The time of fruiting of Ulota Bruchii Hornsch. and U. crispa Brid.

Abstract

For obscure reasons, sporophytes of the dioicous Hamatocaulis vernicosus are currently unknown in France. With the aim of understanding this failure of sporophyte production we investigated (1) sexual phenology, (2) sex ratio pattern at different spatial scales, (3) limitation of sporophyte formation by the availability of only one sex, (4) limitation of sporophyte formation caused by the distance between male and female, (5) location of sporophytes and (6) the consevation implications of this failure of sporophyte production. Principal results show that phenology cannot explain the failure of sporophyte production. At the regional scale, sporophyte formation is prevented by the fact that several mountain ranges have single-sex populations. The production of spore capsules relies on a strict combination of factors at the local scale: occurrence of sexually expressed mixed-sex colonies, a short distance between male and female individuals and favourable environmental factors (light grazing, high water table). In the Massif Central of France spore production is unlikely because of current unfavourable conditions. Fertility patterns appeared useful in a conservation context.     

Arbuscular mycorrhiza and water and nutrient supply differently impact seedling performance of dry woodland species with different acquisition strategies

Background: We recently demonstrated that dwarf male plants are much more common among mosses than hitherto thought. Dwarf plants, producing functional male sexual organs occur in 10–20% of the moss species worldwide.

Aims: We investigated how an inadvertent omission of dwarf males affects estimates of male sex expression rates and sex ratios.

Methods: We recorded frequencies of specimens with female-expressing plants, male-expressing normal-sized and/or dwarf plants, and sporophytes in herbarium material of five model species from Macaronesia. We analysed how frequency estimates of male occurrence and sex ratios were affected by excluding dwarf males.

Results: Occurrence of dwarf males was positively associated with sporophyte formation. When dwarf males were excluded, male mate availability was missed in 51–61% of specimens (in 74–76% of sporophyte-containing specimens). Specimen-level sex ratio was balanced in material with sporophytes in the three species exhibiting male dwarfism, but strongly female biased if dwarf males were omitted.

Conclusions: We provide the first quantitative evidence that sex-expressing dwarf males are essential for sexual reproduction and sporophyte production in mosses that form such males. Considering its abundance, male dwarfism therefore deserves more attention among plant biologists. Innovative approaches are required to further study the minute male plants.     

Sex distribution and sporophyte frequency in a population of the clonal moss Hylocomium splendens

Abstract

The sex ratio and factors that affect sporophyte production were studied by following one population of the clonal moss Hylocomium splendens growing on top of a boulder in a S.E. Norwegian boreal spruce forest valley for 5 years. Sex of ca 75% of all segments recorded during the study period could be determined from records of occurrence of sporophytes, gametangia and branching patterns. The population had a 4:1 female-biased sex ratio. Of 1005 female segments recorded during the 5-year study period, 29.8% carried sporophytes. The frequency of sporophyte production in female segments varied more than threefold among years, and was apparently related to climatic favourability for bryophyte growth and development. At the scale of H. splendens segments, sporophyte production was favoured by large size of segments and high relative growth rates. A distinct low-size threshold for sporophyte production occurred in all years. This threshold was higher in years of low sporophyte frequency, with an absolute low limit of ca 2 mg segment dry weight. The fraction of female segments with sporophytes, and the number of sporophytes per sporophyte-carrying segment, increased with segment size from the low threshold to an upper size threshold, beyond which further increase in segment size did not imply increased sporophyte frequency. This may be caused by the spatial distribution of sex organs. Sporophyte production increased strongly upwards in the bryophyte carpet, even after vertical variation in size had been taken into account. This indicates that radiation plays a key role in initiation of sexual branches, and that the processes previously demonstrated to regulate vegetative branching apply to branching in general.     

The male gametophores of Leucobryum glaucum (Hedw.) Ångstr. and L. juniperoideum (Brid.) C. Muell. in two Welsh woodlands

Abstract

The distribution and variation in size of male gametophores associated with fertile, female cushions of Leucobryum glaucum and L. juniperoideum in two Welsh localities are described. Both dwarf and larger males were present in fruiting cushions of both species; no independent male plants were found. It is suggested that the scarcity of functional males might be a factor limiting sporophyte production in these species.     

