Species Of Diplazium Research Articles

Common Pteridophytes of Shimla (Himachal Pradesh) Part II : Athyriaceae

The general characters of the family, genus and species alongwith their keys for identification have been presented. Six species of Athyrium, namely Athyrium flabellulatum, A. foliolosum, A. pectinatum, A. rupicola, A. schimperi, A. setiferum and A. strigillosum; two species of Deparia, namely Deparia boryana and D. japonica and two species of Diplazium, namely Diplazium esculentum and D. maximum of family Athyriaceae have been dealt with.

Phylogenetics of Hawaiian Diplazium (Athyriaceae: Polypodiales)

Hawaii is home to four species in the fern genus Diplazium: Diplazium arnottii, D. esculentum, D. molokaiense, and D. sandwichianum. Three are endemic to the Hawaiian Islands, and one is introduced and naturalized (D. esculentum). They vary in frequency, from very abundant (D. sandwichianum) to rare and critically endangered (D. molokaiense). Prior to this work, the phylogenetic relationships of only D. esculentum had been estimated due to lack of information available about these species. In this study, we inferred phylogenetic relationships of the Hawaiian Diplazium based on six chloroplast regions – atpA, atpB, matK, rbcL, rps4+rps4–trnS IGS, and trnL intron+trnL–trnF IGS. We downloaded these plastid markers from GenBank for an additional 83 Diplazium species and two Athyrium species, and used the latter as outgroups. The resulting phylogeny inferred from combined data indicated that D. arnottii and D. sandwichianum are sister taxa, likely with origins in the Paleotropics. Morphologically, the two species can be distinguished by leaf dissection and lengths of sori. Diplazium molokaiense is sister to the Asian D. heterocarpum (among our sampled taxa). Diplazium molokaiense is distinguished from other species in the clade by its large sori and longer fronds. Our results reveal the phylogenetic placements of three Diplazium species and corroborate the placement of a fourth species.

<p><strong>Rediscovery of <em>Diplazium procumbens</em> (Athyriaceae) in Peninsular Malaysia after more than 70 years, with notes on the morphology, spore ornamentation and anatomy</strong></p>

Recent floristic sampling in Fraser’s Hill, Pahang, Malaysia resulted in recollection of a rare Diplazium species in Peninsular Malaysia, D. procumbens, a poorly known species which was first described by R.E. Holttum in 1940. In this paper, we provide comprehensive morphological and anatomical description of the species based on our thorough examination of specimens including the type, and designate a lectotype and isolectotype. We also studied the spore ornamentation of D. procumbens using scanning electron microscopy (SEM) and discovered that the spores have folds and non-fenestrate perine. Descriptions of the habitat and ecology are also presented.

Species delineation of the genus Diplazium Swartz (Athyriaceae) using leaf architecture characters

The present study was conducted to delineate Diplazium Swartz species based on leaf architecture. Using PAleontological STatistics (PAST), a cluster and Principal Component Analysis of leaf architecture characters of 27 selected Diplazium species at the Philippine National Herbarium (PNH) was done. The dendogram (cophenetic correlation = 0.8436) and principal component analysis supported the four clusters of Diplazium using leaf architecture characters. At Gower distance of 0.25, Diplazium species were categorized as: Cluster 1 (Cladodromous – short stalked, stout and massive 1° vein); Cluster 2 (Reticulodromous – long stalked, moderate 1° vein); Cluster 3 (Craspedodromous – long stalked, stout to massive 1° vein); and Cluster 4 (Craspedodromous – short stalked, stout to massive 1° vein). The unifying characters were apex shape, base symmetry and 1° vein category, while the significant differentiating characters were 2° vein angle of divergence and variation in the 2° vein angle of divergence, 3° vein category, 3° vein angle of divergence, variation in 3° vein angle of divergence, 3° vein spacing and lobation. The successful delineation ofDiplazium species proved that leaf architecture can be a good taxonomic marker and could be an alternative way of identifying species in the absence of sori.

