Sporophytic Characters Research Articles

Evidence for entomophily in “Knothole Moss” (Anacamptodon splachnoides)

Anacamptodon splachnoides is an uncommon moss almost entirely restricted to water-filled treeholes in deciduous trees in eastern North America and Europe. There has been uncertainty regarding taxonomic placement of the genus because of conflicts between gametophytic characters, in which it resembles Amblystegiaceae, and sporophytic characters, which seem to ally it with Fabroniaceae, Campyliaceae, or other families. Recent evidence from DNA sequencing clearly places Anacamptodon in Amblystegiaceae despite features of the sporophyte such as an erect capsule, rostrate lid, reflexed peristome teeth, and low endostome membrane. All these unusual features, including sticky spores (which seem to have been overlooked), are characteristic of species of Splachnaceae, until now the only group of mosses whose spores are known to be dispersed by flies. Field observations of A. splachnoides over a period of 16 months revealed that the moss mat and sporophytes are regularly visited by many species of flies that are also treehole specialists. Of 12 species of flies captured, nine carried spores of the moss. Many of these are strong fliers with hairy legs and bodies that inadvertently pick up the sticky spores, dispersing them in a directed fashion to other treeholes, where the females lay eggs that develop into aquatic larvae that later emerge as adults. Though differing in some respects from the adaptations seen in Splachnaceae, the parallel evolution of sporophytic characters related to entomophily is remarkable. In addition, we consider other aspects of the ecology of this moss that may help explain its rarity, such as treehole location, pH of rainfall versus stemflow and treehole water, and a possible beneficial relationship with certain wood-rotting fungi.

Encalypta driva (sp. nov.) and its relationship to E. vulgaris in Scandinavia

Encalypta vulgaris is well known for its wide morphological variation, here we study morphological and genetic variation of E. vulgaris in Scandinavia and compare this with the closely related E. rhaptocarpa and the morphologically similar E. mutica. Our morphological and genetic results indicate that Scandinavian plants of E. vulgaris s.l. includes two taxa, E. vulgaris s.s. and a new species here described as E. driva. The two species differ genetically by plastid markers while the included nuclear markers do not differentiate between the two. Morphologically they can be separated both by gametophytic and sporophytic characters. We also clarify the distinction between E. mutica and the newly described E. driva which have been confused in Scandinavia.

Study on Gametophytes of Four Species of Maiden Hair Ferns (Adiantum L.) and Their Systematic Implications

Matured spores of four (4) species of maiden hair ferns namely, Adiantum capillus-veneris L., A. caudatum L., A. edgeworthii Hook. and A. incisum Forssk. were collected and grown in petri-dishes containing natural media formed by mixing dead wood of tree fern (Cyathea Sm.), sand and charcoal powder (2:1:1). The cultures in triplicates for each species were monitored every day from the stage of sowing to the chimera formation. After disintegration of gametophytes, juvenile sporophytes were shifted to pots and morphological characters were studied till the dehiscence of sporangia. Spore germination initiated between 4-6 days for A. capillus-veneris, 7-9 days in A. caudatum and A. incisum and 10-12 days in A. edgeworthii. Prothallia in all the Adiantum are autotrophic, Vittaria type, cordate-shaped, homosporous with antheridia developing earlier than archegonia and remain confined on the adaxial surface. The gametophytes showed considerable variation in their shapes, orientation of lobes, presence or absence of hairs and placement of sex organs on the prothallia. The gametophytic and sporophytic characters are correlated and used in structuring key to species. The study concludes that gametophytic characters also constitute a dependable criterion for assessing their systematics.