Reproductive traits in the tropical moss Octoblepharum albidum Hedw. differ between rainforest and coastal sites

Abstract

Octoblepharum albidum Hedw. is an autoicous moss commonly occurring in tropical savannas, dry forests, rainforests and coastal habitats. It frequently reproduces by spores and asexual structures (gemmae and protonemata or buds at leaf tips), making it a good model for understanding how reproductive traits change with respect to habitat type. Our aims were to characterize the different life-history traits in O. albidum relative to sexual and asexual cycles and to detect variations in reproductive performance among the different habitats, trade-offs among these traits, and relationships among reproductive traits and plant length. We studied colonies from two Atlantic rainforests and two coastal sites in north-eastern Brazil. Shoots in the coastal sites, compared to those of the forest sites, had higher numbers of sporophytes, male and female branches per shoot, male gametangia per sexual branch, and longer setae. Numbers of female gametangia per sexual branch did not differ between forest and coastal sites. A male-biased sex ratio of branches and gametangia occurred in all sites. Compared to gemmae, sporophytes and protonemata or buds were more likely to be found on longer shoots than on shorter ones, but this relationship was only applicable to forest sites. The abundant production of gemmae and protonemata or buds at leaf tips, and sporophytes (spores) in O. albidum are important components in explaining colonization success and maintenance in this tropical moss. Longer sporophytic setae in addition to a higher reproductive performance (especially for number of male gametangia and sporophytes per shoot) may favour spore dispersal and colonization in plants of coastal sites.     

The life history ofAcrochaete wittrockii (Ulvellaceae,Chlorophyta)

Acrochaete wittrockii (Wille) Nielsen is a heteromorphic diplohaplont. The haplophase consists of isomorphic, dioecious filamentous epiphytes on brown algae. Several generations follow each other by triflagellate zoospores from spring to early summer. By late summer and throughout autumn, quadriflagellate zoopores are produced by the epiphytic thalli; they give rise to male and female gametophytes of a globular, pseudoparenchymatic appearance in culture. The gametophytes produce anisogamic biflagellate gametes which, after gametic union, develop into diploid unicellular sporophytes. After 6–7 days, the sporophyte produces triflagellate zoospores, repeating the life history when germinating on brown algal hosts. Alternatively, triflagellate zoospores which settle on the bottom of petri dishes, develop into unicellular, autonomous sporangial plants. Their triflagellate spores repeat the epiphytic stage on brown algal hosts, or the sporangial plant cycle on non-living substrate, respectively.     

Comparison of spatial scale variability of shoot density and epiphytic leaf assemblages of Halophila stipulacea and Cymodocea nodosa on the Eastern Coast of Tunisia

Abstract

A study was undertaken to compare the patterns of spatial variability, epiphytic percentage cover, and distribution of epiphytic fauna and flora between the two adjacent seagrasses Cymodocea nodosa and the invasive species Halophila stipulucea. Samples were taken at six stations separated by 600?m and exposed to different current conditions. The stations G2 and G5 were affected by a high current tide, while G4 was directly exposed to the northern marine currents. The station G3 was situated in the middle of a Posidonia oceanica bed and was less exposed to hydrodynamism, whereas the other stations were relatively protected. Results indicate that for both H. stipulacea and C. nodosa, shoot density and epiphytic cover biomass decreased when exposed to high levels of hydrodynamic activity. In terms of epiphytic leaf assemblages, our results showed significant differences for the two host plants in their mean cover and for the six stations at the largest and smallest spatial scale. Our observations highlight the dominance of Rhodophyta and the low number of epiphytic species and the epiphytic cover on H. stipulacea compared to C. nodosa. In addition, results indicate the absence of two taxa Hydrozoans and Annelida in the epiphytic assemblage of H. stipulacea leaves.     

Factors influencing reproductive success in the clonal moss, Hylocomium splendens

Female reproductive success in the unisexual perennial clonal moss Hylocomium splendens was examined by recording, if the segment was reproductive [produced sporophyte(s)] or not, together with several distance-to-male and male density variables, and segment size. This was done for every female segment in a population over a 5 year study period. A high fraction of the population could be sexed because we monitored the population in situ for 5 years, and thereafter harvested the population for electrophoretic analysis from which the clonal identity and expressed sex could be deduced. Fertilization distances in H. splendens were short, indicated by the fact that as many as 85% of the female segments with sporophytes were situated within a distance of 5.0 cm from the nearest male. The longest distance measured between a sporophytic female and the closest male was 11.6 cm. However, analysed within a generalized linear modelling (GLM) framework, the year was the best single predictor for the presence of H. splendens sporophyte although female-segment size and distance to the closest situated male were also strongly significant. The two latter factors explained larger fractions of variation in sporophyte presence in a GLM model with three predictors than in single-predictor models. This is because (i) the large variation in sporophyte production among years partly obscures the strong general increase in sporophyte production with increasing female-segment size and vitality, and (ii) the between-year variation and the size obscure the effect of the distance to the most proximate male. To our knowledge, this study is the first to incorporate into one model the relative importance of several factors for bryophyte reproductive success. Our results demonstrate the value of multiple-predictor approaches in studies of reproductive success.     

