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Entomophily in the Splachnaceae
Abstract
The Splachnaceae is the only moss family in which entomophily has been observed. There are three kinds of adaptation for entomophily: (1) adaptations to substrates of animal origin, (2) morphological adaptations and (3) chemical adaptations.
Species of Sptachnum and the entomophilous species of Tayloria are restricted to the dung of herbivorous mammals. Species of Tetraplodon grow on skeletal remains, on antlers, on stomach pellets of predatory birds, or on dung, which contains both bone and hair. The entomophilous species Aplodon wormskioldii grows on corpses, on caribou (reindeer) dung, bones and antlers, on owl pellets, or on enriched gravel.
Cultivation experiments indicate that protonema and shoots of Splachnum sphaericum have a greater tolerance for concentrated dung liquid than the control species Physcomitrium pyrij'orme, Funaria hygrometrica and Pohlia nutans. Tetraplodon mnioides, A. wormskioldii and Splachnum vasculosum have higher nitrogen contents in their tissues than do other arctic bryophytcs and the levels of nitrogen, phosphorus and calcium in the plant tissues of T. mnioides and A. wormskioldii reflect those of their substrata. The adaptation to grow on osmotically concentrated substrates confers a selective advantage upon these species.
Morphological adaptations of the sporophyte are the enlarged, coloured neck (hypophysis), the coloured upper region of the seta, and the hygroscopic movements of the urn wall and peristome which help the spores to leave the urn. The small, thin-walled spores are dispersed in clumps on the hairs of visiting flies.
Chemical adaptations are the odours produced and released by the sporophyles. Several volatile compounds have been found in the urn and hypophysis of entomophilous species, volatile octane derivatives and organic acids such as acetic, propionic and butyric acids particularly in the hypophysis. Volatile compounds may be released through the many stomata of the hypophysis. The seta, the gametophyte and the substratum do not contain these compounds. Flies which are attracted to and visit species of Splachnum in Europe and North America include Scatophaga furcala, (Scatophagidae), Pyrellia cyanicolor, Myospila metida (Muscidae) and Delia platura (Anthomyidae). The most common visitors to Tetraplodon mnioides are Delia platura and species of the Muscidae. Scatophagids have been found to benefit from a possible increase in copulatory success after visiting these plants.
Entomophilly in the Splachnaceae can be regarded as chemical mimicry. Other organisms that mimic faecal/carrion odours to attract flies to disperse the spores include the macrofungus Phallus impudkus. Some species in the angiospcrm families Ramcsiaccae and Araceae, which arc pollinated by flies, produce odours resembling those found in the Splachnaceae.
... Sin embargo, hay casos de anemocoria ante cambios demográficos del vector biótico, falta de disponibilidad de sustratos, o severidad y restricciones en el hábitat (Cox, 1991;Culley et al., 2002). Koponen (1990) describió para la familia Splachnaceae las distintas adaptaciones morfológicas y químicas a la entomocoria. Entre estas adaptaciones, son importantes el tipo de sustrato, los esporofitos con cápsulas muy coloridas con modificaciones a nivel de la hipófisis a menudo agrandada, que emanan compuestos volátiles de olores que imitan la materia animal en descomposición. ...
... Entre estas adaptaciones, son importantes el tipo de sustrato, los esporofitos con cápsulas muy coloridas con modificaciones a nivel de la hipófisis a menudo agrandada, que emanan compuestos volátiles de olores que imitan la materia animal en descomposición. La familia Splachnaceae era la única en la que se había observado este modo de dispersión de las esporas a través de los insectos (Koponen, 1990). Pero Ignatov & Ignatova (2001), observaron en bosques de coníferas de la región Holártica que el musgo Schistostega pennata (Hedw.) ...
... Para el hemisferio sur, son escasos los trabajos publicados sobre las especies de esta familia (Jofre et al., 2010(Jofre et al., , 2011Mighell, 2011;Piñeiro & Solan, 2018). No obstante, Koponen (1977Koponen ( , 1990, Goffinet et al. (2004) y Glime (2017), ya habían advertido la falta de pruebas empíricas y experimentos de captura que permitan comprobar la dispersión de esporas a través de insectos en las especies del hemisferio sur. ...
The Splachnaceae family is a bryological component of the temperate forests of Nothofagus in the Fuegian region of Argentina. It is represented by the Tayloria genera with three species, T. dubyi (endemic), T. magellanica, T. mirabilis, and Tetraplodon, with a single specie Tetraplodon fuegianus. They grow on organic substrates of animal origin and are the only family among mosses in the area in which entomochory is observed (i.e. dispersion of spores through insects). From herbarium material, the taxonomic features of gametophytes and sporophytes which allow species to be identified are described. Spores were studied with OM and SEM. Dispersal vectors for Tayloria mirabilis and morfo-ecological adaptations associated with entomochory were observed and analyzed. Mosses are differentiated from their leaves and the morphology and color of the sporophyte capsules. The spores, similar in the studied species, are dispersed in sticky masses, they are spheroidal, monoletes, 8-13 μm of diameter with a pitted-reticulate ornamentation. The dispersing agents mostly correspond to the order Diptera. The Splachnaceae family has developed adaptive strategies in relation to substrate (coprophilous gametophytes), in striking and showy sporophytes morphologies and in a particular dispersal mode of the spores by insects. All these morpho-ecological adaptations contribute to an effective action of the dispersing agents in the muscinal biocenosis.
... An exception to the pattern that species having adhesive propagules do not actively recruit their dispersers is found in the moss family Splachnaceae, in which the spores of all Splachnum, Tetraplodon and several Tayloria species are dispersed to decaying substrates (feces, carrion) by diverse families of flies (Koponen, 1990). The sporophytes of these mosses are brightly coloured and distinctively scented, but provide no nutritional reward. ...
... The bryophyte family Splachnaceae is a globally distributed (Koponen, 1990) monophyletic lineage (Goffinet et al., 2004) of 73 species with genera distributed among three subfamilies: Aplodon, Tetraplodon and Splachnum (Splachnoideae), Moseniella and Tayloria (Taylorioideae) and Voitia (Voitioideae) (Crosby et al., 1999;Bai and Tan, 2000;Fife and Goffinet, 2003;Goffinet et al., 2004). Most species are confined to north or south temperate habitats, whereas others are endemics to isolated mountains or montane forests at (sub)tropical latitudes. ...
... Like stinkhorn fungi (Sleeman et al., 1997;Tuno, 1998), the Splachnaceae attract flies not to promote sexual reproduction, but to disperse asexual spores to appropriate substrates (Koponen, 1990). Although entomophily previously has been demonstrated in only a small number of taxa of the Northern hemisphere (Koponen and Koponen, 1977;Pyysalo et al., 1978;Cameron and Troilo, 1982;Pyysalo et al., 1983;Marino, 1988a;Koponen, 1990;Marino, 1997;Jofre, 2007), in all putatively entomophilous Splachnaceae, sporophytes emit odours from the inflated and/or elongated and often coloured (e.g., yellow, magenta, red) apophysis, the sterile region of the capsule below the sporangium ( Fig. 1b; Pyysalo et al., 1978;. ...
The use of sensory attractants is central to most animal-mediated pollination and seed dispersal interactions. Approximately
half the 73 species of mosses’ in the family Splachnaceae are entomophilous (have their spores dispersed by flies) and are
coprophilous (grow on feces and carrion). When mature, entomophilous species often produce brightly coloured, scented sporophytes
which, for several species, have been shown to attract flies. In a number of cases, sporophyte colours and odours, as well
as the flies that visit them, have been shown to be species-specific, suggesting that the mosses co-exist by signal diversification,
just as flowering plants are thought to reduce competition for pollinators. Analyses of scent chemistry identified an odour
contrast between generations; gametophytes were either unscented or weakly scented in most species, whereas sporophyte odours
were universally stronger per unit mass and much more chemically complex. Sporophyte odours of North and South American species
sampled were both complex and diverse, with an apparent inverse relationship between the size and showiness of the apophysis
and its odour complexity. Furthermore, phylogenetic evidence suggests that fly dispersal of spores through visual and olfactory
signals has evolved multiple times in the Splachnaceae and that modifications of sporophyte morphology may have followed,
rather than triggered, the transitions to coprophily and entomophily. We review the ecological and evolutionary aspects ofentomophily,
with particular emphasis on the chemistry of sporophyte odours and the means by which they mimic decaying organic matter.
KeywordsDirected dispersal-Diptera-key innovations-mosses-sensory signals-Splachnaceae-spore dispersal
... Approximately half of the species in the moss family Splachnaceae are entomophilous, having their spores dispersed by flies (Diptera ;Koponen 1990;Marino 1991). ...
... The sporophytes of all entomophilous species so far examined release volatile chemicals that attract flies (Pyysalo et al. 1978(Pyysalo et al. , 1983Marino et al. 2009). Sporophyte odour in the Splachnaceae is novel among mosses, with both visual and olfactory cues appearing integral to fly attraction (Koponen 1990;Marino et al. 2009). The complex and diverse volatile compounds released by mature Splachnaceae hypophyses suggest chemical mimicry of herbivore dung among several genera (e.g. ...
... The cells in mature S. ampullaceum hypophyses resembled those of yellowing birch leaves, where the cytoplasm retreats to the edges of cell walls (Dodge 1969). The tissues of mature sporophytes appear to be undergoing senescence, which is to be expected as the capsule dehisces at this stage (Koponen 1990). ...
