Frond development and phenology of Thelypteris limbosperma, Athyrium distentifolium, and Matteuccia struthiopteris in Western Norway

Odland, A. 1995. Frond development and phenology of Thelypteris limbosperma, Athyrium distentifolium, and Matteuccia struthiopteris in Western Norway. — Nord. J. Bot. 15: 225–236. Copenhagen. ISSN 0107–055X. The pattern of growth and phenology of Thelypteris limbosperma, Athyrium distentifolium, and Matteuccia struthiopteris fronds has been investigated, with particular emphasis on height increments of the sporophytes and the development of fertile fronds. In order to study both interspecific and intraspecific differences, fern stands along altitudinal gradients have been monitored. To investigate the initiation of fertility, fronds representing different developmental stages have been sampled. Climatic data have been collected with a data-logger during the study periods. The ferns show significant differences in growth pattern and phenology. Matteuccia struthiopteris has a growth curve that is best described as monomolecular, characterised by high growth rates during the early developmental stage, while the development of Thelypteris limbosperma and Athyrium distentifolium follow a logistic growth curve. The species need different periods of time to develop their fronds. The main frond elongation does not start before soil temperature has reached 7 C. After that, the growth is mainly controlled by air temperature. Within the interval when mean maximum temperatures are 8–20 C, all three ferns showed increased growth rates with increasing air temperatures, but the increase was greatest in M. struthiopteris. At higher temperatures, the growth rate of M. struthiopteris and T. limbosperma decreased. Athyrium distentifolium is characterised by producing sori at an early developmental stage. Fertile Matteuccia strurhiopteris fronds are developed at a much later developmental stage. Thelypteris limbosperma needs a longer period of time to produce mature fronds than the other species. The investigation indicates that plant growth and development are determined by both internal and external factors. It is concluded that growth rate, phenology, and the ability of ferns to produce mature fronds reveal close similarities with their response to environmental variables, and hence with their broad-scale geographical distributional patterns.     

Arsenic hyperaccumulation in the Chinese brake fern (Pteris vittata) deters grasshopper (Schistocerca americana) herbivory

Brake fern, Pteris vittata, not only tolerates arsenic but also hyperaccumulates it in the frond. The hypothesis that arsenic hyperaccumulation in this fern could function as a defense against insect herbivory was tested. Fronds from control and arsenic-treated ferns were presented to nymphs of the grasshopper Schistocerca americana. Feeding damage was recorded by visual observation and quantification of the fresh weight of frond left uneaten and number of fecal pellets produced over a 2-d period. Grasshopper weight was determined before and after 5 d of feeding. Grasshoppers consumed significantly greater amounts of the frond tissue, produced more fecal pellets and had increased body weight on control plants compared with grasshoppers fed arsenic-treated ferns. Very little or none of the arsenic-treated ferns were consumed indicating feeding deterrence. In a feeding deterrent experiment with lettuce, sodium arsenite at 1.0 mm deterred grasshoppers from feeding whereas 0.1 mm did not. In a choice experiment, grasshoppers preferred to feed on lettuce dipped in water compared with lettuce dipped in 1.0 mm sodium arsenite. Our results show that arsenic hyperaccumulation in brake fern is an elemental defense against grasshopper herbivory.     

Lepidopteran Soral Crypsis on Caribbean Ferns1

Insect‐fern interactions are not as well characterized as insect‐angiosperm interactions. For example, the imitation of unique fern structures by insects has not been observed. On a recent trip to Puerto Rico, we collected ferns that bore small Lepidoptera imitating fern sori. Herbarium specimen searches indicate that these Lepidoptera are restricted to the Caribbean on ferns with highly visible sporangia. A possible selective pressure for the evolution of lepidopteran soral crypsis is wasp parasitism, as evidence of parasitoid wasps was found. However, it is more likely that soral crypsis evolved to avoid visually oriented predators such as birds or lizards.     