Infrageneric Revision of the Fern Genus Deparia (Athyriaceae, Aspleniineae, Polypodiales)

Current molecular phylogenetic analyses support the monophyly and circumscription of the athyrioid fern genus Deparia (Athyriaceae), which includes previously recognized genera including Athyriopsis, ×Depazium, Dictyodroma, Dryoathyrium (= Parathyrium), Lunathyrium, and Neotriblemma (= Triblemma Ching), and ×Neotribleparia. This broad generic concept has been adopted in several recent taxonomic treatments, including the Pteridophyte Phylogeny Group I. However, the infrageneric taxonomy of Deparia still needs further revision. In this study, we provide a new infrageneric classification with five sections and three subsections based on the phylogenetic evidence. We recognize two new sections, sect. Dictyodroma and sect. Erectus, and two new subsections, sect. Deparia subsect. Caespites and subsect. Athyriopsis. In addition, we provide a key to the sections and subsections, a table of diagnostic characters, new combinations for three Deparia and one Diplazium species, and a list of species assigned to the sections/subsections. Hybrids derived from crosses between the sections and subsections are also noted.

Systematics ofDiplazium laffanianum(Athyriaceae), a Fern Species Endemic to Bermuda

Diplazium laffanianum was last collected from its natural habitat in Bermuda in 1905. Now the species is considered extinct in the wild as it exists only in greenhouses, tissue cultures, and as a few young reintroduced sporophytes. For the last 13 years, a collaborative team of researchers from the Government of Bermuda and the Lab for Rare & Endangered Plants at Omaha's Henry Doorly Zoo in Nebraska, USA has been working to bring this species back from the brink of extinction. In this project, we developed propagation methods for D. laffanianum, and we studied molecular and morphological characters to determine the phylogenetic placement of D. laffanianum. We sequenced plastid DNA from D. laffanianum using high-throughput sequencing, downloaded six plastid markers (atpA, atpB, matK, rbcL, rps rps4-trnS, and trnL intron trnL-F) from GenBank for 21 Diplazium and outgroup taxa, and inferred a phylogeny. We examined 185 collections of five Diplazium species closely related to D. laffanianum, and we scored 15 morphological characters to test whether D. laffanianum is a distinct species. The phylogeny placed D. laffanianum in the D. cristatum group, suggesting a Neotropical ancestor, and two morphological characters distinguished D. laffanianum from close relatives. Our results provide evidence for the phylogenetic placement of D. laffanianum, and we are actively propagating and reintroducing individuals to the wild, aiding conservation of the species.

Novelties in <I>Diplazium</I> (Athyriaceae) from South America

Abstract Two new species of Diplazium from Brazil, D. adnatum and D. brachycarpum , are described and illustrated. A new name and status— Diplazium peruvianum —is proposed for Diplazium celtidifolium var. puberuletum. A list of characters differentiating these taxa and a key to the 2–3-pinnate-pinnatifid species of Diplazium from Brazil is provided.

Diplazium fimbriatum (Athyriaceae), a New Species from Brazil

Abstract A new species, Diplazium fimbriatum (Athyriaceae), is described and illustrated. It is endemic to the humid montane forests of eastern Brazil, a region known for its high level of endemism and species richness. A comparative table to distinguish it from similar species of Diplazium occurring in Brazil is provided.

Diplazium condorense (Athyriaceae), a new species from Ecuador

Abstract A new species of Diplazium, D. condorense from Ecuador, with free veins and concolorous blade scales, is described and illustrated. It is apparently most closely related to species of Callipteris, having anastomosing veins and scales with black margins and teeth.

Lectotypification of Several Names Currently Placed in Diplazium (Woodsiaceae)

Abstract Lectotypifications are made for the following names that apply to species of Diplazium in the Old World: D. atratum, D. conterminum, D. crinipes, D. megaphyllum, D. polypodioides var. vestitum, D. sechellarum, and D. sikkimense. The types of these names have scales with black borders and bifid marginal teeth, a scale type characteristic of the diplazioid segregate Callipteris.