Many mosses of the family Splachnaceae are entomophilous and rely on flies for spore dispersal. Splachnum ampullaceum produces a yellow- or pink-coloured hypophysis that releases volatile compounds, attracting flies to the mature moss. The biosynthetic sources of the visual and aromatic cues within the hypophysis have not been identified, and may be either symbiotic cyanobacteria or chromoplasts that break down lipids into volatile compounds. Here, we used transmission electron microscopy and gas chromatography-mass spectrometry (GC-MS) to investigate the sources of these attractants, focusing on different tissues and stages of maturation. Microscopy revealed an abundance of plastids within the hypophysis, while no symbiotic bacteria were observed. During plant maturation, plastids differentiated from amyloplasts with large starch granules to photosynthetic chloroplasts and finally to chromoplasts with lipid accumulations. We used GC-MS to identify over 50 volatile organic compounds from mature sporophytes including short-chain oxygenated compounds, unsaturated irregular terpenoids, fatty acid-derived 6- and 8-carbon alcohols and ketones, and the aromatic compounds acetophenone and p-cresol. The hypophysis showed localised production of pungent volatiles, mainly short-chain fermentation compounds and p-cresol. Some of these volatiles have been shown to be produced from lipid oxidase degradation of linolenic acid within chromoplasts. However, other compounds (such as cyclohexanecarboxylic acid esters) may have a microbial origin. Further investigation is necessary to identify the origin of fly attractants in these mosses.This article is protected by copyright. All rights reserved.
... Insects are only recruited as dispersal agents by the Splachnaceae, and only those that grow on substrates of animal origin. Olfactory and visual cues attract flies foraging or looking for these substrates to lay their eggs (Koponen 1990). The chemical adaptation is complemented by morphological innovations such as sticky spores, a pseudocolumella which acts as a piston to elevate the spore mass to the mouth of the capsule or, more strikingly, the expansion of the sterile base of the urn either to amplify the visual cue or to provide a suitable landing platform for insects (Koponen 1990). ...
... Olfactory and visual cues attract flies foraging or looking for these substrates to lay their eggs (Koponen 1990). The chemical adaptation is complemented by morphological innovations such as sticky spores, a pseudocolumella which acts as a piston to elevate the spore mass to the mouth of the capsule or, more strikingly, the expansion of the sterile base of the urn either to amplify the visual cue or to provide a suitable landing platform for insects (Koponen 1990). Recent phylogenetic investigation suggests that entomophily, insect-mediated spore dispersal, arose early in the evolutionary history of the Splachnaceae, and was subsequently lost, maybe due to the severe biotic constraints shaping this system (Goffinet & Buck 2004). ...
... Insects are only recruited as dispersal agents by the Splachnaceae, and only those that grow on substrates of animal origin. Olfactory and visual cues attract flies foraging or looking for these substrates to lay their eggs (Koponen 1990). The chemical adaptation is complemented by morphological innovations such as sticky spores, a pseudocolumella which acts as a piston to elevate the spore mass to the mouth of the capsule or, more strikingly, the expansion of the sterile base of the urn either to amplify the visual cue or to provide a suitable landing platform for insects (Koponen 1990). ...
... Olfactory and visual cues attract flies foraging or looking for these substrates to lay their eggs (Koponen 1990). The chemical adaptation is complemented by morphological innovations such as sticky spores, a pseudocolumella which acts as a piston to elevate the spore mass to the mouth of the capsule or, more strikingly, the expansion of the sterile base of the urn either to amplify the visual cue or to provide a suitable landing platform for insects (Koponen 1990). Recent phylogenetic investigation suggests that entomophily, insect-mediated spore dispersal, arose early in the evolutionary history of the Splachnaceae, and was subsequently lost, maybe due to the severe biotic constraints shaping this system (Goffinet & Buck 2004). ...
With approximately 13 000 species, the Bryophyta compose the second most diverse phylum of land plants. Mosses share with the Marchantiophyta and Anthocerotophyta a haplodiplobiontic life cycle that marks the shift from the haploid-dominated life cycle of the algal ancestors of embryophytes to the sporophyte-dominated life cycle of vascular plants. The gametophyte is free-living, autotrophic, and almost always composed of a leafy stem. Following fertilization a sporophyte develops into an unbranched axis bearing a terminal spore-bearing capsule. The sporophyte remains physically attached to the gametophyte and is at least partially physiologically dependent on the maternal plant. Recent phylogenetic reconstructions suggest that three lineages of early land plants compose an evolutionary grade that spans the transition to land and the origin of plants with branched sporophytes (see Chapter 4). The Bryophyta seem to occupy an intermediate position: their origin predates the divergence of the ancestor to the hornworts and vascular plants but evolved from a common ancestor with liverworts (Qiu et al. 2006). The origin of the earliest land plants can be traced back to the Ordovician and maybe the Cambrian (Strother et al. 2004). Although unambiguous fossils of mosses have only been recovered from sediments dating from younger geological periods (Upper Carboniferous), divergence time estimates based on molecular phylogenies suggest that the origin of mosses dates back to the Ordovician (Newton et al. 2007) and thus that their unique evolutionary history spans at least 400 million years.
... y S. luteum (Hewd.) (Koponen 1990). ...
... Sin embargo, este mecanismo de dispersión biótica de esporas en musgos ha sido estudiado en forma sistemática exclusivamente en el Hemisferio Norte. Datos preliminares sugieren que este fenómeno podría ocurrir en la especie Tayloria purpurascens de Nueva Zelanda (Koponen 1990). La concentración del período de liberación de esporas desde los esporofitos de T. dubyi a comienzos del verano austral en coincidencia con los meses de mayor actividad de dípteros potencialmente dispersantes de esporas, sugiere que la entomocoría también podría ocurrir en la Reserva de Biosfera Cabo de Hornos, en la ecorregión subantártica de Magallanes (Jofre 2009). ...
The sub-Antarctic Magellanic ecoregion harbors a high diversity of bryophytes, greater than the species richness of vascular plants. Despite this fact, phenological studies on bryophytes are lacking for this ecoregion and Chile. Based on the study of the sporophytic phase of Tayloria dubyi, an endemic moss from the sub-Antarctic Magellanic ecoregion, we propose a methodology for phonological studies on austral bryophytes. We defined five phenophases, easily distinguishable with a hand-lens, which were monthly recorded during 2007 and 2008 in populations of T. dubyi at the Omora Ethnobotanical Park and Mejillones Bay on Navarino Island (55o S) in the Cape Horn Biosphere Reserve. The sporophytic (or reproductive) phase of T. dubyi presented a clear seasonality. After growing in November, in three months (December- February) of the austral reproductive season the sporophytes mature and release their spores; by March they are already senescent. T. dubyi belongs to the Splachnaceae family for which entomochory (dispersal of spores by insects, specifically Diptera) has been detected in the Northern Hemisphere. The period of spores release in T. dubyi coincides with the months of highest activity of Diptera which are potential dispersers of spores; hence, entomochory could also take place in sub-Antarctic Magellanic ecoregion. In sum, our work: (i) defines a methodology for phenological studies in austral bryophytes, (ii) it records a marked seasonality ion the sporophyte phase of T. dubyi, and (iii) it proposes to evaluate in future research the occurrence of entomochory in Splachnaceae species growing in the sub-Antarctic peatlands and forest ecosystems in the Southern Hemisphere.
... In the order Splachnales, most species belong to the family Splachnaceae, with numerous entomophilous species growing on dung or other animal substrates. Adaptations to coprophily and spore dispersal by flies have evolved several times within the family and are linked with several morphological characters, such as colorful expanded apophyses, volatile compounds attractive to insects, and sticky spores (Koponen, 1990. The sporophyte structure in the Splachnaceae is usually characteristic, with a more or less widened apophysis, or capsule neck. ...
Mosses (division Bryophyta) are characterized by the dominance of haploid, poikilohydric gametophytes, and relatively persistent sporophytes that are dependent on the gametophyte generation. Because they are poikilohydric, tend to be desiccation tolerant, and have primarily ectohydric water uptake mechanisms, the ecological requirements of mosses tend to differ from those of vascular plants. We review recent research on phylogenetic relationships and morphological and ecological diversity among mosses. Ancestral character state reconstructions illustrate the evolution of habitat preferences and the relationships between these and morphological variation. These reconstructions reveal the convergent evolution of both epiphytic and aquatic habitat preferences, as well as several reversals from epiphytic to other terrestrial habitats. Morphological character states connected to ectohydry, such as lack of water-conducting stem central strands, may be more prone to adaption driven by environmental conditions, while connections between endohydry and environmental conditions remain ambiguous and require further study. The distribution of the most elaborate endohydric water-conducting structures may be phylogenetically determined rather than resulting from adaptation to habitats. Among the early diverging lineages in the Bryophyta, shifts to cladocarpy may be connected to epiphytic lifestyles. We discuss ecological drivers of aspects of plant architecture including acrocarpous, pleurocarpous and cladocarpous perichaetial positions, and sporophytic reductions, the latter being common in dry, frequently disturbed terrestrial environments and in epiphytic habitats, but also occasionally found in aquatic habitats.
