Spore fitness components do not differ between diploid and allotetraploid species of Dryopteris (Dryopteridaceae)

BACKGROUND AND AIMS: Although allopolyploidy is a prevalent speciation mechanism in plants, its adaptive consequences are poorly understood. In addition, the effects of allopolyploidy per se (i.e. hybridization and chromosome doubling) can be confounded with those of subsequent evolutionary divergence between allopolyploids and related diploids. This report assesses whether fern species with the same ploidy level or the same altitudinal distribution have similar germination responses to temperature. The effects of polyploidy on spore abortion and spore size are also investigated, since both traits may have adaptive consequences. METHODS: Three allotetraploid (Dryopteris corleyi, D. filix-mas and D. guanchica) and three related diploid taxa (D. aemula, D. affinis ssp. affinis and D. oreades) were studied. Spores were collected from 24 populations in northern Spain. Four spore traits were determined: abortion percentage, size, germination time and germination percentage. Six incubation temperatures were tested: 8, 15, 20, 25 and 32 degrees C, and alternating 8/15 degrees C. KEY RESULTS: Allotetraploids had bigger spores than diploid progenitors, whereas spore abortion percentages were generally similar. Germination times decreased with increasing temperatures in a wide range of temperatures (8-25 degrees C), although final germination percentages were similar among species irrespective of their ploidy level. Only at low temperature (8 degrees C) did two allotetraploid species reach higher germination percentages than diploid parents. Allotetraploids showed faster germination rates, which would probably give them a competitive advantage over diploid parents. Germination behaviour was not correlated with altitudinal distribution of species. CONCLUSIONS: The results of this study suggest that (i) relative fitness of allopolyploids at sporogenesis does not differ from that of diploid parents and (ii) neither does allopolyploidization involve a change in the success of spore germination.     

中国云南鳞毛蕨属(鳞毛蕨科)13种植物的染色体数目

报道了中国云南产鳞毛蕨科Dryopteridaceae鳞毛蕨属Dryopteris Adanson 13种植物的染色体数目。其中4个种：多雄拉鳞毛蕨D．alpestris（2n=82）、哈巴鳞毛蕨D．habaensis（2n=82）、脉纹鳞毛蕨D．lachoongensis（2n=82）和永自鳞毛蕨D．yungtzeensis（2n=82）的染色体数目为首次报道;无融合三倍体的川西鳞毛蕨Dryopteris rosthornii、无融合四倍体的大羽鳞毛蕨D．wallichiana以及无融合二倍体的栗柄鳞毛蕨D．yoroii为3个新发现的细胞类型。细胞学研究结果支持高山鳞毛蕨组sect．CaespitosaeS．G Lu应该为鳞毛蕨属内一个独立的组;中国喜马拉雅地区的鳞毛蕨属植物中存在着大量的无融合二倍体细胞类型。对蕨类植物中同一植株产生不同倍性孢子的现象进行了讨论。     

Morphological and chromosomal variation of the Dryopteris varia (L.) Kuntze complex (Dryopteridaceae) in Korea

The Dryopteris varia complex is highly variable in morphology, resulting in taxonomic confusion in delimiting taxon boundaries and determining relationships. We have examined the variation in morphology, chromosome number and mode of reproduction of the Korean members of the D. varia complex to clarify their taxonomic identities. Landmark analysis of the leaf blades and pinnae and the principal components analysis of 31 morphological characters revealed seven entities within the D. varia complex in Korea; these comprise D. varia s. str., D. pacifica, D. sacrosancta, D. bissetiana, D. saxifraga, D. saxifragivaria, and the Suak population which is considered to be a new taxon. Mitotic chromosome counts and examination of reproduction modes indicated that D. bissetiana in Korea appears to be agamosporous with diploid (2n = 82) or triploid (2n = 123) chromosome numbers. However, D. saxifraga is sexual diploid or agamosporous triploid, and the other taxa are agamosporous triploid. Dryopteris bissetiana, D. saxifraga and D. saxifragivaria are similar in major morphological characteristics, but show differences in attachment and shape of rachis scales. The results also suggest that agamosporous triploid D. saxifragivaria was probably derived from hybridization between sexual diploid D. saxifraga and D. bissetiana.     