The adaptive background of nannandry: dwarf male distribution and fertilization in the moss Homalothecium lutescens

Dwarf males (nannandry) occur in many unrelated, mostly aquatic, groups of organisms. Among land plants they are unique to bryophytes. In this study our aim was to explain variation in frequency of dwarf males and fertilization within populations in the moss Homalothecium lutescens. We compared parameters related to dwarf male presence and sporophyte production in 90 colonies from three localities. Dwarf male density was positively associated with colony moisture at two of the localities, suggesting increased spore germination and dwarf male survival with moist conditions. At one of these localities, dwarf male density was also positively associated with the presence of perichaetia (female sexual branches). Dwarf male density and fertilization frequency were positively associated in two of the localities. Furthermore, in one population, fertilization was also positively associated with canopy cover, which could be attributed to improved nutrient status as a result of throughfall, increased sperm‐dispersal efficiency because of larger water drops, or more favourable moisture conditions as a result of shading. Nannandry thus appears to strongly reduce the problem of short fertilization distances in bryophytes, but the presence of water is still critical because the dwarf males are dependent on a certain level of humidity for recruitment and/or development. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 74–84.     

La Posidonia oceanica: problemi di ecologia,di funzionamento e di biologia

Abstract

The functional roles of seagrasses in the economy of coastal waters are to be fully realised and more clearly delineated. The biology, ecology and energetics of seagrasses meadows of the Mediterranean Sea, and in particular of Posidonia oceanica inclusing the stabilization of coastal sediments, this role as a base for major detrital food chains, nutrient resources contributory to general productivity of the open sea and the support of dense epiphytic communities, are discussed.     

Observations on Archilejeunea apiculifolia St. from India

Abstract

The occurrence of microphyllous branches, variable disposition of inflorescences, structural range in female bract-lobule, perianth keels, seta anatomy and sporophyte details of Archilejeunea apiculifolia St., which were previously not known, are described.     

A method to establish an “immortalized F2” sporophyte population in the economic brown alga Undaria pinnatifida (Laminariales: Alariaceae)

Studies of quantitative trait loci based on genetic linkage maps require the establishment of a mapping population. Permanent mapping populations are more ideal than temporary ones because they can be used repeatedly. However, there has been no reported permanent sporophyte population of economically important kelp species. Based on the characteristics of the kelp life cycle, we proposed a method to establish, and then constructed experimentally, an “immortalized F₂” (IF₂) population of Undaria pinnatifida. Doubled-haploid “female” and “male” sporophytes were obtained through the parthenogenesis of a female gametophyte clone and the selfing of a “monoicous” gametophyte clone (originally male), respectively, and they were used as the parents. The F₁ hybrid line was generated by crossing the female and male gametophytes derived from the respective female and male parents. Full-sibling F₂ gametophyte clones, consisting of 260 females and 260 males, were established from an F₁ hybrid sporophyte. Thirty-five females and 35 males were randomly selected and paired to give rise to an IF₂ population containing 35 crossing lines. A parentage analysis using 10 microsatellite markers confirmed the accuracy of the IF₂ population and indicated the feasibility of this method. This proposed method may be adapted for use in other kelp species, and thus, it will be useful for genetic studies of kelp.     

Calymperastrum,a new genus of Pottiaceae

Abstract

Annual growth rate and spore and capsule production was studied in 50 colonies of the epixylic hepatic, Ptilidium pulcherrimum in northern Sweden. Radial growth rate was independent of colony size but varied from year to year (mean value 5.2 mm year^?1, range 3.5–6.3 mm year^?1). Size at first sporophyte production was estimated to be 68 cm² and the age of first reproduction was estimated to be about 9 years. It was estimated that antheridia were formed the third year. No correlation between growth rates and reproduction was observed. Capsule density as well as capsule and spore production increased significantly with increased colony size. Differences between years in spore and capsule production (up to six-fold) were observed. The mean number of spores per capsule was 27 400 but significant differences between colonies (range from 18 000 to 44 000) were observed.     