... Otro caso de interacciones dinámicas se registra en el musgo del estiércol Tetraplodon fuegianus, cuya distribución depende de la interacción con animales. La red de interacciones asociadas a la presencia o la ausencia de estiércol o restos de animales en descomposición, las moscas que se posan sobre éstos y dispersan las esporas de Tetraplodon sobre sustratos ricos en nutrientes, influyen directamente sobre los patrones de distribución de este musgo en los Bosques en Miniatura (Koponen, 1990). ...
Bryophytes (mosses, liverworts, and hornworts) and lichens have been undervalued in conservation and environmental education programs, and by modern culture in general. So much so, that most bryophytes and lichens lack a common name. We present a new methodology and activity to foster an ecological understanding of biodiversity, as well as of biocultural conservation and ethics, which includes little, under-perceived, living-beings --such as bryophytes. If something has no name, it does not exist in the cultural realm. If something is named, then it exists in the cultural sphere; additionally, its existence is decisively influenced by its name. At the southern end of the Americas, the UNESCO Cape Horn Biosphere Reserve in Chile protects a world’s biodiversity hotspot for bryophytes. Here, the research team at the Omora Ethnobotanical Park has developed an innovative educational and ecotourism activity to appreciate bryophytes: Cultivating a garden of names. This activity has been developed with the methodological approach of Field Environmental Philosophy. It encourages students and park’s visitors to engage physically as well as culturally (including the symbolic-linguistic dimension) with bryophytes and lichens, their life-habits and habitats. Park’s visitors are invited to observe, draw, and create names for bryophytes and lichens by using: (i) a magnifying glass or hand-lens to amplify the biophysical features of small plants, and (ii) the conceptual lenses of the biocultural ethics to broaden their understanding about how to interpret and respect the natural world. Through this activity, visitors connect to living beings that were previously under-perceived, and they come to see, value and care for a biophysical reality that is diverse, beautiful, and performs ecological functions that are essential to ecosystem integrity and human well-being. Cultivating a garden of names fosters a change of scientific and ethical perspectives to include little-perceived groups of organisms into the goals of conservation and environmental education.
... The exoskeleton of insects is rather well-studied (Hepburn 1972;Andersen 1979;Gorb 2001;Vincent & Wegst 2004), but there are no data to support the idea that such infraspecific variation exists. Yet, symbiotic associations involving morphological and ecological adaptations have been found among mosses and flies, such as in the Splachnaceae (Koponen 1990), and symbioses involving phoresy of plants or other sessile organisms on mobile animals are frequent among marine organisms (Stachowicz & Hay 1999). ...
Epizoic foliicolous liverworts and lichens are reported for the first time from insects, two species of the Shield Mantis Choeradodis, C. rhombicollis and C. rhomboidea (Mantodea: Mantidae: Choeradodineae) in Costa Rica. Five species of liverworts, 23 species of lichens, and several unidentified fungi were found growing mainly on the pronotum, but also on the fore wings, of 60 out of 84 individuals collected at various locations in Costa Rican lowland rainforests. The amount of epizoic colonization and the development of the epizoic “epiphyllous” communities were more pronounced in C. rhombicollis versus C. rhomboidea and in females versus males. These differences are potentially explained by a longer lifespan of females, which according to the development of the lichen thalli is suggested to be up to 24 months or more, while males show colonization that mostly corresponds to a lifespan of 12 months or less. While the occurrence of epizoic “epiphylls” in leaf-like insects suggests an enhanced effect of camouflage, we conclude that this occurrence is opportunistic rather than indicating an evolutionary phenomenon.
... obs.). However, the comparatively high moisture requirements of many coprophilous species (Koponen, 1990) imply they would require sheltered microhabitats with sufficient moisture at harvested sites. Although other studies have documented detrimental effects of harvesting on tree-colonizing epiphytes (Hazell et al., 1998;Caners et al., 2010), this study showed that epiphytes were unrelated to retention level. ...
Biological traits are potentially important for understanding mechanisms of plant species responses to alteration of local habitat conditions through natural and anthropogenic disturbance. Forest harvesting is a prominent disturbance in the circumpolar boreal biome, influencing stand- and landscape-scale patterns of forest structure and biodiversity. We examined a range of variable retention harvesting intensities (10%, 50%, and 75% dispersed green-tree retention harvesting and unharvested controls) in terms of their effectiveness for maintaining mosses and liverworts with differing biological traits. Bryophytes were sampled in 20 m radius plots 5–6 years post-harvest in 24 forest stands (each 10 ha) of two forest types (broadleaf-coniferous mixedwood, coniferous-dominated). We first examined the environmental factors that were the strongest predictors of species composition across the forest types and retention levels. We then used fourth-corner analysis to relate differences in the forest environment to species traits. Selected traits included bryophyte group, life form, habitat requirements, and reproductive and dispersal characteristics. The strongest predictors of species composition were ground-level moisture (estimated using growth of the moss Hylocomium splendens) and degree of canopy cover. Fourth-corner analysis showed that forest type, retention level, and their associated moisture conditions were closely related to the abundances of species characterized by different biological traits. Species with rare sporophyte production, larger spores, dioicous sexuality, or that require greater moisture or shade, were affiliated with higher retention and forest moisture. Reduced abundances of species with these traits after harvesting may detrimentally affect their capacity to disperse and re-establish, and suggests that moisture limitation is an important environmental filter that may restrict their representation at harvested sites. Coniferous-dominated forests supported higher abundances of several species types compared to mixed forests, including liverworts, acrocarpous mosses, and species that have greater moisture requirements, dioicous sexuality, or infrequent sporophyte production. This conveys the importance of coniferous forests as bryophyte habitat in mixedwood landscapes and the influence of canopy composition on regional species distributions. Understanding the tolerances of species exhibiting particular traits after harvesting may improve predictions about species extirpation risk and inform approaches to ensure their continued survival.
... Some of the main characteristics of the Splachnaceae are linked to entomophily, namely the large, brightly coloured apophysis and the emission of volatile compounds ( Koponen et al., 1990;Pyysalo et al., 1978Pyysalo et al., , 1983; capsule walls that shrink and the presence of a pseudocolumella which ensures that spores are pushed out of the capsule; "sticky" spores which clump together and attach to the bodies of insects (Koponen & Koponen, 1978;Koponen, 1982a); and reflexed or erect peristome teeth after dehiscence so that spores come into contact with the insects' bodies (Koponen, 1977(Koponen, , 1982a(Koponen, , b, 1990Koponen & Koponen, 1978;Vitt, 1981). The relationships between members of the Splachnaceae and their organic substrates have been investigated in some detail (Cameron & Wyatt, 1989;Gonzalez et al., 2006;Koponen, 1990;Marino, 1988a, b;Marino et al., 2009). In contrast, gametogenesis and vegetative propagation in members of this family have not been widely documented (Duckett et al., 2004;Mallón et al., 2006). ...
Tayloria rudolphiana (Garov.) Bruch et Schimp. was grown in axenic culture, from spores and gametophyte fragments to study its development patterns for the first time. It was grown for 16 months on Parker’s growth media under a 14h light/16°C-10h dark/14°C cycle with daylight fluorescent lighting. The expansion of protonemata filaments and branch formation in T. rudolphiana followed the typical tip growth pattern seen in mosses. All types of protonemata cells were observed (chloronemata, caulonemata and rhizoids) in specific developmental sequences, depending on their origin. Protonemata (caulonema) derived brood cells were observed for the first time in T. rudolphiana. Brood cells formed at the ends of the caulonemal filaments as chains of short, relatively thick-walled, spherical cells, containing abundant chloroplasts and some lipid droplets. Brood cells developed after 4 months in culture on colonies initiated from spores.
... By convergently mimicking the odour, appearance, and sometimes elevated temperature of these substrates, members of the Asclepiadaceae (Stapelia spp. and relatives), Aristolochiaceae (Burgess et al. 2004), Rafflesiaceae, Araceae (Lack & Diaz 1991;Yafuso 1993;Albre et al. 2003;Chouteau et al. 2008;Urru et al. 2010), Malvaceae (van der Pijl 1953), Hydnoraceae, Taccaceae, Burmanniaceae, Orchidaceae (Dressler 1981;Pemberton 2010) and Splachnaceae (a family of Bryophyta, see Koponen 1990) successfully attract these pollen (or spore) vectors, and mostly provide no reward (Faegri & van der Pijl 1979). ...
This paper comprises Part II of a review of flower visitation and pollination by Diptera (myiophily
or myophily). While Part I examined taxonomic diversity of anthophilous flies, here we consider the rewards and
attractants used by flowers to procure visits by flies, and their importance in the lives of flies. Food rewards such as
pollen and nectar are the primary reasons for flower visits, but there is also a diversity of non-nutritive rewards such
as brood sites, shelter, and places of congregation. Floral attractants are the visual and chemical cues used by Diptera
to locate flowers and the rewards that they offer, and we show how they act to increase the probability of floral
visitation. Lastly, we discuss the various ways in which flowers manipulate the behaviour of flies, deceiving them to
visit flowers that do not provide the advertised reward, and how some flies illegitimately remove floral rewards
without causing pollination. Our review demonstrates that myiophily is a syndrome corresponding to elements of
anatomical, behavioural and physiological adaptations of flower-visiting Diptera. The bewildering diversity of
anthophilous Diptera and of the floral attractants and rewards to which they respond allows for only broad
generalizations on myiophily and points to the need for more investigation. Ecological relationships between flies
and flowers are critical to the survival of each group in many habitats. We require greater understanding of the
significance of flies in pollination, especially in the face of recent pollinator declines.