Lectins of Dryopteris Adans fern species

Lectin activity of three ferns species (Dryopteris bushiana Fom., D. laeta (Kom.) C. Chr, D. pseudomas (Wollaston) Holub et Pouzar) has been studied. We used hemagglutination reaction albumin and globulin fractions of fronds and rhizomes extracts with rat erythrocytes. The frond extract of D. pseudomas have shown higher activity as compared with other extracts. The lectin activity of D. laeta leaves was absent. The rhizomes of all three fern species could be characterized as high activity. Based on lectin activity the species were placed as follows order: D. buschiana > D. pseudomas > D. laeta. The dependence with lectin activity and elements (Mn, Fe, Cu, Ni, Zn, Pb, Rb, Sr) content were not found.     

粗茎鳞毛蕨孢子萌发研究

以经过3年低温储藏的粗茎鳞毛蕨孢子为实验材料,从孢子离心、孢子消毒、培养基种类、光质等4方面对孢子萌发进行研究,结果表明:在离心转数≤14 000 r.min-1、离心时间≤30 min条件下,离心处理对孢子萌发基本无影响;对孢子进行1%NaClO水溶液浸泡处理20～30 min为最佳消毒条件;改良Knop’s培养基为最佳孢子萌发培养基;黑暗条件下孢子不能萌发,但是黑暗处理能够明显提高孢子萌发整齐性;红光比白光能促进孢子提早萌发1 d左右,但对提高萌发率效果不显著。     

Molecular phylogenetics and historical biogeography of Hawaiian Dryopteris (Dryopteridaceae)

The fern genus Dryopteris (Dryopteridaceae) is represented in the Hawaiian Islands by 18 endemic taxa and one non-endemic, native species. The goals of this study were to determine whether Dryopteris in Hawai'i is monophyletic and to infer the biogeographical origins of Hawaiian Dryopteris by determining the geographical distributions of their closest living relatives. We sequenced two chloroplast DNA fragments, rbcL and the trnL-F intergenic spacer (IGS), for 18 Hawaiian taxa, 45 non-Hawaiian taxa, and two outgroup species. For individual fragments, we estimated phylogenetic relationships using Bayesian inference and maximum parsimony. We performed a combined analysis of both cpDNA fragments employing Bayesian inference, maximum parsimony, and maximum likelihood. These analyses indicate that Hawaiian Dryopteris is not monophyletic, and that there were at least five separate colonizations of the Hawaiian Islands by different species of dryopteroid ferns, with most of the five groups having closest relatives in SE Asia. The results suggest that one colonizing ancestor, perhaps from SE Asia, gave rise to eight endemic taxa (the glabra group). Another colonizing ancestor, also possibly from SE Asia, gave rise to a group of five endemic taxa (the exindusiate group). Dryopteris fusco-atra and its two varieties, which are endemic to Hawai'i, most likely diversified from a SE Asian ancestor. The Hawaiian endemic Nothoperanema rubiginosum has its closest relatives in SE Asia, and while the remaining two species, D. wallichiana and D. subbipinnata, are sister species, their biogeographical origins could not be determined from these analyses due to the widespread distributions of D. wallichiana and its closest non-Hawaiian relative.     

Anatomy of six species of Dryopteris ferns (Dryopteridaceae) from Mexico

Rhizome and foliar anatomy of the Mexican Dryopteris Adans. species were studied and compared with other Dryopteridaceae and other fern families to identify anatomical features with diagnostic value. The anatomy of rhizome, stipe, and blade is similar in species of the Dryopteris patula complex. The cells with un-lignified, thickened wall, with cap or U-shape around the meristeles belong to the collenchyma, in contrast with other fern families. Dryopteris wallichiana (Spreng.) Hyl. is anatomically distinguished from the other studied species by having more layers of sclerenchyma and meristeles on the stipe, and by the lack of sclereid nests on the rhizome. Dryopteris rossii C. Chr. and D. maxonii Underw. & C. Chr. are characterized by the presence of crystals on the periphery of rhizome nests. D. maxonni and D. wallichiana lack blade glands.     