Status,conservation and ecology of Pallavicinia lyellii (Hook.) Carruth. in a key region of England

ABSTRACT

Introduction. Pallavicinia lyellii is recognised as being of principal importance for conservation in England and is threatened with extinction in Europe. The aims of this study were to investigate the current status, conservation and ecology of the species in S Hampshire, a key region for the plant.

Methods. All recent and historic locations for Pallavicinia lyellii in S Hampshire were searched and information collected on distribution, abundance and ecology.

Results. Pallavicinia lyellii was re-found at three sites. It is almost certainly extinct at two sites and may have been lost from a third. Whilst few colonies were sterile, most comprised a single sex. Colonies with both male and female plants were confined to Cadnam Bog, which supported 49 (91%) of the colonies found. Overall, female shoots occurred at 41 (76%) colonies and males at 26 (48%), producing a female sex bias (1.6:1). All colonies occur in boggy woodland, mostly on the sides of low mounds of damp, acidic, peaty soil developed, due to poaching by ponies and cattle, at the base of trees, usually I. aquifolium. The community mainly comprises a species-poor mix of acidophilous mosses and liverworts, most frequently Mnium hornum, Pseudotaxiphyllum elegans and Leucobryum juniperoideum.

Conclusions. Cadnam Bog appears to support the largest sexually reproducing population of Pallavicinia lyellii in England. The site is specially protected, actively managed with consideration to P. lyellii and seemingly in favourable condition for the plant. Research is needed to quantify canopy shade favoured by the plant and assess implications for woodland management. Future monitoring should include re-survey of populations every 3?yr.     

CUTICLES FROM CHONDRUS CRISPUS (RHODOPHYTA)1

A simple, nondestructive physical process was developed for routinely isolating the outermost layers from female, male, and sporophyte fronds of Chondrus crispus Stack-house. Yields of pure cuticles from apical segments ranged from 0.74 to 2.35% on a dry weight basis after 5–7 d of culture. These undegraded cuticles were examined by electron microscopy (scanning and transmission electron microscopy), spectroscopy (infrared and X-ray), and chemical means. Cuticles isolated from female or male fronds were characterized by parallel arrays of electron-dense lamellae (typically 6–14) alternating with more electron-transparent regions. The thickness and uniformity of these lamellae provide the physical basis for the iridescence characteristic of C. crispus fronds. Sporophyte fronds are not iridescent. This phenomenon may be explained by the fewer electron-dense cuticular lamellae (usually three to seven) and the fact that these lamellae anastomose freely to form a thin cuticle with a highly irregular substructure. Elements detected by X-ray analysis, in addition to carbon and oxygen, included Mg, Br, S, and Ca in both gametophyte and sporophyte cuticles. Major features of FTIR spectra of all cuticles were absorbances due to proteins. A strong band, indicative of sulfate ester, occurred near 1250 cm^?1 in all cuticle preparations. Gametophyte, but not sporophyte, cuticles absorbed at 935, 846, and 800 cm^?1 consistent with the presence of kappa and/or iota carrageenan. Amino acid analyses showed that sporophyte and gametophyte cuticles were generally similar in gross composition. All contained proline as the principal residue together with significant amounts of cysteine, methionine, and lysine. Protein contents calculated from these analyses ranged from 37.6 to 44.4% on a dry weight basis as compared to 51.5–56.7% calculated from total nitrogen values. Up to 75% of the cuticle mass was solubilized by sodium dodecyl sulfate-β-mercaptoethanol. Three similar migrating bands were seen in female and male cuticle extracts on sodium dodecyl sulfate–polyacrylamide gel electrophoresis; however, none of the three weaker bands from sporophyte cuticles comigrated with those from gametophytes. Chloroform-methanol extraction removed < 3.3% of the cuticle mass, suggesting that lipids were minor components.     

