013) can be distinguished from other groups by its entirely reticulate venation, forming irregularly-joined areoles, sometimes with simple or forked included veinlets (Ching 1963;Iwatsuki 1963). ...
... Iwatsuki 1963), mainly based on venation characters. Among them, the Dictyocline group (sensu genus Dictyocline in Lin et al. 2013) can be distinguished from other groups by its entirely reticulate venation, forming irregularly-joined areoles, sometimes with simple or forked included veinlets (Ching 1963;Iwatsuki 1963). Four species are usually recognized in the Dictyocline group: S. griffithii (T.Moore) K.Iwats., S. mingchegensis (Ching) X.C.Zhang & L.J.He, S. sagittifolia (Ching) L.J.He & X.C.Zhang, and S. wilfordii (Hook.) ...
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We describe a new species of Stegnogramma (Thelypteridaceae; Polypodiales) as Stegnogramma australis. Morphologically, it is most similar to S. griffithii, sharing the character of pinnate fronds with ternate terminal pinnae. However, S. australis can be distinguished from S. griffithii by having more free pinna pairs, shorter distal stipe hairs, and fewer included veinlets in the laminar areoles. A description, photographs of key characters, and the known distribution of the new species are presented. We also infer genome size and reproductive mode of the new species using flow cytometry. In addition, the occurrence of S. aspidioides in Vietnam is confirmed and a key to all known species of Stegnogramma in Vietnam is given.
... Taxonomically, Stegnogramma s.l. can be further divided into four sections (Iwatsuki, 1963): sect. Stegnogramma, sect. ...
... Stegnogramma, Dictyocline and Haplogramma include species with anastomosing venation. Of these, Dictyocline is the most unusual in that areoles of Dictyocline form irregularly reticulated hexagons with included veinlets (Ching, 1936Iwatsuki, 1963;Lin et al., 2013). In contrast, leaves of sect. ...
... In contrast, leaves of sect. Leptogramma species are free-veined (Ching, 1936Iwatsuki, 1963;Lin et al., 2013). In spite of such morphological variation, current phylogenetic evidence supporting previous generic or infrageneric delimitation is weak. ...
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The thelypteroid fern genus Stegnogramma s.l. contains around 18–35 species and has a global, cross‐continental distribution ranging from tropical to temperate regions. Several genera and infrageneric sections have been recognized previously in Stegnogramma s.l., but their phylogenetic relationships are still unclear. In this study, we present a global phylogeny of Stegnogramma s.l. with the most comprehensive sampling to date and aim to pinpoint the phylogenetic positions of biogeographically and taxonomically important taxa. Based on the reconstructed historical biogeography and character evolution, we propose a new (infra)generic classification and discuss the diversification of Stegnogramma s.l. in a biogeographical context. New names or combinations are made for 12 (infra)species, including transferring the monotypic species of Craspedosorus to Leptogramma. Finally, we discuss a possible link between leaf architecture and ecological adaptation, and hypothesize that the increase in leaf dissection and free‐vein proportion is an adaptive feature to cool climates in Stegnogramma s.l.
... In presence of raphides in the leaf epidermis, .4. cardiophyllum resembles the condition in sect. Hymenasplenium excepting .4. cheilosorum (Iwatsuki, 1975). Raphides are also lacking in .4. normale which Holttum (1955) considered to be related to sect. ...
... Raphides are also lacking in .4. normale which Holttum (1955) considered to be related to sect. Hymenasplenium but Iwatsuki (1975) excluded from the section. The chromosome number of 2n= 156 indicates that .4. cardiophyllum is a tetraploid with a basic number of x-39. ...
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Asplenium cardiophyllum is a morphologically unusual species with simple leaves and anastomosing venation, and is often placed in the segregate genusBoniniella. To determine its systematic position, character comparisons were made of vascular anatomy, raphides in leaf epidermis, chromosome number and perispore of this species and those ofAsplenium sect.Hymenasplenium. Asplenium cardiophyllum conforms with sect.Hymenasplenium in its dorsiventral dictyostele, the presence of raphides, a chromosome number of 2n=156 (x=39), and lophate peristore with spinulate projections on the lumina. We therefore propose to includeA. cardiophyllum in that section.