Quantitative vegetation-environment relationships in west norwegian tall-fern vegetation

The aims of this paper are to detect floristic variation within different types of tall-fern dominated vegetation and to interpret these patterns in terms of environmental variables. Numerical approaches have been applied to a large and varied vegetational data-set with associated environmental data from stands dominated by Athyrium distentifolium, Thelypteris limbosperma , and Matteuccia struthiopteris in different parts of western Norway. The numerical procedures of two-way indicator species analysis, simple discriminant functions, and canonical correspondence analysis have been used, and the strengths and weaknesses of these as tools in discerning vegetational-environmental relationships are discussed. For each of the 96 quadrats investigated, 17 environmental variables were measured. The investigation shows that some of the observed differences in vegetational composition can be explained in terms of relatively simple soil and climatic variables measured for each quadrat. The ferns appear to be ecologically well separated. T. limbosperma-dominaled stands are mainly characterised by low soil fertility, high January temperature, and high humidity. A. distentifolium-dominated stands are associated with low winter temperatures, and M. struthiopteris-dominated stands have high soil fertility and high summer temperatures.     

Elevational diversity patterns as an example for evolutionary and ecological dynamics in ferns and lycophytes

Evolutionary processes such as adaptation, ecological filtering, and niche conservatism involve the interaction of organisms with their environment and are thus commonly studied along environmental gradients. Elevational gradients have become among the most studied environmental gradients to understand large-scale patterns of species richness and composition because they are highly replicated with different combinations of geographical, environmental and historical factors. We here review the literature on using elevational gradients to understand evolutionary processes in ferns. Some phylogenetic studies of individual fern clades have considered elevation in the analysis or interpretation and postulated that fern diversification is linked to the colonization of mountain habitats. Other studies that have linked elevational community composition and hence ecological filtering with phylogenetic community composition and morphological traits, usually only found limited phylogenetic signal. However, these studies are ultimately only correlational, and there are few actual tests of the evolutionary mechanisms leading to these patterns. We identify a number of challenges for improving our understanding of how evolutionary and ecological processes are linked to elevational richness patterns in ferns: i) limited information on traits and their ecological relevance, ii) uncertainties on the dispersal kernels of ferns and hence the delimitation of regional species pools from which local assemblages are recruited, iii) limited genomic data to identify candidate genes under selection and hence actually document adaptation and selection, and iv) conceptual challenges in developing clear and testable hypotheses to how specific evolutionary processes can be linked to patterns in community composition and species richness.     

Allelopathic effects of interrupted fern on northern red oak seedlings: Amelioration bySuillus luteus L.: Fr.

Summary The allelopathic effects of interrupted fern frond leachates on ectomycorrhizal (inoculated) and nonmycorrhizal (noninoculated) northern red oak (Quercus rubra L.) seedlings were investigated. Container-grown northern red oak was inoculated with vegetative mycelium ofSuillus luteus L. Fr. following acorn germination. Noninoculated control seedlings were also maintained. Seedlings were grown in a glasshouse under full sunlight or shaded (25% of full sunlight) conditions. Leachate or deionized water solutions were applied to seedlings eleven times over a 91-day period to simulate a rainfall induced transfer of allelopathic chemicals from fern fronds to the soil. Fern frond leachates significantly reduced seedling survival, however, inoculated seedlings showed less mortality. Chromium concentrations of pooled lateral root or leaf tissue were comparatively higher in tissues exhibiting greater mortality. Root biomass was reduced by fern fern frond leachate applications. Seedling biomass was not significantly affected by fungal inoculation. Our results confirm previous documentation of the allelopathic potential of ferns, and suggest that ectomycorrhizal fungi may ameliorate allelopathic effects of ferns on northern red oak seedling survival and growth.     