... Another type of relationship is that of zoochory, only recorded in moss families located in temperate regions: Splachnaceae, whose spores are dispersed by Diptera in Europe (Koponen 1990, Lloret 1990) and Chile (Jofre et al. 2011), and Schistostegaceae, by several arthropods (spiders, insects) in Europe (Ignatov and Ignatova 2001), or liverworts and Cyanobacteria by Arachnida (Machado and Vital 2001) and Algae and bryophytes by lizards (Gradstein and Equihua 1995) in México, and mosses by Coleoptera in Oceania (Gradstein et al. 1984). ...
In a Nature Reserve called Reserva Natural Río Nambí (Nariño, Colombia), a platyrhacid millipede population of Psammodesmus bryophorus Hoffman, Martínez & Flórez, 2011 was discovered with 10 epizoic bryophyte species from five families: Fissidentaceae, Lejeuneaceae, Metzgeriaceae, Leucomiaceae and Pilotrichaceae. The inspected sample included 22 P. bryophorus individuals of which 15 were carrying mosaics of different bryophyte species on their dorsa, principally Lepidopilum scabrisetum, Lejeunea sp. 1 and Fissidens weirii. This finding constitutes the first record of epizoic plants on Diplopoda.
... In addition, forest floor bryophytes have been shown to respond to variation in pH (Zamfir et al. 1999), microtopography (Økland 1994), and soil moisture (Robinson et al. 1989). Although these studies have been integral to our understanding of habitat preferences and requirements of many bryophyte species (Kimmerer 1993;Koponen 1990;Söderström 1988Söderström , 1993 their focus on either specific microsite types or certain bryophyte taxa have limited our ability to untangle the relative importance of various factors in determining bryophyte species assemblages. ...
This study examined bryophyte community composition in relation to microsite and microenvironmental variation at different scales in three conifer-dominated stands in the boreal forest of Alberta, Canada. We documented bryophyte assemblage on specific microsite types (physiognomic forms providing substrates for moss colonization: logs, stumps, tree bases, undisturbed patches of forest floor, disturbed patches of forest floor), and at coarser scales: mesosites (625 m2 plots within stands), and stands (10 ha). Patterns of variation in bryophyte composition arising from the microsite sampling were clearly related to microsite type and, for woody substrates, to microsite quality (decay class; hardwood vs. softwood). Microenvironment (moisture, pH, temperature, light) also had some influence on bryophyte composition of woody microsite types. Forest floor moisture, pH, and light were related to bryophyte composition of undisturbed patches of forest floor while forest floor moisture and temperature were significant correlates for disturbed forest floor. At the coarser-scale, surface moisture and forest floor moisture were related to bryophyte assemblage of mesosites; this was partially reflective of differences among stands. We conclude that bryophyte species composition in these forests is related to a hierarchy of factors including fine scale variation in the type and quality of available microsites along with microenvironmental variation at different scales. Management efforts to preserve bryophyte biodiversity will need to incorporate this complexity.
... For some species of Splachnum Hedw. evidence of entomophily was provided [1][2][3][4][5][6][7][8][9]. They have many unusual adaptations for the dispersal of spores that allure coprophilous flies, including brightly coloured and expanded hypophyses of the capsules which emanate odours and volatile compounds and eject secretions [10][11][12][13]. ...
Splachnum pensylvanicum (Brid.) Grout ex H.A.Crum is recorded for the first time in Lithuania and it is its fourth discovery at a third locality in Europe. It was found for the first time in 2000 in Kamanos mire, the largest peatland complex in the northern part of this East Baltic country. Targeted investigations at this site in 2017 resulted in the discovery of 14 populations and it is apparently the largest and most abundant locality of the species in Europe. Splachnum pensylvanicum is briefly described and illustrated along with some taxonomic notes and a detailed description of its habitat requirements. The global geographical distribution of S. pensylvanicum is reviewed and mapped. It is a Euro-Eastern North American temperate species and deeply penetrates into the Neotropics at montane stations in Venezuela and SE Brazil in South America.
... Bryophytes disperse by small spores with a diameter that typically ranges between 10 and 30 µm, and asexual propagules . They primarily disperse by wind (Barbé et al., 2016), although evidence for the role of animal-mediated dispersal, including epizoochory (Chmielewski and Eppley, 2019; Koponen, 1990;Pauliuk et al., 2011) and endozoochory (Boch et al., 2015;Parsons et al., 2007;Russo et al., 2020;Wilkinson et al., 2017), has been increasingly acknowledged. It has therefore been widely accepted that bryophytes exhibit high dispersal capacities (Lönnell et al., 2012) and in fact, similar values of turnover were actually observed among island bryophyte communities and those expected under a null model, according to which species can randomly disperse among islands (Liu et al., 2020). ...
Climate change has and will have a dramatic impact on species ranges. Terrestrial species have accordingly already migrated poleward at a median speed of 16.9km per decade since the beginning of the industrial era. However, many species are not equipped to efficiently track the geographic changes of the conditions
matching their climatic niche, and are consequently prone to extinction. No less than about 20% of land plant species are hence threatened with extinction in the future, with major consequences on human food resources and health. In this context, Species distribution models (SDMs) offer an appealing framework to test the potential effects of climate change on species ranges. Like many biodiversity analyses, SDMs have traditionally been conducted at the species level. Cryptic speciation, which results in taxa that cannot rapidly be distinguished morphologically, but underwent divergent evolutionary histories, has been, however, increasingly reported, raising the question of whether SDMs should be fitted at the level of species (clade models), cryptic species or intraspecific lineages (subclade models). Projecting models through time further raises several questions and relies on several assumptions. In particular, projecting
species potential ranges in the future based on their niche inferred from extant climate conditions onto future climatic layers involves that (I) species climatic niches are conserved through time (niche
conservatism hypothesis) and that (ii) species are at equilibrium with their environment (i.e. their entire niche is filed), implying that they are not limited by their dispersal capacities, and are immediately able to
colonize any newly suitable area.
Focusing on bryophytes, whose ecophysiological characteristics, such as poikilohydry and reliance on rainfall for water uptake, make them excellent candidates to study the impact of climate change, but which exhibit reduced morphologies, raising concerns about broadly defined morphological species concepts, we address here the following questions:
1 At which taxonomic level should SDMs be computed? We compare the extent to which model projections generated at the level of species differ from those obtained for intraspecific lineages. Modelling at the level of infraspecific lineages raises a second issue, which is associated with the very small sample sizes that typically characterize molecularly defined lineages, that is: how can ensemble of small models calibrated from very small datasets be evaluated? In the light of analyses of niche overlap, we finally determine whether models should be calibrated at the level of the species or intraspecific lineages.
2 Is there climatic niche conservatism in bryophytes, and how does the tendency for closely related taxa to share the same climatic niche vary at increasing taxonomic depth?
3 To what extent will such efficient dispersers as bryophytes successfully track the shift of their suitable areas during the next decades? To address Q1, Ensembles of Small Models were evaluated by null models calibrated from randomly sampled presence points. We compared the extent of suitable area predicted by the projections of clade and subclade models. Niche overlaps were quantified using Schoener's D and Hellinger's I metrics, and the significance of these metrics in terms of niche conservatism or divergence was assessed by niche similarity tests. Combined predictions from subclade models contributed, on average, five times more than clade models to the total suitable area predicted by the combination of both subclade and clade models. Niche overlap was 0.71 on average, with evidence for niche conservatism in half of the species and no signal for niche divergence. Given the poor performance of models based on small datasets, we pragmatically suggest that, in the absence of evidence for niche divergence during diversification of closely related intraspecific
lineages, SDMs should be based on all available occurrence data at the species level. The hypothesis of climatic niche conservatism and its evolutionary ‘labillity’ was further tested at the level of an entire phylum of land plants, the Marchantiophyta, through analyses of the relationship between the spatial turnover of floras and macroclimatic variation. Phylogenetic turnover among floras was quantified
through πst statistics. πst-through-time profiles were generated at 1 myr intervals along the phylogenetic time-scale and were correlated with current geographic distance and macroclimatic variation with Mantel tests based on Moran spectral randomization to control for spatial autocorrelation. The contribution of macroclimatic variation to phylogenetic turnover was about four-times higher than that of geographic distance. The correlation between phylogenetic turnover and geographic distance rapidly decayed at increasing phylogenetic depth, whereas the relationship with macroclimatic variation remained constant until 100 myrs. Our analyses reveal that changes in the phylogenetic composition among liverwort floras across the globe are primarily shaped by macroclimatic variation. They demonstrate the relevance of macroclimatic niche conservatism for the assembly of liverwort floras over very large spatial and evolutionary time scales, which may explain why such a pervasive biodiversity pattern as the increase of species richness towards the tropics also applies to organisms with high dispersal capacities.