Origin of Dryopteris shibipedis (Dryopteridaceae), a fern species extinct in the wild

Dryopteris shibipedis was once treated as an extinct species in the Red List (2007) by the Japanese Ministry of the Environment, but 'rediscovered' in the Tsukuba Botanical Garden. To clarify its origin, using 'overlooked' cultivated stocks we analyzed nuclear PgiC intron sequences. As the PgiC genotype of D. shibipedis can be explained by a combination of alleles of D. kinkiensis and those of D. pacifica, the hypothesis of hybrid origin of the species is supported.     

Phylogeny, divergence times, and historical biogeography of New World Dryopteris (Dryopteridaceae)

? Premise of the study: Dryopteris is a large, cosmopolitan fern genus ideal for addressing questions about diversification, biogeography, hybridization, and polyploidy, which have historically been understudied in ferns. We constructed a highly resolved, well-supported phylogeny for New World Dryopteris and used it to investigate biogeographic patterns and divergence times. ? Methods: We analyzed relationships among 97 species of Dryopteris, including taxa from all major biogeographic regions, with analyses based on 5699 aligned nucleotides from seven plastid loci. Phylogenetic analyses used maximum parsimony, maximum likelihood, and Bayesian inference. We conducted divergence time analyses using BEAST and biogeographic analyses using maximum parsimony, maximum likelihood, Bayesian, and S-DIVA approaches. We explored the monophyly of subgenera and sections in the most recent generic classification and of geographic groups of taxa using Templeton tests. ? Key results: The genus Dryopteris arose ca. 42 million years ago (Ma). Most of the Central and South American species form a well-supported clade which arose 32 Ma, but the remaining New World species are the result of multiple, independent dispersal and vicariance events involving Asia, Europe, and Africa over the last 15 Myr. We identified six long-distance dispersal events and three vicariance events in the immediate ancestry of New World species; reconstructions for another four lineages were ambiguous. ? Conclusions: New World Dryopteris are not monophyletic; vicariance has dominated the history of the North American species, while long-distance dispersal prevails in the Central and South American species, a pattern not previously seen in plants.     

Genetic variation in the allotetraploid Dryopteris corleyi (Dryopteridaceae) and its diploid parental species in the Iberian Peninsula

Studies on genetic diversity help us to unveil the evolutionary processes of species and populations and can explain several traits of diploid-polyploid complexes such as their distributions, their breeding systems, and the origin of polyploids. We examined the allozyme variation of Dryopteris aemula and D. oreades, diploid ferns with highly fragmented habitats, and the allotetraploid D. corleyi to (1) analyze the putative relationship between both diploids and the tetraploid, (2) compare the levels of genetic variation among species and determine their causes, and (3) assess the breeding system of these taxa. The allozymic pattern of D. corleyi confirms that it derived from D. aemula and D. oreades. The lack of genetic diversity in D. aemula, a species of lowland habitats, may be due to genetic drift associated with the contraction of populations in the last glaciation. By contrast, the alpine D. oreades had moderate intrapopulation genetic variation, which may derive from the expansion of populations during the last glaciation. In the latter species, low interpopulational variation suggested effective gene flow (spore exchange), and genotype frequencies in Hardy-Weinberg equilibrium indicated cross-fertilization of gametophytes. Evolutionary history appears to be an essential element in the interpretation of genetic variation of highly fragmented populations.     

Leaf morphology and anatomy of Manilkara Adans. (Sapotaceae) from northeastern Brazil

Aiming to complement the group’s taxonomy, the objective of this work is to describe and characterise, searching for morpho-anatomical patterns, the leaf anatomy of the Manilkara species occurring in the northeast region of Brazil. The leaves of all Manilkara species were analysed, using herbaria specimens and collections made in the field by the authors. The characterisation of the leaf blades was performed according to the usual protocols in vegetal anatomy. As diagnosed by anatomical characters, the Manilkara Adans. genus presents petioles with a thick cuticle, a uniseriate epidermis, a sclerenchymatous sheath involving the vascular bundle, and laticifers on the cortical and medullar regions. The anatomical data increase the M. elata and M. huberi synonymisation and show the importance of the anatomy as a complementary tool to the taxonomy based on external morphology.     