Molecular tools to identify tropical mosses: a case study of the Brazilian species of Schlotheimia Brid. (Bryophyta, Orthotrichaceae)

Species of the moss genus Schlotheimia are often difficult to identify morphologically, because it is necessary to study sporophytic characters and fertile plants are quite rare. In this paper, we aim to infer the potential of trnG-R, trnL-F, and ITS markers to resolve phylogenetic relationships and species delimitations within Schlotheimia in Brazil, using different tree-based analysis methods and Automatic Barcode Gap Discovery (ABGD). For the first time in bryophytes, the trnG-R spacer was sequenced together with the trnG intron (trnG-R region). Furthermore, we aim to evaluate the morphological species concept based on the molecular data, to aid the on-going taxonomic revision of Schlotheimia in Brazil. The combined analysis of all three markers resolved 11 clades corresponding to Schlotheimia species, which was corroborated by ABGD and morphological characters. The best candidate marker for DNA barcoding was trnG-R due to its easy amplification and ability to discriminate all but one species. While trnG-R is sufficient for routine identification, the combination trnG-R + ITS should be used if all Brazilian Schlotheimia species are to be identified with high statistical support. The nuclear marker ITS was easy to amplify and more variable than the plastid markers, but a higher percentage of low quality sequences due to polynucleotide stretches or fungal contamination is a potential drawback. TrnL-F had a low discrimination potential. The analysis of the molecular markers investigated provides a baseline for the taxonomic revision of Schlotheimia and is an important tool for the identification of sterile specimens.

Taiwanobryum undulatifolium (Neckeraceae), a new combination based on sporophytic features

Taiwanobryum undulatifolium (Tixier) W.Z.Ma, Enroth & Shevock is proposed as a new combination, based on the morphological characters of the newly discovered capsules. Lack of cross-striolate ornamentation at the exostome base, long-exserted capsules and mammillose setae are the most important sporophytic characters that support the generic transfer from Neckera. A detailed description and photomicrographs are provided for the sporophyte and perichaetial leaves.

Material fructificado de Chenia lorentzii (Bryophyta, Pottiaceae) hallado en Argentina y una evaluación del valor taxonómico del esporofito en el género Chenia

Material fructificado de Chenia lorentzii (Bryophyta, Pottiaceae) hallado en Argentina y una evaluación del valor taxonómico del esporofito en el género Chenia. En la presente contribución, el esporofito de Chenia lorentzii es descrito en detalle y se proveen fotomicrografías. Además, el rol de los caracteres esporofíticos es discutido en un contexto filogenético. Para ello, primero, se lleva a cabo un análisis filogenético de un dataset reducido de géneros de Pottiaceae empleando marcadores moleculares (nad5, rps4, 26S) y datos combinados (morfología y marcadores moleculares). Luego, los caracteres esporofíticos y gametofíticos son mapeados en el árbol de moléculas y su ajuste a la filogenia es medida en base a su número de pasos extras y el número de nodos diagnosticados. Los resultados sugieren que los caracteres esporofíticos deben ser evaluados con precaución al momento de proponer cambios nomenclaturales a nivel genérico.

Multigene evidence reveals the systematic position of Pleurocladopsis simulans (C. Massal.) R. M. Schust. within Schistochila Dumort., Schistochilaceae

Abstract The monotypic Pleurocladopsis, endemic to Chile, was established by Schuster in 1964 based on an earlier poorly known species Cephalozia (?) simulans C. Massal. The phylogenetic position of Pleurocladopsis simulans had been considered uncertain until it was placed in the family Schistochilaceae on account of the gynoecial and sporophytic characters. It has been assumed that Pleurocladopsis represents the starting point of evolution in Schistochilaceae. In the present study, the phylogenetic position and taxonomic status of Pleurocladopsis simulans are inferred from phylogenetic analysis of three chloroplast DNA sequence data. The result suggests that the genus was established solely based on the autapomorphic characters, thus obscuring its actual phylogenetic relationship with Schistochila and that these characters are later derived rather than ancestral. The result also confirms that the gynoecial and sporophytic characters are important in taxonomy, but they may be not sufficient at the infrafamilial level and at other lower taxonomic levels. In accordance with the results of the present study, Pleurocladopsis is synonymised with Schistochila, and the new combination Schistochila simulans (C. Massal.) Xiao L. He & Yu Sun is made.