Finally, we developed a newly designed spatially-explicit model of dispersal by wind in the context of changing climate and presented an example of application in the case of the European flora. A grid of pixel specific environmental conditions and dispersal kernels, combining information on species dispersal traits, local wind conditions, as well as landscape features affecting dispersal by wind, was generated and used as input in simulations of species dispersal in the landscape under changing climate conditions. In European
bryophytes, the median ratios between predicted range loss vs expansion by 2050 across species and climate change scenarios ranged from 1.6 to 3.3 when only shifts in climatic suitability were considered, but increased to 34.7–96.8 when species dispersal abilities were added to our models. This highlights the importance of accounting for dispersal restrictions when projecting future distribution ranges and suggests that even highly dispersive organisms like bryophytes are not equipped to fully track the rates of ongoing climate change in the course of the next decades.
... This is the only moss family with entomophilous spore dispersion in some species (Marino et al. 2009). Koponen (1990) in a review of the topic "Entomophily in the Splachnaceae" cited several authors who demonstrated that several species of this family possess volatile compounds for attracting flies and butterflies to facilitate its spore dispersion. It is a very interesting group to studies in ecology and co-evolution. ...
We recognize four genera and six species of Splachnaceae in Brazil. Tetraplodon itatiaiae Müll.Hal. is proposed as a new synonym of Tetraplodon mnioides (Hedw) Bruch & Schimp., and Tetraplodon tomentosus Sehnem new synonym of Leptodontium capituligerum Müll.Hal. Three species are recognized as endemic: Moseniella ulei (Müll.Hal. ex Broth.) A. Kop., M. brasiliensis Broth., and Tayloria arenaria (Müll.Hal.) Broth. The species Tayloria scabriseta (Hook.) Mitt. is a new record for the country. We provide identification keys, comments, and diagnostic illustrations, as well as comments on their conservation status.
... About one half of extant representatives of the moss family Splachnaceae are known to develop on decaying organic matter, particularly animal dung (Koponen and Koponen 1978; Cameron and Wyatt 1986; Marino 1988 Marino , 1991a Marino , 1991b Marino , 1997 Koponen 1990). These so-called dung mosses have evolved unique suites of morphological and chemical adaptations to insect-mediated spore dispersal. ...
Chemical mimicry is an essential part of certain interspecific interactions, where the outcome for both species may depend on the degree to which the original signals are mimicked. In this review, we discuss a number of specific cases relating to pollination and obtaining nutrient resources that we believe exemplify recent advances in our understand- ing of chemical mimicry. Subsequently, we suggest avenues for future ecological and chemical research that should allow us to gain further insight into the evolution of chemical mimicry.
... The flies are attracted to the moss through both visual and chemical cues that mimic dung and or carrion and inadvertently pick-up the sticky spores, which may fall off when flies land on the natural substrate. This fly mediated spore dispersal syndrome provides an efficient means of local to regional dispersal (Koponen, 1990;Marino et al., 2009). ...
... Moss spores are anemochorous with the only two known exceptions for zoochory by Splachnaceae (Koponen, 1990;Marino, 2009;McCuaig et al., 2015) and Schistostega (Ignatov & Ignatova, 2001). Despite recent literature reported a number of cases of moss spore carriage by birds (Lewis et al., 2014;Chmielewski & Eppley, 2019) and small mammals (Barbé et al., 2016), nothing is still known about the efficiency of this way of dispersal. ...
... This assumption is supported by the revealed fact of a limited ability of spores for wind dispersal (Ignatov & Ignatova 2001). Its spores are sticky, exposed at the capsule mouth and have a structure extremely similar to that of the well-known entomochorous spores of the genera Splachnum Hedwig (1801: 51), Tetraplodon Bruch & Schimp. in Bruch, Schimper & Gümbel (1844: 211) and Aplodon Brown (1823: 41) of the Splachnaceae (Koponen 1990). Even after long drying, Schistostega spores cannot be blown out by wind (Fig. 1C). ...
This is the first record of the intentional consumption of moss spores by an arthropod. A moss mite Kunstidamaeus lengersdorfi (Oribatida: Damaeidae) was observed in Moscow Province, Russia, as regularly visiting mature capsules of the moss Schistostega pennata (Schistostegaceae) and eating spores from the opened capsules. The spores of Schistostega are thin-walled, sticky, not dispersed by wind, and exposed at the capsule mouth for easy consumption, being very similar in this respect to the spores of the entomochorous mosses of the Splachnaceae family. Walking through Schistostega population, these mites carry a lot of its spores of their bodies. According to published data, Kunstidamaeus is mostly feeding on saprotrophic fungi and green algae. Behavioral observations as well as the gut content study show that Kunstidamaeus lengersdorfi can also feed on spores of Schistostega pennata. This is the first record of Kunstidamaeus lengersdorfi in Russia. Previously it was known in Central Europe at cave entrances. In a parallel way, Schistostega grows in Central Europe mostly in caves, while it recently explosively spread in boreal, especially spruce-dominated forests in the Central European Russia. The available data indicate that the mite range expansion had followed the expansion of its forage plant.
... Species within some taxonomic groupings of bryophytes have strong ecological similarities. For example, a large proportion of the species in the Family Splachnaceae is dispersed by insects to the dung of herbivorous animals (Koponen 1990). In general, mosses of the genus Sphagnum are found in wet acidic habitats (Schofield 1985). ...
The sub-Antarctic Magellanic ecoregion harbors a high diversity of bryophytes, greater than the species richness of vascular plants. Despite this fact, phenological studies on bryophytes are lacking for this ecoregion and Chile. Based on the study of the sporophytic phase of Tayloria dubyi, an endemic moss from the sub-Antarctic Magellanic ecoregion, we propose a methodology for phonological studies on austral bryophytes. We defined five phenophases, easily distinguishable with a hand-lens, which were monthly recorded during 2007 and 2008 in populations of T. dubyi at the Omora Ethnobotanical Park and Mejillones Bay on Navarino Island (55º S) in the Cape Horn Biosphere Reserve. The sporophytic (or reproductive) phase of T. dubyi presented a clear seasonality. After growing in November, in three months (December-February) of the austral reproductive season the sporophytes mature and release their spores; by March they are already senescent. T. dubyi belongs to the Splachnaceae family for which entomochory (dispersal of spores by insects, specifically Diptera) has been detected in the Northern Hemisphere. The period of spores release in T. dubyi coincides with the months of highest activity of Diptera which are potential dispersers of spores; hence, entomochory could also take place in sub-Antarctic Magellanic ecoregion. In sum, our work: (i) defines a methodology for phenological studies in austral bryophytes, (ii) it records a marked seasonality ion the sporophyte phase of T. dubyi, and (iii) it proposes to evaluate in future research the occurrence of entomochory in Splachnaceae species growing in the sub-Antarctic peatlands and forest ecosystems in the Southern Hemisphere.
Many dung mosses (Splachnaceae) are characterized by insect-mediated spore dispersal. All of the entomophilous 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. Communicating Editor: Aaron Liston
The moss Tayloria dubyi (Splachnaceae) is endemic to the subantarctic Magallanes ecoregion where it grows exclusively on bird dung and perhaps only on feces of the goose Chloephaga picta, a unique habitat among Splachnaceae. Some species of Splachnaceae from the Northern Hemisphere are known to recruit coprophilous flies as a vector to disperse their spores by releasing intense odors mimicking fresh dung or decaying corpses. The flies land on the capsule, and may get in contact with the protruding mass of spores that stick to the insect body. The dispersal strategy relies on the spores falling off when the insect reaches fresh droppings or carrion. Germination is thought to be rapid and a new population is quickly established over the entire substrate. The objectives of this investigation were to determine whether the coprophilous T. dubyi attracts flies and to assess the taxonomic diversity of the flies visiting this moss. For this, fly traps were set up above mature sporophyte bearing populations in two peatlands on Navarino Island. We captured 64 flies belonging to the Muscidae (Palpibracus chilensis), Tachinidae (Dasyuromyia sp) and Sarcophagidae (not identified to species) above sporophytes of T. dubyi, whereas no flies were captured in control traps set up above Sphagnum mats nearby.
The moss family Splachnaceae is characterized by half of its members relying on insects for spore dispersal. These species grow on dung or other animal substrates. They produce small and aggregated spores, and their capsule is modified to attract coprophilous insects or carrion flies using olfactory and visual cues. Systematic concepts and implicit evolutionary inferences have relied much on variation in characters associated with the spore dispersal syndrome. Phylogenetic reconstructions based on sequence variation of two chloroplast loci (trnL-trnF region and the rps4 gene) suggest that most supraspecific taxa are poly- or paraphyletic. Transformations in morphological characters associated to the syndrome thus offer little if any phylogenetically informative signal. Brachymitrion is resolved in a nested position within Tayloria. A new combination, Tayloria immersa (Goffinet) Goffinet, Shaw & Cox is proposed for B. immersum. Only one of the five subgenera of Tayloria (subg. Orthodon) is potentially monophyletic. Voitia shares a common ancestor with Tetraplodon and is thus nested within the Splachnoideae. The affinities of Aplodon remain ambiguous. Reconstruction of shifts between wind and insect spore dispersal syndromes suggests that entomophily arose more than once and may have been followed by a reversal to the generalist strategy in two lineages.