Diplazium subsinuatum and Di. tomitaroanum should be Moved to Deparia According to Molecular, Morphological, and Cytological Characters

sequence data revealed that the putative intergeneric hybrid, Diplazium tomitaroanum Masam. belongs in Deparia, as also does Diplazium subsinuatum (Wall, ex Hook, et Grev.) Tagawa, one of the putative parents. An examination of rachis, scale and spore morphology, and chromosome data provide support for this placement. We propose a new taxonomic treatment of the two Diplazium species as Deparia.

Organization of the vascular system in the stems ofDiplaziumandBlechnum (Filicales)

Detailed analysis of the three‐dimensional vascular organization in species of Diplazium and Blechnum indicates that these ferns possess reticulate (dictyostelic) vascular systems that closely reflect the helical phyllotaxis of the shoot. In each species, the vascular pattern shows a specific relationship to the phyllotaxis, so that the phyllotactic fraction can be determined by examination of the number of cauline vascular bundles (meristeles) in cross section of the stem. The number of meristeles in a cross section equals the denominator of the phyllotactic fraction, i.e., the number of foliar orthostichies on the stem. The same numerical relationship also exists in the eusteles of seed plants between the number of axial (sympodial) stem bundles and the phyllotaxis. There is a further parallel between the three‐dimensional reticulate pattern of fern dictyosteles and the reticulate patterns that characterize some herbaceous dicotyledons. However, the hypothesized separate origins of seed plant eusteles and fern dictyosteles from protostelic precursors preclude any direct homologies between these similar patterns. The parallel evolution of presumably more physiologically efficient reticulate systems in herbaceous seed plants and in ferns that have only a primary plant body is noteworthy. The similar relationships between the primary stem vascular patterns and phyllotaxy in both ferns and seed plants further emphasize the likely similarity of the morphogenetic events that occur at the shoot apex in these taxonomically disparate groups.

Organization of the Vascular System in the Stems of Diplazium and Blechnum (Filicales)

Detailed analysis of the three-dimensional vascular organization in species of Diplazium and Blechnum indicates that these ferns possess reticulate (dictyostelic) vascular systems that closely reflect the helical phyllotaxis of the shoot. In each species, the vascular pattern shows a specific relationship to the phyllotaxis, so that the phyllotactic fraction can be determined by examination of the number of cauline vascular bundles (meristeles) in cross section of the stem. The number of meristeles in a cross section equals the denominator of the phyllotactic fraction, i.e., the number of foliar orthostichies on the stem. The same numerical relationship also exists in the eusteles of seed plants between the number of axial (sympodial) stem bundles and the phyllotaxis. There is a further parallel between the three-dimensional reticulate pattern of fern dictyosteles and the reticulate patterns that characterize some herbaceous dicotyledons. However, the hypothesized separate origins of seed plant eusteles and fern dictyosteles from protostelic precursors preclude any direct homologies between these similar patterns. The parallel evolution of presumably more physiologically efficient reticulate systems in herbaceous seed plants and in ferns that have only a primary plant body is noteworthy. The similar relationships between the primary stem vascular patterns and phyllotaxy in both ferns and seed plants further emphasize the likely similarity of the morphogenetic events that occur at the shoot apex in these taxonomically disparate groups.

A comparison of the flavonoids in Athyriaceae and Aspleniaceae

Flavonoid patterns were examined for 18 species of Athyriaceae and 15 species of Aspleniaceae, mainly of Malaysian origin. In the Athyriaceae, flavonol 3- O-glycosides based on both quercetin and kaempferol were the most common flavonoids found, and additionally in some species of Diplazium and Deparia flavone C-glycosides were detected, based on apigenin. Flavonoid data supports the generic separation of Athyrium, Diplazium and Deparia based on morphological and other characters. In the Aspleniaceae, the flavonoid profiles are more complex. As to flavonols, kaempferol derivatives appeared to predominate, especially the 3,7-glycosides, but 3,4′-diglycosides and 3,7,4′-triglycosides were also detected. O-Methylated kaempferol glycosides were found in A. marinum. Flavone C-glycosides were based on both luteolin and apigenin, and the C-sugars were sometimes O-glycosylated. In three species of Asplenium, flavone O-glycosides were encountered, including scutellarein 6- O-glucoside in A. belangeri. Thus the overall tendency of the flavonoid chemistry of the Aspleniaceae is different from that of the Athyriaceae, and this supports the taxonomic separation of the two families.