... One of Bir's collections from Lachen, North Sikkim, in PAN was identified by Fraser-Jenkins (1997) as Stegnogramma leptogrammoides and Fraser-Jenkins (2008b) accepted this (as T. kingii) as a distinct species, though he pointed out that it was probably only temporarily endemic as to be expected in Tibet etc. Dixit (1984) mistakenly gave its distribution as "Eastern India, Japan", not realising that Sikkim is in the Central Himalayan region and presumably thinking that as Iwatsuki had described it might be in Japan, but this was corrected by Chandra (2000). Inspie of initial report on endemic nature of T. kingii or S. leptogrammoides from Sikkim (Iwatsuki 1963, Reed 1968,Jarrett 1983, Fraser Jenkins 1997) this fern has not been mentioned in the Indian Red-lists (Nayar and Sastry, 1983–1990; Rao et al., 2003), along with many other threatened pteridophytes that were not brought to attention. It was also inadvertently omitted by Chandra et al. (2008). ...
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A rare and little known, endemic fern Thelypteris kingii C.F.Reed was first described from a 19th Century collection from an unspecified locality in what was then "British Sikkim", now Darjeeling District, India. Recently a small population of the plant was rediscovered by the author from present-day Sikkim State. Herbarium records and field study reveal that this species is very rare and under great threat of extinction due to natural and man-made impact. This species is therefore little known to pteridologists and is readily confused with the similar, but common species, T. mollissima (Kunze) N.Thapa as well as with another S. E. Asian species T. leptogrammoides (Ross.) C.F. Reed due to nomenclatural similarities or being later homonymous. Although only known so far from India, and thus an endemic Indian species, its presence in Tibet and China etc. is probably to be expected. A brief history of its discovery, taxonomy, morphology, habitat and ecology is presented here to assist taxonomists and conservation biologists with its identification and conservation. It has been identified as Endangered and Globally Threatened and efforts have also been made by the present author to include it in the forthcoming volume of the Red Data Book of Indian plants.
... This result was expected, as we did not increase the sampling of these groups. For a more detailed discussion about the morphological rationale regarding the recognition of Dictyocline, Leptogramma, and Stegnogramma as a single genus, see Iwatsuki (1963) and He and Zhang (2012). ...
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Thelypteridaceae is one of the largest fern families, having about 950 species and a cosmopolitan distribution but with most species occurring in tropical and subtropical regions. Its generic classification remains controversial, with different authors recognizing from one up to 32 genera. Phylogenetic relationships within the family have not been exhaustively studied, but previous studies have confirmed the monophyly of the lineage. Thus far, sampling has been inadequate for establishing a robust hypothesis of infrafamilial relationships within the family. In order to understand phylogenetic relationships within Thelypteridaceae and thus to improve generic reclassification, we expand the molecular sampling, including new samples of Old World taxa and, especially, many additional neotropical representatives. We also explore the monophyly of exclusively or mostly neotropical genera Amauropelta, Goniopteris, Meniscium, and Steiropteris. Our sampling includes 70 taxa and 121 newly generated sequences from two plastid genomic regions (rps4-trnS and trnL-trnF), plus 71 rps4 and 70 trnL-trnF sequences from GenBank. These data resulted in a concatenated matrix of 1980 molecular characters for 144 taxa. The combined data set was analyzed using maximum parsimony and bayesian inference of phylogeny. Our results are consistent with the general topological structure found in previous studies, including two main lineages within the family: phegopteroid and thelypteroid. The thelypteroid lineage comprises two clades; one of these included the segregates Metathelypteris, Coryphopteris, and Amauropelta (including part of Parathelypteris), whereas the other comprises all segregates of Cyclosorus s.l., such as Goniopteris, Meniscium, and Steiropteris (including Thelypteris polypodioides, previously incertae sedis). The three mainly neotropical segregates were found to be monophyletic but nested in a broadly defined Cyclosorus. The fourth mainly neotropical segregate, Amauropelta, was found to comprise species considered to be part of the Parathelypteris. In Old World thelypteroids, which correspond to nearly half the diversity in the family, an increase in sampling is still needed to resolve relationships and circumscription of genera, particularly in the christelloid clade (i.e., Amphineuron, Chingia, Christella, Pneumatopteris, Pronephrium, and Sphaerostephanos). Based on currently available knowledge, we propose the recognition of 16 genera in the family.