The Antarctic Peninsula has experienced some of the most accelerated warming worldwide, resulting in the retreat of glaciers and creation of new areas for plant development. Information regarding the plant dispersal processes to these new niches is scarce in Antarctica, despite birds being important vectors elsewhere. Many bird pellets (with feed remains such as bones and feathers) are generated annually in Antarctica, which are light and easily transported by the wind and include vegetation that is accidentally or purposely swallowed. The aim of this study was to analyze the presence of plant fragments within skua (Stercorarius/Catharacta spp.) pellets collected from two sampling areas in the Maritime Antarctic: Stinker Point (Elephant Island, 17 samples) and Byers Peninsula (Livingston Island, 60 samples), in the South Shetland Archipelago, during the austral summers of 2018 and 2020. In both study areas, five species of Bryophyta were found that were associated with the pellets and viable in germination tests in a humid chamber. The ingestion of Bryophyta for the skuas contribute to the dispersion of different moss species, including to areas recently exposed by the ice retreat. This is the first demonstration that skua pellets effectively act in the dispersion of Antarctic mosses.
Mathematical models for bit life of polycrystalline diamond compact (PDC) drill bits were developed for drilling in hard abrasive media. Based on the wear-out criterion of an average 0.060 in. (1.5 mm) flank wear land, bit life equations were formulated in three forms: bit life versus rotary speed and feed rate, bit life versus rotary speed and penetration rate, and wear rate versus cutting speed and cutter engagement area. The traditional linear-logarithmic model provided the best bit life prediction equation. Consequently, it would be possible to predict the optimum economical drilling conditions more accurately by employing a quadratic-logarithmic based bit life equation. The equation demonstrated the ability to predict the bit life precisely under different modes of wear.
We evaluate the influences of gametophyte size, taxonomic affiliation (as an indicator of phylogenetic relationship), and water relations on variation and covariation of five life history traits among 335 moss species representing the orders Funariales, Polytrichales, and Pottiales. Size effects, though statistically significant, account for a relatively small proportion of the variation in most life history traits. Canonical Correlation Analyses, conducted across all species as well as within orders and larger families, reveal significant relationships between patterns of covariation in life history traits and variation in morpho-anatomical traits associated with water relations. Water relations account for 40-50% of life history variation depending on taxonomic group, but patterns of relationships between the two sets of variables are broadly similar, irrespective of the taxonomic group in which they were examined. The first water relations variate is a gradient describing water uptake/retention capacity. The life history variate associated with this is an axis that arranges species from those that are monoicous, short-lived and produce few large spores to those having the opposite suite of traits. The second water relations variate represents an endo-ectohydric gradient, which correlates with a life history variate describing differential investment in spores as a function of life expectancy. We used Nested Analyses of Variance and Covariance to partition variance in the life history traits into taxonomic components and evaluate whether size and water relations differences among particular groups mediate apparent taxonomic effects. Generic influences on all traits are strong, and independent of size and water relations effects; together, genus and water relations account for >50% of the variation in each of the life history traits considered. Apparent ordinal effects largely disappear after adjustment for size and water relations, while substantial family level effects exist for spore number and spore size. Variation among species within genera is modest. These results suggest that most of the diversification in moss life histories occurred at the level of genus and that life histories have been relatively conserved in the process of subsequent cladogenesis. The persistent occurrence of water relations effects at several taxonomic levels, and the relatively large proportion of variance in life history traits, independent of taxonomic affiliation, for which these effects account, may imply adaptive coevolution of water relations and life history. These ideas can best be tested in an explicitly phylogenetic framework.
AimMany intercontinental disjunctions, especially among spore-producing plants, are shaped by long-distance dispersal (LDD) via wind currents. Amphitropical disjunctions are most commonly explained through LDD, but other vectors and dispersal scenarios must also be considered. To interpret the New World amphitropical disjunction in the dung-moss genus Tetraplodon, we compared stepwise migration along the Andes, direct LDD and ancient vicariance.LocationGlobal, specifically high-latitude and high-elevation localities, with a focus on the New World.Methods
Phylogenetic relationships were inferred from four loci sampled from 124 populations representing the global range of Tetraplodon, and analysed using maximum-likelihood and Bayesian optimality criteria, with divergence dates estimated in beast.ResultsThe monophyletic T. mnioides complex diversified between the early Miocene and early-to-mid Pliocene into three well-supported clades, each with a unique geographical distribution: Laurasian, primarily high-elevation tropical, and amphitropical. Populations from southernmost South American were reconstructed as a monophyletic lineage that diverged from high-latitude Northern Hemisphere populations around 8.63 Ma [95% highest posterior density (HPD) 3.07–10.11 Ma].Main conclusionsDirect LDD has resulted in the American amphitropical disjunction in Tetraplodon. A lack of modern or historical wind connectivity between polar regions and the poor resistance of Tetraplodon spores to the conditions associated with wind-dispersal suggest that bird-mediated LDD provides the best explanation for the establishment of amphitropicality.
The genus Pseudoptilolepis SNYDER, 1949 and the species P. confusa SNYDER, 1949 are recorded for the first time in Argentina, extending the Neotropical distribution of both taxa. The specimens were collected in Corrientes province, flying around the fruiting body of a stinkhorn fungus (Phallaceae). A modification to the key to Argentine Muscidae is given. Zusammenfassung Ein Fund von Pseudoptilolepis confusa SNYDER, 1949 in der Provinz Corrientes ist der erste Nachweis dieser Gattung und Art in Argentinien. Damit erweitert sich die bislang bekannte Verbreitung in der Neotropis. Exemplare von P. confusa wurden gesammelt, als sie den Frucht-körper einer Stinkmorchel (Phallaceae) umflogen. Der Bestimmungsschlüssel für die Muscidae Argentiniens wird modifiziert, um diesen Neufund aufzunehmen.
Many plants have geographic disjunctions, with one of the more rare, yet extreme being the amphitropical, or bipolar disjunction. Bryophytes (namely mosses and liverworts) exhibit this pattern more frequently relative to other groups of plants and typically at or below the level of species. The processes that have shaped the amphitropical disjunction have been infrequently investigated, with notably a near absence of studies focusing on mosses. This dissertation explores the amphitropical disjunction in the dung moss Tetraplodon, with a special emphasis on the origin of the southernmost South American endemic T. fuegianus. Chapter 1 delimits three major lineages within Tetraplodon with distinct yet overlapping geographic ranges, including an amphitropical lineage containing the southernmost South American endemic T. fuegianus. Based on molecular divergence date estimation and phylogenetic topology, the American amphitropical disjunction is traced to a single direct long-distance dispersal event across the tropics. Chapter 2 provides the first evidence supporting the role of migratory shore birds in dispersing bryophytes, as well as other plant, fungal, and algal diaspores across the tropics. Chapter 3 describes the complete chloroplast and mitochondrial genomes and nuclear ribosomal repeat across seven patches of the endemic T. fuegianus. Screening of variation within distinct patches of moss revealed inter-individual polymorphism within single patches of moss, and intra-individual variation in the nuclear ribosomal repeat. Chapter 4 employs a RAD-seq approach to sequence thousands of loci across the range of the amphitropical lineage inferred in Chapter 1 allowing for resolution of a monophyletic T. fuegianus, which shares an ancestor with populations from northwestern North America. Within the lineage, geographic structure is identified, suggesting a complex phylogeographic history for this group, likely shaped by Pleistocene glaciations in northwestern North America.
Life on Earth originated in the sea; thus, land is a frontier for aquatic organisms. Although colonization of land by plants occurred in the Ordovician about 450 million years ago (ma; Field et al. 2015), aquatic microorganisms colonized land as early as 3500 mya (Beraldi-Campesi 2013). When aquatic microorganisms (including cyanobacteria) headed to land, they encountered adverse terrestrial conditions such as drought, extreme diurnal and seasonal temperature changes, low nutrient supply, and strong sunlight. These microorganisms are considered to have achieved terrestrialization by developing a tough, pigmented cell wall, an agglutinated colony structure, and a symbiotic lifestyle. Although modern lichens are associations of internal algae with external advanced fungi belonging to Ascomycetes and Basidiomycetes, colonylike associations of microorganisms, including basal algae and fungi, are thought to have colonized land before the Phanerozoic. Colonization of terrestrial habitats by photosynthesizing multicellular organisms occurred in a clade of green plants containing chlorophyll a and b (Delaux et al. 2012). The order Charales in the division Charophyte adapted to life in ephemeral freshwater pools, which frequently dry up. The body of the Charales is a monoploid gametophyte and comprises a main axis and lateral branchlets occurring in whorls on each node of the axis (Fig. 2.1). The reproductive organs consist of the nucleus (i.e., archegonia-producing ova) and globules (i.e., antheridia-producing flagellate sperm; Fig. 2.2). The sperm swim to the archegonium and fertilize the ovum. The zygote develops into an oospore, which is protected by its tough cell wall containing an unusually stable polymer, sporopollenin. The oospore can resist desiccation by remaining in a dormant state, and undergoes meiosis before germination of protonema. Another charophyte order, Coleochaetales, is a parenchymatous disclike alga that grows on substrata in waterfront habitats (Fig. 2.1). In the Coleochaete, plural meiospores are produced from a zygote, suggesting that embryos are protected and nourished in maternal tissue (Graham and Wilcox 2000).