Taxonomic and Phytogeographic Relationships of Diplazium flavoviride, D. pycnocarpon, and Diplaziopsis

Diplazium, a worldwide genus of about 400 species, is the largest genus of the athyrioid group, although Ching (1964b) placed the majority of species of Diplazium in the genus Allantodia. Several monotypic or oligospecific genera, such as Callipteris, Dictyodroma, Diplaziopsis, Hemidictyum, and Monomelangium, are often segregated from Diplazium and recognized on the basis of a few specialized characters (Copeland, 1947; Crabbe et al., 1975; Ching, 1978). All of these are closely related to and probably derived from Diplazium. Diplaziopsis, comprising two species, is defined by its pinnate leaves with a conform terminal pinna, thin lamina texture, anastomosing veins, and sausage-like sori (Copeland, 1947). Ching (1964a) added two new species from China to Diplaziopsis.

Diplazium delitescens and the Neotropical Species of Asplenium sect. Hymenasplenium

Since the original description of Diplazium delitescens Maxon, no one has seriously questioned its generic disposition. However, a preliminary study of herbarium material of this uncommon neotropical species suggests that it might be better placed in Asplenium. Maxon relied chiefly on the back-to-back arrangement of linear sori in ascribing this species to Diplazium. In contrast, spleenworts generally have only single, linear sori on the ultimate veins. Although soral arrangement is the primary (and usually most reliable) character used to separate the two genera, I report here a survey of additional characters that provide good evidence for transferring D. delitescens to Asplenium. Sporangia.-The species of Asplenium consistently have one-rowed sporangial stalks, at least at the base (Bower, 1928, p. 140; Tardieu-Blot, 1932, p. 363) and sporangial capsules that often split divaricately at the tip of the fully extended or backwardly flexed annulus. Diplazium delitescens possesses both these characteristics (Fig. 1). Other species of Diplazium have shorter, twoor three-rowed sporangial stalks, with annuli often not extended and capsules not splitting divaricately at the tip, e.g., D. werckleanum Christ (Fig. 2). I have been unable to find any reference to this distinctive type of sporangial opening in Asplenium, but I believe it may be a very useful character in distinguishing asplenioid species from other groups of ferns. The number of annular cells inAsplenium is generally higher than in Diplazium. Tardieu-Blot (1932, p. 364) reported 15-20 annular cells for species of Diplazium and 20-25 annular cells for species of Asplenium. Copeland (1947, p. 147) listed Diplazium (inAthyrium) as having an of 12-20 (commonly 16) thickened cells, whereas Asplenium was described as having an usually of 20-28 cells. I have made five counts each onA. laetum Swartz (Breedlove 33893, DS), A. abscissum Willd. (Breedlove 22178, DS), A. harpeodes Kunze (Breedlove 22504, DS), and A. auriculatum Swartz (Breedlove 22506, DS); these species averaged 23, 20, 21, and 20 annulus cells per sporangium, respectively. Diplazium franconis Liebm. (Breedlove 22421, DS), D. lonchophyllum Kunze (Breedlove 22461, DS), D. acutale Fee (Breedlove 22760, DS), and D. werckleanum Christ (Breedlove 33664, DS) all averaged 15 annular cells per sporangium. Diplazium delitescens (Breedlove 33853, DS) averaged 21 annular cells per sporangium, which agrees with Asplenium. Spores.-Diplazium delitescens spores have numerous, sharp folds in the perispore (Fig. 3), and are similar to, or even indistinguishable from, spores of several species of Asplenium from southern Mexico and Central America, e.g., A.