C3 photosynthesis in the gametophyte of the epiphytic CAM fern Pyrrosia longifolia (Polypodiaceae)

Sporophytes of some epiphytic species in the fern genus Pyrrosia exhibit Crassulacean acid metabolism (CAM), generally considered to be a derived physiological response to xeric habitats. Because these species alternate between independent sporophytic and gametophytic generations yet only the sporophyte has been characterized physiologically, experiments were conducted to determine the photosynthetic pathways present in mature sporophytes, immature sporophytes, and gametophytes of Pyrrosia longifolia. Diurnal CO₂ exchange and malic acid fluctuations demonstrated that although the mature sporophytes exhibited CAM, only C₃ photosynthesis occurred in the gametophytes and young sporophytes. Consideration of the above results and those from previous studies, as well as the life cycle of ferns, indicates that the induction of CAM probably occurs at a certain developmental stage of the sporophyte and/or following exposure to stress. Elucidation of the precise mechanisms underlying this C₃-CAM transition awaits further research.     

Reproductive features and gonad development cycle of the soft bottom-gravel gorgonian Leptogorgia sarmentosa (Esper, 1791) in the NW Mediterranean Sea

Abstract

Leptogorgia sarmentosa (Cnidaria: Octocorallia) is a common soft bottom-gravel gorgonian in the northwestern Mediterranean Sea. This study examined the annual reproductive cycle of a L. sarmentosa. population at a depth of 30 m, and further determined the sex ratio, the intra-colonial gonadal output variability and the age at first reproduction. L. sarmentosa was gonochoric and the population studied was characterized by a sex ratio slightly biased towards female colonies (1 male:1.6 female). Secondary branches had higher polyp fertility (number of gonads per polyp) compared to primary branches, but because of the number of polyps on the former, most of the sexual effort was actually contributed by the primary branches, as in other gorgonian species. Male colonies became fertile above a height of 21 cm, but female colonies had gonads even in the 1–10 cm height class. Spawning occurred between late August and early September, although there was no direct observation of broadcast spawning or surface brooder strategy. The maximum gonad size was 550 μm in males and 500 μm in females, with a mean diameter of 270 μm for spermaries, and 190 μm for eggs. The number of gonads per polyp prior to spawning reached 4–5 spermaries polyp!1 in male colonies, and 3–4 eggs polyp!1 in female colonies. Spawning did not appear to be synchronized with the full moon. L. sarmentosa appeared to have a different reproductive strategy to other Mediterranean gorgonians (living on hard bottom substrata), probably because of its different habitat preferences and biological (morphology, diet, prey capture rates, growth, biochemical levels) features.     

Male-male Competition and Female Mate Choice through Courtship Display in the Territorial Damselfish Stegastes nigricans

Both sexes of the herbivorous damselfish Stegastes nigricans maintain individual feeding territories. These territories are distributed contiguously, forming distinct colonies. Females visit male territories to spawn, and eggs are guarded by males until hatching. Male-male competition and female mate choice were studied in two colonies of different size compositions. Only larger individuals bred in both colonies. Some males in the large colony, that were larger than the breeding males in the small colony, did not succeed in reproducing probably because of severe attacks by the larger males while courting. However, females did not choose large size among breeding males. The most important male characteristic in female choice was the frequency of courtship displays in both colonies. Females in the large colony chose males mainly on the basis of the frequency of displays conducted in the females' territories, whereas females in the small colony chose males on the basis of the frequency of displays conducted in the males' territories. This difference may be a result of the difference in colony size. The distances between females' and males' territories were much greater in the large colony, and, because females cannot see courtship displays conducted in distant male territories, males in the large colony may have had to visit female territories frequently in order to conduct courtship near the females.     

Family affiliation,sex ratio and sporophyte frequency in unisexual mosses

Patterns of sex expression and sex ratios are key features of the life histories of organisms. Bryophytes are the only haploid‐dominant land plants. In contrast with seed plants, more than half of bryophyte species are dioecious, with rare sexual expression and sporophyte formation and a commonly female‐biased sex ratio. We asked whether variation in sex expression, sex ratio and sporophyte frequency in ten dioecious pleurocarpous wetland mosses of two different families was best explained by assuming that character states evolved: (1) in ancestors within the respective families or (2) at the species level as a response to recent habitat conditions. Lasso regression shrinkage identified relationships between family membership and sex ratio and sporophyte frequency, whereas environmental conditions were not correlated with any investigated reproductive trait. Sex ratio and sporophyte frequency were correlated with each other. Our results suggest that ancestry is more important than the current environment in explaining reproductive patterns at and above the species level in the studied wetland mosses, and that mechanisms controlling sex ratio and sporophyte frequency are phylogenetically conserved. Obviously, ancestry should be considered in the study of reproductive character state variation in plants. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 163–172.