On the circumscription of the mossRhynchostegium hookeri(Brachytheciaceae)

Rhynchostegium hookeri and R. calderii are found conspecific and hence the latter is reduced to a synonym under the former. The orthographic error made by Vohra (1980) while latinising the species name “calderii” is rectified here to “calderi”. The description of R. hookeri is amended to include all the variations now observed, and sporophytic characters are added here for the first time. An illustration including all variations and sporophytic characters is provided.

<p class="HeadingRunIn"><strong>Common but new: <em>Bartramia rosamrosiae</em>, a “new” widespread species of apple mosses (Bartramiales, Bryophytina) from the Mediterranean and western North America</strong></p>

Recent phylogenetic analyses challenged the traditional generic concept of the Bartramiaceae (apple mosses), especially with regard to the largest genus Bartramia . Although molecular analyses revealed the three Bartramia sections ( Bartramia , Strictidium and Vaginella ) to be monophyletic, they appeared in different parts of the inferred phylogenies and thus rendered the genus itself polyphyletic. In addition, Anacolia laevisphaera , a tropical montane species, appeared in the section Strictidium , weakening its character as a typical Mediterranean element. Although morphologically similar to members of the section, such as B. stricta , A. laevisphaera has been generally treated within the genus Anacolia based on sporophytic characters. In fact, Anacolia laevisphaera is often confused with B. stricta in sterile condition. A thorough analysis revealed another surprise, in placing A. laevisphaera sister to the Southern South American samples of Bartramia stricta, while the Mediterranean populations of “ B. stricta ” cluster with the Australian B. breutelii . Subsequent morphometric studies revealed three morpho-species in accordance with the polyphyletically resolved B. stricta . Here we describe the new species Bartramia rosamrosiae to accommodate the Mediterranean and western North American populations of what was traditionally treated as B. stricta .

Some Medicinal Bryophytes: their Ethnobotanical Uses and Morphology

Medicinal bryophytes and their morphological features and ethnic uses were examined and documented. Field collections of bryophytes were made through transect walks along the two mountain sites of Bukidnon, namely: Mt. Kalatungan and Mt. Kitanglad. The number of taxa currently determined with medicinal properties was based from the collected species. The species with medicinal uses was generated from a pool of resources, literature review and scientific journals. Description on their morphological characters was made using microscopic examinations. Taxonomic characters used include leaf orientation, stem structure, sporophyte characters and their habit to classify into families, genera and species. Species composition between study sites was also compared. Visual habitat as to habitat preference and identification for each species was employed. Results of the study highlight the difference in the morphological features among species of medicinal bryophytes, their ethnic uses, and adaptations within the forested area. Mt. Kitanglad exhibited seventeen (17) species of medicinal bryophytes. Of these, eleven (11) are mosses belonging to seven (7) families in eleven (11) genera and six (6) liverworts belonging to five (5) families in six (6) genera. Mt. Kalatungan exhibited fourteen belonging to six (6) families in eight (8) genera and six (6) liverworts belonging to five (5) families in six (6) genera. The moss species with medicinal properties belongs to the genera Sphagnum, Polytrichum, Rhodobryum, Fissidens, Bryum, Mnium, Dawsonia, Philonotis, Pogonatum, Barbula and Plagiomnium . The liverworts species include Marchantia, Pallavicinia, Herbertus, Riccardia, Dumortiera and Plagiochila . The species under study exhibited antimicrobial activity, anti-tumor, anti-cancer, antileukemic activity and healing effects based from secondary data. Â Keywords - ethnobotany, morphology, bryophytes, taxonomic treatment, floristic, ethnic uses

Phylogenetic inference rejects sporophyte based classification of the Funariaceae (Bryophyta): Rapid radiation suggests rampant homoplasy in sporophyte evolution