It is an exciting time for the study of obligate pollination mutualisms. New mutualisms continue to be discovered, and information on individual systems is rapidly growing. Presently, there are at least seven plant lineages apart from Phyllanthaceae that contain plants pollinated by seed-parasitic insects (Fig. 13.1, Table 13.1). There is little doubt that more such lineages will be discovered, inasmuch as one or two new mutualisms continue to be uncovered each decade. The abundance and heterogeneity of documented cases of obligate pollination mutualism offer an unprecedented opportunity to examine key topics of broad ecological and evolutionary relevance. In this chapter, we review the basic natural history of the seven mutualisms known outside of Phyllanthaceae (Fig. 13.1, Table 13.1) and address the following questions that are critical to our understanding of obligate pollination mutualisms.1.
Why do plants specialize to seed-parasitic pollinators despite the high cost imposed by the seed-feeding pollinator larvae?
2.
How are mutualisms maintained despite the potential for selfish partners to disrupt the interaction?
3.
Is pollinator specificity reinforced, and if so, why?
4.
Do obligate pollination mutualisms drive the reciprocal diversification of plants and pollinators?
Rapid advances in the field of bryophyte population dynamics include the development of non-destructive techniques to mark and monitor individual shoots, which allowed the construction and sophisticated use of matrix models and the analysis of the consequences of fine-scale dynamics and sexual reproduction for shoot survival and growth. Dissection of turfs of desert bryophytes yielded additional evidence, as well as more detailed knowledge on phenological patterns and year-to-year variation in investments in reproduction. Studies of population dynamics at larger spatial (metapopulations) and temporal (diaspore banks) scales included experimental and modelling work on spore dispersal and on conditions favourable to spore germination and establishment, as well as extension of current metapopulation models to explicitly incorporate the dynamics of the habitable patches themselves. Together with the rapid advance of molecular techniques allowing genet recognition and the analysis of parent-offspring relations, these new developments now open the way to study the dynamics of populations of genets, which is a necessary pre-requisite for a more in-depth analysis of rates and directions of processes of natural selection and evolutionary change.
Tayloria hornschuchii (Grev. et Arnott) Broth. is described for the first time as having either axillary or rhizoidal propagula or both and is reported for the first time from the Canadian Arctic Archipelago on Northeastern Ellesmere Island, N. W. T. A gametophytic comparison is made between T. hornschuchii and T. froelichiana (Hedw.) Mitt. to distinguish specimens without sporophytes. Leaf cell lengths prove to be a dependable character, with T. hornschuchii having significantly shorter basal and apical cell lengths. Propagula have never been found on T. froelichiana. North American distribution maps show T. froelichiana as low arctic-alpine and T. hornschuchii as high arctic-alpine.
Insect-mediated spore dispersal was quantified for Splachnum ampullaceum, S. rubrum, S. sphaericum, S. vasculosum, and Tetraplodon mnioides. Dipterans from the genera Scatophaga (Scatophagidae), Delia (Anthomyidae), Myospila (Muscidae), and Pyrellia (Muscidae) visited colonies and contacted sporophytes. Scatophagids, the most frequent and effective visitors, benefited from a possible increase in copulatory success as a result of visiting these plants. Measurements indicated that wind is not an effective agent of spore dispersal in S. rubrum. Consequently, spore dispersal adaptations and insect behavior are major factors restricting colonies of these mosses to organic substrates.
Insect-microbial interactions were studied during the first month of dung decomposition in Wyoming and Michigan. At both sites, screen cones placed over fresh dung were used to exclude insect colonists and confine normal field densities of Aphodius beetle adults or enough sarcophagid adults to produce normal larval densities. The effect of these insects on bacterial and hyphal densities as well as on fungal species numbers was assayed after the dung had been in the field 3 to 4 wk. Presence of maggots and Aphodius beetles increased bacterial and decreased hyphal density in Wyoming but not in Michigan. We hypothesize that these effects are due to insect mixing of the substrate, giving bacteria a competitive advantage over fungi. Normal insect colonization increased the number of fungal species per pat in Michigan and Wyoming, although the toW number of fungal species was not affected by treatments.
Splachnaceae grow primarily on dung and have spores that are dispersed to dung by Diptera. On a local scale, few mechanisms appear to promote coexistence of Splachnaceae on a dropping. Competition between mosses is strong and individual droppings tend to be occupied by a single species. On a regional scale, dispersal may promote habitat separation between the genera while coexistence in Splachnum spp. appears to be influenced by a tradeoff between dispersal and competitive ability. Coexistence in Tetraplodon spp. is facilitated by temporal separation of sporophyte production. Resource fluctuations may also promote coexistence. -from Author
The characteristics of pollen cementing (stickiness; total amount, structure and distribution of the pollenkitt; fine structure of the exine) have been studied in Salix caprea, Populus nigra and P. tremula (Salicaceae), Tilia platyphyllos and T. tomentosa (Tiliaceae), and Erica herbacea, Calluna vulgaris and Andromeda japonica (Ericaceae).
These thousand eyes have also been looking upon naturalists for quite a while, but only few have looked back. Usually they were zoologists interested in specific groups which live on, in or under bryophytes; in the role these animals play during initial land colonization by cryptogams; in freshwater associations and in diverse other aspects. Botanists have published far fewer observations, these dealing mainly with fertilization or spore dispersal by invertebrates. Although scattered and uneven, taken as a whole the compiled data offer suggestive insights into the relationships between bryophytes and invertebrates, especially in regard to their co-evolution.
Splachnum ampullaceum, S. luteum, S. rubrum, S. sphaericum, and S. vasculosum, and Tetraplodon angustatus, T. mnioides, and T. urceolatus are widely distributed from the east to the west coast of northern North America. However, the north-south distributions and the relative abundances of the species are not uniform across this range. Splachnum species differ as to whether they are most abundant in eastern (S. ampullaceum) or western North America (S. luteum and S. sphaericum) and, in their north-south distribution (S. vasculosum in arctic regions, S. luteum in boreal and arctic regions and S. rubrum in boreal regions). Tetraplodon species differ in their north-south distributions (T. urceolatus in arctic regions, T. mnioides in arctic and boreal regions, and T. angustatus in boreal regions) but are uniform in distribution from east-west. Tetraplodon species are more frequent than Splachnum species and of Splachnum species the most frequent are those with the least modified sporophytes. In those specimens where two or more Splachnaceae species occur intermixed, only the species combination of S. luteum and S. sphaericum was frequent.
Voitia hyperborea was first proposed by Greville and Arnott in 1822 on the basis of a specimen collected in 1820 on Melville Island in the Canadian Arctic Archipelago during one of the early British voyages of arctic exploration. Because of its restriction to arctic regions, this moss has remained poorly understood, and has been placed incorrectly in the synonymy of V. nivalis Hornsch., a species of the Alps, the Rocky Mountains, and Turkestan. However, the two species are quite separate and differ conspicuously, especially in their capsule shape, V. nivalis having an elongated, narrowly ovate-cylindrical capsule, and V. hyperborea a much shorter and broader capsule shaped like an inverted funnel. The patterns of geographical distribution of the two species are also entirely different. Illustrations are provided to show these morphological differences, as well as the geographic range of V. hyperborea in North America.
As the anomalous fleshy, hyaline, translucent seta of living sporophytes of Aplodon, resembling that of an hepatic, does not seem to have been reported previously, it is described and illustrated from material collected at Point Barrow, Alaska. At the end of the first season the swollen fleshy seta dehydrates and assumes the delicate, pale-yellow, silky condition seen in herbarium specimens and described in manuals. The known geographic distribution of this species is illustrated by a map that includes many recent collections, the majority of which were made within the Arctic Circle, a fact which thus further confirms the membership of this species in a typically high-arctic, circumpolar floristic element. The nomenclatural history of Aplodon wormskioldii is reviewed in order to elucidate why Robert Brown's original spelling, Aplodon, should be accepted instead of Haplodon, which under the present international code of botanical nomenclature has become an illegitimate name.
Field observations on colonies of Splachnum ampullaceum Hedw. in Isle Royale National Park, Michigan, and laboratory cultures maintained for three years without decline in vigor indicate a possible interrelationship between nutrient and pH factors to explain the habitual decline of colonies in nature.
The dark purple flower in the umbel of Daucus and the black spot on the umbel of Artedia have been shown to play a significant role in the pollination of these umbelliferae, because they attract the fly pollinators by their mimicry of resting insects.
The effect of pH on the growth of protonemata of Tetraplodon mnioides (Hedw.) B.S.G. and Funaria hygrometrica Hedw. was studied for 5 weeks at initial pH levels of 3.5, 5.8, 6.4, and 7.8 in two different media. Protonemata of both species increased in dry weight significantly at each succeeding pH. There was a significant difference in growth of the protonemata in the two media, but there was no significant difference in growth between the two species. Also, there was no significant interaction between any of the sources of error. pH may be important as a limiting factor in restricting the habitat of Tetraplodon mnioides and Funaria hygrometrica.