Major evolutionary events in the origin and diversification of the fern genus Polystichum (Dryopteridaceae)

Recent advances in molecular systematics of the ferns make it possible to address long-standing questions about classification of the major fern genera, such as the worldwide genus Polystichum (Dryopteridaceae), comprising at least 200 species. In this study we examined rbcL sequences and morphological characters from 55 fern taxa: 34 were from Polystichum and 21 were from other genera in the Dryopteridaceae. We found that Phanerophlebia, possibly including Polystichopsis, is the sister group to Polystichum sensu lato (s.l.), including Cyrtomium. Polystichum as commonly recognized is paraphyletic. Our results lead us to suggest recognizing the clade of earliest diverging Polystichum species as a distinct genus (Cyrtomidictyum) and to continue to recognize Cyrtomium as a separate genus, leaving a monophyletic Polystichum sensu stricto (s.s.). We resolved a tropical American clade and an African clade within Polystichum s.s. However, the resemblance between the once-pinnate, bulb-bearing calciphilic species found in Asia and the West Indies appears to be the result of convergent evolution. Optimizing our morphological character transformations onto the combined phylogeny suggests that the common ancestor of Polystichum s.l. and Phanerophlebia had evolved the common features of the alliance, including ciliate petiole-base scales, once-pinnate fronds, ultimate segments with scarious tips, peltate indusia, and microscales.     

Pollen morphology of the genus Orobanche L. (Orobanchaceae)

A comparative palynological study of 25 taxa of the genus Orobanche L. belonging to section Trionychon Wallr. and section Orobanche L. was carried out. A clear difference between the pollen grains of section Trionychon and section Orobanche was observed. The grains of section Trionychon are typically tricolpate, radially symmetrical, more or less oblate spheroidal, with a microreticulate, or scabrate-perforate sculpture. Section Orobanche on the other hand, had spheroidal, more or less inaperturate pollen grains, although some were tricolpate, but without any clear symmetry. The basic surface sculpture of the pollen in section Orobanche was scabrate to rugulate-perforate. Palynoiogicaliy, six groups can be recognized amongst the species of the genus Orobanche , lor which a key is provided.     

Reticulate evolution on a global scale: a nuclear phylogeny for New World Dryopteris (Dryopteridaceae)

Reticulate, or non-bifurcating, evolution is now recognized as an important phenomenon shaping the histories of many organisms. It appears to be particularly common in plants, especially in ferns, which have relatively few barriers to intra- and interspecific hybridization. Reticulate evolutionary patterns have been recognized in many fern groups, though very few have been studied rigorously using modern molecular phylogenetic techniques in order to determine the causes of the reticulate patterns. In the current study, we examine patterns of branching and reticulate evolution in the genus Dryopteris, the woodferns. The North American members of this group have long been recognized as a classic example of reticulate evolution in plants, and we extend analysis of the genus to all 30 species in the New World, as well as numerous taxa from other regions. We employ sequence data from the plastid and nuclear genomes and use maximum parsimony (MP), maximum likelihood (ML), Bayesian inference (BI), and divergence time analyses to explore the relationships of New World Dryopteris to other regions and to reconstruct the timing and events which may have led to taxa displaying reticulate rather than strictly branching histories. We find evidence for reticulation among both the North and Central/South American groups of species, and our data support a classic hypothesis for reticulate evolution via allopolyploid speciation in the North America taxa, including an extinct diploid progenitor in this group. In the Central and South American species, we find evidence of extensive reticulation involving unknown ancestors from Asia, and we reject deep coalescent processes such as incomplete lineage sorting in favor of more recent intercontinental hybridization and chloroplast capture as an explanation for the origin of the Latin American reticulate taxa.     

Reconstructing Dryopteris "semicristata" (Dryopteridaceae): Molecular profiles of tetraploids verify their undiscovered diploid ancestor

? Premise of the study: Discovering missing ancestors is essential to understanding the evolutionary history of biodiversity on Earth. Evidence from extinct species can provide links for reconstructing intricate patterns of reticulate relationships among extant descendents. When fossils are unavailable and other evidence yields competing hypotheses to explain species ancestry, data from proteins and DNA can help resolve conflicts and generate novel perspectives. The identity of a parent shared by two tetraploid species in the cosmopolitan fern genus Dryopteris has remained elusive for more than 50 years. Based on available data, four hypotheses were developed previously, each providing a different resolution to this uncertainty. ? Methods: New molecular evidence from studies of isozymes and restriction site analysis of chloroplast DNA tested the competing hypotheses about the diploid ancestors of these two extant Dryopteris polyploids. ? Key results: The results falsify two of the hypotheses, resolve the uncertainty in the third, and support the fourth. ? Conclusions: Our data validate the prior existence of Dryopteris "semicristata," which was proposed 38 years ago as a diploid progenitor of the allotetraploids D. cristata and D. carthusiana but has never been collected. After developing a phylogeny using the new molecular data, we describe a plausible morphology for D. "semicristata" by extrapolating likely character states from related extant species.     