The moss family Funariaceae, which includes the model systems Funaria hygrometrica and Physcomitrella patens, comprises 15 genera, of which three accommodate approximately 95% of the 250–400 species. Generic concepts are drawn primarily from patterns in the diversity of morphological complexity of the sporophyte. Phylogenetic inferences from ten loci sampled across the three genomic compartments yield a hypothesis that is incompatible with the current circumscription of two of the speciose genera of the Funariaceae. The single clade, comprising exemplars of Funaria with a compound annulus, is congruent with the systematic concept proposed by Fife (1985). By contrast, Entosthodon and Physcomitrium are resolved as polyphyletic entities, and even the three species of Physcomitrella are confirmed to have diverged from distinct ancestors. Although the backbone relationships within the core clade of the Funariaceae remain unresolved, the polyphyly of these genera withstands alternative hypothesis testing. Consequently, the sporophytic characters that define these lineages are clearly homoplasious suggesting that selective pressures (or their relaxation) are in fact driving the diversification rather than the conservation of sporophytic architecture in the Funariaceae.

Orthotrichum pilosissimum (Orthotrichaceae), a new moss from arid areas of Nevada with unique axillary hairs

Abstract Orthotrichum pilosissimum is described from herbarium material and recently collected specimens in the state of Nevada. Its specific epithet refers to the diagnostic hyper-developed axillary hairs, which are frequently longer than 1 mm, multiseriate and branched, covering most of the stems surface and sometimes protruding from the foliose shoots. These hairs are not only unique in the genus but also among mosses, providing a reliable diagnostic character for the new taxon. Additional gametophytic and sporophytic characters that contribute to its taxonomic delimitation are also given.

Studies on the population status of Dumortiera Hirsuta (S.W.) Nees growing in Bhaderwah (North West Himalaya)

Paper includes information on intraspecific variability (ecologial and morpho-anatomical) among 25 accessions of Dumortiera hirsuta collected from diverse habitats in Bhaderwah (J.&K.). Taxon exhibits tremendous diversity in ecological (habitat, pH and altitude) and morpho-anatomical (both gametophytic and sporophytic) characters.

Ephemerella readeri Müll. Hal. (Physcomitrella readeri (Müll. Hal.) I.G. Stone & G.A.M. Scott, Funariidae, Bryophyta): a genus and species new to Europe

A morphological and molecular analysis of a Physcomitrella, collected from a reservoir margin in the north of England, revealed this to be P. readeri, a species new to Europe. The present study clarifies previous confusion over the taxonomy of P. readeri showing it to be clearly distinct in both sporophytic and gametophytic characters from P. patens and uniform across its world range from England to USA, Japan and Australasia. While phylograms of the ITS1 region from both the Physcomitrella species, Physcomitrium pyriforme (Hedw.) Bruch & Schimp., Enthosthodon attenuatus (Dicks.) Bryhn and Funaria hygrometrica Hedw., place the first two in separate clades, in ITS2 phylograms they occur as sister taxa. This, together with previous genealogical studies on the speciation history of the Physcomitrella–Physcomitrium species complex, and morphology, suggests that generic rank is appropriate for P. readeri. We therefore reinstate the original name Ephemerella readeri Müll. Hal. Recent records at several reservoirs in England indicate that E. readeri may be native to UK, though remarkable congruence in ITS1 with Australian plants also suggests recent arrival as an alternative possibility.