Some closely related members of the monocotyledonous familiesAlismataceae, Liliaceae, Juncaceae, Cyperaceae, Poaceae andAraceae with variable modes of pollination (insect- and wind-pollination) were studied in relation to the ultrastructure of pollenkitt and exine (amount, consistency and distribution of pollenkitt on the surface of pollen grains). The character syndromes of pollen cementing in entomophilous, anemophilous and intermediate (ambophilous or amphiphilous) monocotyledons are the same in principal as in dicotyledons. Comparing present with former results one can summarize: 1) The pollenkitt is always produced in the same manner by the anther tapetum in all angiosperm sub-classes. 2) The variable stickiness of entomophilous and anemophilous pollen always depends on the particular distribution and consistency of the pollenkitt, but not its amount on the pollen surface. 3) The mostly dry and powdery pollen of anemophilous plants always contains a variable amount of inactive pollenkitt in its exine cavities. 4) A step-by step change of the pollen cementing syndrome can be observed from entomophily towards anemophily. 5) From the omnipresence of pollenkitt in all wind-pollinated angiosperms studied one can conclude that the ancestors of anemophilous angiosperms probably have been zoophilous (i.e. entomophilous) throughout.
1.
Zur Trennung und zum Nachweis der flchtigen Amine wurde ein papierchromatographisches Verfahren entwickelt.
2.
Fr primre Amine ist ein Papier besonders geeignet, das mit 0,15 molarem Na-Acetat imprgniert ist. Sekundre und tertire Amine dagegen werden besser auf unvorbehandeltem Papier getrennt.
3.
Als Laufmittel werden n-Butanol/Wasser/Eisessig-Gemische (50/49/1 und 40/50/10) empfohlen. Collidin/Wasser (50/50) ist fr die Differenzierung der cyclischen Amine und der aliphatischen Diamine und Alkanolamine von den aliphatischen Monoaminen zu verwenden.
4.
Als Farbreagentien zur Sichtbarmachung der Flecken auf dem Chromatogramm haben sich bewhrt:
a)
Ninhydrin fr primre Amine
b)
Nitroprussidnatrium fr sekundre Amine
c)
Joddampf und Phosphormolybdnsure fr tertire Amine.
5.
Die Nachweisgrenzen liegen fr primre Amine bei 0,6 , fr sekundre bei 2 und fr tertire Amine bei 3 .
6.
Zur Kontrolle der papierchromatographischen Methode und zu ihrer Ergnzung diente das zweckentsprechend weiter entwickelte Identifizierungsverfahren ber die 2,4-Dinitro--naphthol-Salze der Amine.
7.
Es wurde die Flchtigkeit der Amine bei Wasserdampfdestillation aus alkalischem Medium geprft. Es zeigte sich, da die aliphatischen Monoamine bis C10, -Phenylaethylamin und Benzylamin, nicht aber die anderen cyclischen Amine, die aliphatischen Diamine und Alkanolamine ins Destillat bergehen.
The attachment of pollen grains among themselves, on the loculus wall, and on flower-visiting insects is quite different in entomophilous angiosperms using pollenkitt and those using viscin threads as pollen adhesives. The sticky and viscous pollenkitt makes the pollen grains adhere, while the thin, non-elastic, non-sticky, and flexible viscin fibers fasten them like ropes on insect hairs or bristles. Nectar vomited by honey-bees, sticky stigma secretions or other additional sticky substances further improve the pollen adherence to flower-visiting insects.
Tetraplodon paradoxus (R.Br.) Hagen was originally described from arctic Canada in 1823 andT. pallidus Hagen, described from arctic Scandinavia in 1893, was reported from the Canadian Arctic Archipelago in 1907. Although the
two species are distinct, clear-cut and easily distinguished from other members of the genus, they were almost hopelessly
confused both conceptually and nomenclaturally with each other and withT. mnioides (Hedwig) Bruch, Schimper & Gümbel by Hagen in 1910, and his erroneous treatment has been perpetuated by subsequent bryological
authors, especially in Scandinavia. Moreover, these two species have been omitted from all comprehensive North American manuals
and checklists; the purpose of this paper is to clarify the long-standing confusion and to establish a firm basis for their
recognition in the North American bryoflora, from which they have been excluded for far too long. The distinctions between
these two species are made clear by means of photographs made from living material, as well as by maps of their geographical
distribution in North America.
Most chemical signals in nature are volatile, although there is increasing evidence that insects can respond to involatile plant chemicals. Volatile signals, when mixed with other chemicals, may become more active due to synergism and/or become more persistent. Volatile scents and odours are generally efficient, since only trace amounts are needed to produce an effect. They are often multifunctional: a defence odour may also mediate in interactions between competing herbivores. Chemical signals are most important in reinforcing species—species (e.g. Ophrys-Andrena) interactions, but there are many examples of ‘accidental’ responses to plant odours.
Recent work on chemical signals will be reviewed, with respect to both beneficial and hostile plant-animal interactions.
Chemistry and Chemical Taxonomy of Bryophytes. Saarbrcken, 31 Aug. - 2 Sept. 1988, 25
Establishment and population dynamics of Tayloria tenuis in a Pyrenean forest University of Edinburgh 19-22
- F 8 Lloret
- J Lussenhop
- K Rabinder
- D T Wicklow
- J E Lloyd
LLORET, F., 1988. Establishment and population dynamics of Tayloria tenuis in a Pyrenean forest. International Symposium on Bryophyte Ecology, University of Edinburgh 19-22.July 1988. Abstracts: 8. LUSSENHOP, J., RABINDER, K., WICKLOW, D. T. & LLOYD, J. E., 1980. Insect cffccts on bacteria and fungi in cattle dung. Oikos, 34: 54-58.
Organngraphie der pfrnzen insbesondere der Archcgnniaten und SamenpJlanzen. 2. Auflage 11: HABERLANDT, G., 1886. Beitrage zur Anatornie und Physiologie der Laubmoose. Jahrbiicher fur HARBORNE Chemical signals in the ecosystem
- K Goebel
GOEBEL, K., 1915. Organngraphie der pfrnzen insbesondere der Archcgnniaten und SamenpJlanzen. 2. Auflage 11: HABERLANDT, G., 1886. Beitrage zur Anatornie und Physiologie der Laubmoose. Jahrbiicher fur HARBORNE, J. B., 1987. Chemical signals in the ecosystem. Annals of Botany, 60 (Suppl. 4 ) : 39-57.
Manual gf Byology. The Hague: Martinus NijhofT. GERSON, U., 1982. Bryophytes and invertebrates
- H Gams
GAMS, H., 1932. Bryocenology: In F. Verdoorn (Ed.), Manual gf Byology. The Hague: Martinus NijhofT. GERSON, U., 1982. Bryophytes and invertebrates. In A. J. E. Smith (Ed.), Bryophyte Ecologr: 291-332.
The peristome and spores in Splachnaceae and their evolutionary and systematic significance
- KOPONEN
KOPONEN, A., 1978. 'The peristome and spores in Splachnaceae and their evolutionary and systematic significance. Bryophytorum Bibliotheca, 13: 535-567.
Coexistence on dividend habitats: mosseS in the family Splachndceae The North American distribution of the circumboreal Splachnum and Tetraplodon
- P C Marino
- Annales
- P C Marino
MARINO, P. C., 1988a. Coexistence on dividend habitats: mosseS in the family Splachndceae. Annales <oologici MARINO, P. C., 1988b. The North American distribution of the circumboreal Splachnum and Tetraplodon. PFAEHLER, A., 1904. Etude biologique et morphologique sur la disskmination des spores chez les mousses.
Studies on the generic concept in the classification of the moss family Splachnaccac. Publicatiom from the Departmmt nf Botany
- A Koponen
KOPONEN, A., 1983. Studies on the generic concept in the classification of the moss family Splachnaccac. Publicatiom from the Departmmt nf Botany, Universify of Helsinki, 11: 1-48.
Evidence of entomophily in Splachnaceae (Bryophyta)
- KOPONEN
KOPONEN, A. & KOPONEN, T., 1978. Evidence of cntomophily in Splachnaceae (Bryophytd). Bryophytorum Bibliotheca, 13: 569-577.
Beobachtungen fiber das Ausstreuen der Sporen bei den Splachnaceen
- BRYHN
Mesofaunal responses to cattle dung with particular reference to Collembola
- DAVIDSON
Zur Verbreitungsokologie von Splachnum spkaericum (L. fil.) Swartz
- FISCHER
The genus Tetraplodon in Norway. A taxonomic revision
- FRISVOLL
Beiträge zur Anatomie und Physiologie der Laubmoose
- HABERLANDT
Etude biologique et morphologique sur la dissemination des spores chez les mousses
- PFAEHLER
Spore dispersal in Splachnum ovalum Hedw
- WALSH
Fly-mediated spore dispersal in Splachnum ampullaceum (Musci)
- CAMERON
Der Spaltöffnungsapparat von Brugmansia und Rafflesia
- CAMMERLOHER
Establishment and population dynamics of Tayloria tenuis in a Pyrenean forest Internalional Symposium on Bryophyte Ecology
- F Lloret
Studies on entomophily in Splachnaceae (Musci). II. Volatile compounds in the hypophysis
- PYYSALO
Untersuchungen iiber die fluchtigen Amine dcr Pfianzen
- STEIN VON KAMIENSKI
On the morphology of the sporophyte of Splachnum luteum
- VAIZEY
Elemental analysis of dung mosses (Splachnaceae) and their substrates
- WEBSTER
Pollination of Rqfflesia princei
- BEAMAN
Studies on the generic concept in the classification of the moss family Splachnaceae
- KOPONEN
Comparative behavior of Pyrellia cyanicohr (Diptcra:Muscidac) on the moss Splachnum ampullaceum and on substrates of nutritional value
- TROILO