... These should be investigated to examine the monophyly of S. gymnocarpa. S. tottoides is distributed in Taiwan and Fukien and is distinguished from S. pozoi by the long-attenuate fronds (Iwatsuki 1963). S. tottoides from Taiwan formed a clade with Atami-1 and Atami-2 types (Fig. 3). ...
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In Japanese Stegnogramma pozoi subsp. mollissima (Fisher ex Kunze) K. Iwats. there is the intrasubspecific variation among rbcL sequences. Northern and southern plants are genetically differentiated for maternally inherited cpDNA. In the present study we examined allozyme polymorphisms to test the hypothesis that northern and southern plants may be separate species. Based on allozyme data, the degree of gene flow among populations was estimated to be large. The artificial crossing experiments between cpDNA haplotypes also suggested that isolation has not developed among these cpDNA haplotypes. However, interpopulation genetic differentiation in cpDNA was observed even in the small area at the foot of Mt. Hakone, and the cpDNA haplotypes appear to have different habitat preferences.
Article
The northwest region of Yunnan province (southwest China), belonging to the Pan-Himalaya, is a hotspot of biodiversity. We report Stegnogramma leptogrammoides from Yunnan as a new discovery for China. This species was identified based on morphological comparison with the type specimens of all similar species, and DNA barcoding was performed to obtain additional evidence. As it was previously recorded to be endemic to Sikkim, the new record provides a case for confirming floristic affinities between Sikkim and Yunnan within the Pan-Himalaya at the species level.
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The northwest region of Yunnan province (southwest China), belonging to the Pan-Himalaya, is a hotspot of biodiversity. We report Stegnogramma leptogrammoides from Yunnan as a new discovery for China. This species was identified based on morphological comparison with the type specimens of all similar species, and DNA barcoding was performed to obtain additional evidence. As it was previously recorded to be endemic to Sikkim, the new record provides a case for confirming floristic affinities between Sikkim and Yunnan within the Pan-Himalaya at the species level.
Chapter
A worldwide family of five genera including Cyclosorus, with c 650 species, one of the largest genera among the derived ferns, treated in 20 subgenera. The classification of the family is not resolved since several systems are proposed recognizing different genera and subgeneric groups. The system of A.R. Smith, followed here, is a tentative one that treats most of the taxa under Cyclosorus. Some species in this or other subgenera have incomplete authors because they have not yet been formally transferred to the genus in which they are treated.
Chapter
Terrestrial (rarely epiphytic) ferns with branched or unbranched stems, these long- to short-creeping, suberect, or erect and trunk-like, bearing scales at apex, these entire, usually pubescent along margin and sometimes surface, basally attached. Vascular system a radially symmetrical dictyostele. Petioles non-articulate, scaly at base, with two lateral pneumatic lines along the length, in cross-section nearly always with two hippocampus-shaped vascular strands at base, these fusing into a U-shaped strand distally. Lamina simple to pinnate to most often pinnate + pinnatifid, catadromous, infrequently twice-pinnate or more divided (but never with basitonically enlarged pinnae), monomorphic or less often dimorphic. Pinnae usually numerous, gradually reduced distally into a confluent, lobed, apical segment, sometimes with a conform apical division. Indument of lamina and axes almost always of hairs, these variously unicellular or multicellular, acicular, hooked, furcate, or stellate (Iwatsuki 1962), sometimes also with sessile or stalked glands, occasionally also with scales. Rachis adaxially grooved, this groove not continuous with adaxial grooves of costae. Aerophores often present at bases of pinnae abaxially, these swollen or elongate. Veins simple or less often forked, reaching the margin or not, ± cdistinctly catadromously pinnately arranged in ultimate segments; free, connivent below the sinus, or united to form one to many regular areoles between costa and pinna margin, then with (usually only) ex-current free veinlets. Sporangia nearly always in discrete sori, these orbicular to occasionally elongate, dorsal (occasionally submarginal) on the veins, with a short to long stalk three cells in cross-section, with a well-developed vertical annulus, dehiscing by a horizontal slit, maturation mixed. Indusia present or absent, if present then reniform or infrequently athyrioid, often very reduced and obscured in mature sori. Spores 64 (infrequently 32) per sporangium, nearly always bilateral with a monolete scar (tetrahedral with trilete scar in Trigonospora),non-chlorophyllous; perispore variously ornamented (reticulate, winged, spinulose); gametophytes surficial, cordate, often with marginal glands or hairs; x=27, 29–36.
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