Spore morphology of some Brachytheciaceae Schimp. species (Bryophyta) from Turkey

The spores of Homalothecium sericeum (Hedw.) Schimp., Brachythecium populeum (Hedw.) Schimp., B. velutinum (Hedw.) Schimp. var. salicinum (Schimp.) Mönk., B. velutinum (Hedw.) Schimp. var. validum C. Jens. and Eurhynchium hians (Hedw.) Sande Lac. were studied by light and scanning electron microscopy for the first time. The apertural region consists of a leptoma in all spores. Two spore types, characterized by their surface ornamentation, are established, reflecting the species taxonomic relationships. While the surface ornamentation is of the insuloid type in Eurhynchium hians, it consists of a granuloid type in Homalothecium sericeum , Brachythecium populeum , B. velutinum . var. salicinum, B. velutinum var. validum. The spore wall of the family Brachyteciaceae includes sclerine (the dinstinction between exine and perine may be difficult to define) and intine. The taxonomy of the genera Brachytecium and are discussed on the basis of their spore morphology.     

Reticulate evolution in the apogamous Dryopteris varia complex (Dryopteridaceae,subg. Erythrovariae,sect. Variae) and its related sexual species in Japan

Apogamous fern species are often difficult to distinguish from related species because of their continuous morphological variations. To clarify the genetic relationships among the members of the Dryopteris varia complex, we analyzed the nucleotide sequences of the plastid gene rbcL and the nuclear gene PgiC. We also analyzed the diploid sexual species D. caudipinna and D. chinensis, which have not been included in the complex, but were recently shown to be closely related to the complex in a molecular phylogenetic study. The PgiC sequences of the diploid sexual species, D. varia, D. saxifraga, D. sp. ‘protobissetiana’ (undescribed diploid sexual species), D. caudipinna, and D. chinensis, were well differentiated and hence designated A, B, C, D, and E, respectively. Thus, the PgiC constitution of apogamous species in the complex was as follows: D. bissetiana, B + C; D. kobayashii, B + C + E); D. pacifica, A + C, A + B + C, or A + C + D; D. sacrosancta, A + C + E; and D. saxifragivaria, B + C. These results suggest that these apogamous species are formed by hybridizations of species including not only the three diploid sexual species of the D. varia complex (A, B, and C) but also the two diploid sexual species D. caudipinna (D) and D. chinensis (E), which do not belong to the complex.     

Pollen morphology of the genus Paliurus (Rhamnaceae)

The pollen morphology of the genus Paliurus and the 5 species it comprises is described. Consistent with the general morphological pattern of the family Rhamnaceae, the pollen grains of Paliurus are uniformly isopolar, radially symmetrical, 3-(zono)-colporate and angulaperturate. Despite this general infrageneric uniformity, certain differences and trends can be recognized with respect to pollen size, shape, and tectum architecture.     

Morphogenesis of the sexual phase of six Mexican species of the Dryopteris (Dryopteridaceae), Part II]

The development of the sexual phase of six Mexican species of Dryopteris is described and compared. Spores of all studied species are monolete, ellipsoid and have a rugose surface; the perine is folded, brown to dark brown, with a tubercled outline. Germination pattern is of the Vittaria-type and the development pattern of the prothallia is of the Aspidium-type. Gametangia are of the common type for the leptosporangiate advanced ferns. First leaves of the sporophytes appear 258-265 after sowing and apparently in Dryopteris pseudo-filix-mas the sporophyte have an apogamic origin (80 days). To make a comparative analysis of gametophytic characteristics in the twelve Mexican species and conclude of germination is of the Vittaria-type and development pattern prothallial is of the Aspidium-type, and unicelular trichomes on margin and superficial gametophytic to yield irregular aspect are characteristics to yield unit and characteristic to genera to conform Dryopteridaceae family (sensu Moran 1995) with the exception of Didymochlaena genus.