Independent Origins of Cleistocarpy in the Splachnaceae: Analyses of cpDNA Sequences and Polyphyly of the Voitioideae (Bryophyta)

Many dung mosses (Splachnaceae) are characterized by insect-mediated spore dispersal. All of the ento- mophilous species are coprophilous, whereas anemophilous species are humicolous or epiphytic. The three species of the Voitioideae are coprophilous but are distinguished from other members of the family by sporangia that remain closed (cleistocarpous) and lack a peristome. Spores are released when the sporangial wall disintegrates. Phylogenetic analyses of nucleotide sequences of the trnL-trnF region and the rps4 locus (cpDNA) for 25 species of Splachnaceae suggest that this combination of characters arose twice within the Splachnaceae and that Voitia grandis is more closely related to species of Tayloria subgenus Tayloria rather to the other species of Voitia, which are nested within the genus Tetraplodon. Hence the new combination Tayloria grandis (Long) Goffinet & Shaw is made. Although the optimal trees were left unrooted, our results resolve the Voitioideae (i.e., the genus Voitia ) as nested within the Splachnoideae. The phylogenetic significance of sporophytic characters within the family Splachnaceae is briefly discussed. Some members of the moss family Splachnaceae are unique among seedless land plants: they display fea- tures considered linked with insect-mediated spore dispersal. Such putative adaptations include an en- larged, brightly colored hypophysis, emission of vola- tile compounds that attract insects, sticky spores, and a pseudocolumella (Koponen 1990; Demidova and Fi- lin 1994). The systematics of the family is based pri- marily sporophytic features, and in particular of com- binations of entomophily related characters and the ar- chitecture of the peristome (Koponen 1982). The

Funaria convexaSpruce (Musci: Funariaceae) New to North America

Funaria convexa Spruce is reported new for the North American moss flora from the North Coast Ranges of Lake County, California. It is closely related to Funaria muhlenbergii Turn, and F. pulchella Philib., but differs in several gametophytic and sporophytic characters. A distribution map showing the Mediterranean climates disjunction is included.

Sciuro-hypnum tromsoeense (Kaurin & Arnell) Draper & Hedenäs, a distinct species from the European mountains

Morphological and molecular evidence reveal that Sciuro-hypnum tromsoeense, nowadays considered conspecific with S. starkii, is actually a distinct species that can be distinguished from related species by both gametophytic and sporophytic characters, such as branch shape, stem leaf shape and areolation and seta roughness. An amended description of this moss is presented and its diagnostic characters are discussed. The species grows on moist soil, litter and rocks, and is here confirmed for several boreal and mountainous areas of Europe. In addition, Hypnum reflexum Starke is neotypified in this paper.

Molecular and morphological evidence on the status of Bryum mildeanum Jur. and records revision in the Iberian Peninsula

Bryum mildeanum, a rare species in the Iberian Peninsula, has frequently been confused with B. alpinum. In this study we provide new gametophytic and sporophytic (obtained by SEM) characters that can be used to separate the two and B. mildeanum is described and lectotypified. A molecular study based on two chloroplast DNA regions (trnL-F and trnG) was performed. The results suggest the evolutionary independence of B. mildeanum and B. alpinum, and the separation of both from B. gemmiparum, a species generally placed in the same subsection (Section Bryum, subsection Alpiniformia) as the other two.

Orthotrichum Norrisii (Orthotrichaceae), A New Epiphytic Californian Moss

A new Orthotrichum species, O. norrisii, is described from California. The new taxon is recognized by a unique combination of sporophytic characters: capsule narrowly cylindrical and not contracted below mouth when dry, exothecial bands narrow and formed by 2(–3) cell rows, peristome with eight pairs of exostome teeth and eight endostome segments, stomata cryptopore and located in the lower part of the urn, lid plane and rostrate and vaginula hairy. Widely distributed throughout California, the new taxon has been overlooked and confused with O. tenellum.

Reëvaluation of Orthotrichum Rhytiore (Orthotrichaceae)

Examination of the holotype of Orthotrichum rhytiore B. H. Allen reveals gametophytic characters which support the placement of the species in Ulota. The species is peculiar within the genus due to its unusual sporophyte morphology, and the occurrence of these sporophytic characters in Ulota is discussed. Ulota rhytiore (B. H. Allen) F. Lara, Garilleti, Albertos & Mazimpaka comb. nov. is the first known species of the genus in tropical America.