Anatomically preserved Woodwardia virginica (Blechnaceae) and a new filicalean fern from the middle Miocene Yakima Canyon flora of central Washington, USA

Anatomically preserved Woodwardia virginica (Blechnaceae) and a newly recognized onocleoid fern are described from the middle Miocene Yakima Canyon flora of central Washington State, USA. Identification of the W. virginica fossils is based on a combination of vegetative pinnules, rhizome and stipe anatomy, and fertile pinnules with indusiate sori and sporangia like those of extant W. virginica. Fronds are isomorphic. Vegetative pinnae are elongated and pinnatifid, with a secondary vein paralleling the midvein. Secondary veins of the pinnule lobe anastomose to form primary areoles and are either simple or dichotomize toward the margin. Rhizomes have a simple dictyostele with 3-5 cauline vascular bundles and often a sclerotic hypodermis. Leaf traces contain two large adaxial vascular bundles that occur laterally and adaxially, flanking an arc of 4-6 smaller bundles. Fertile pinnules have linear sori that are somewhat embedded in the laminae and are enclosed by a thin indusium. Leptosporangia display a vertical annulus and an elongated stalk. A second fern, Wessiea yakimaensis gen. et sp. nov., is represented by anatomically preserved branching rhizomes and attached frond bases that conform to the Onoclea-type pattern of rhizome and frond-base vasculature. Rhizomes have a simple dictyostele of 4-5 cauline meristeles. Leaf divergence is helical, with paired hippocampiform rachial traces. These two ferns occur in the same matrix with specimens of Osmunda wehrii. They demonstrate that filicalean fern assemblages similar to those of extant temperate floras were well established in western North America by the middle Miocene and further emphasize the exceptional species longevity of some homosporous pteridophytes.     

Experimental and Analytical Studies of Pteridophytes: XL. Factors in the Formation and Distribution of Sori in Leptosporanglate Ferns

In further studies of soral morphogenesis in common Britishferns, consideration is given to environmental and other factorswhich may be causally involved. Accounts are given of the normalcourse of sporophyll development in Dryopteris austriaca andof the effects of more or less extensive defoliation on theinception of sori in several ferns. Some anomalous developmentsin sporophylls of Blechnum spicant are described and illustrated.Various general ideas regarding factors involved in the onsetof the reproductive phase are discussed. It is concluded that,on the evidence which is admittedly very incomplete, the onsetof the reproductive phase in the fern sporophyte is determinedby the interaction of a number of factors, rather than by thespecial action of some particular factor. Because the nature,functional sorus is a complex and highly organized structure,its further investigation is considered to be a task of veryconsiderable magnitude.     

The classification of ferns

Intensive morphological studies have been devoted to the more primitive ferns, which represent a small minority of living species, but too little is yet known about the vast majority of other ferns, with the result that recent attempts at a natural classification show considerable differences of treatment.
The problem is complicated by convergent evolution in the characters of almost all parts of a fern plant. Not only similar soral form, but also similar frond form, types of venation, scales, etc. have been developed on different evolutionary lines.
To illustrate the nature of the problem an attempt has been made to state the probable characters of a primitive leptosporangiate fern, and the kinds of ways in which existing ferns have developed from this condition. Evolutionary change in different parts of the plant has proceeded in different ways and to different degrees in the many genera of existing ferns. Primitive characters of one kind or another are shown by a great number of ferns, along with highly advanced characters of other kinds.
Recent schemes of classification are briefly compared, and a summary is given of the author's own scheme, with notes on evolutionary trends in the various groups as he sees them.
Much more information is needed on which to establish a really satisfactory scheme. The present one is put forward in the hope that others will take up the work. With modern facilities for travel, it is to be hoped that more botanists will come to the tropics and see ferns and other too-little-known plants in their native habitats. Morphological study needs to be undertaken with an understanding of the living plant and of its environment.     

Diversity and distribution of ferns in sub-Saharan Africa, Madagascar and some islands of the South Atlantic

Aim This paper reports the diversity and endemism patterns of African ferns, and explores the potential role of diversity refuges and environmental and historical factors in the shaping of these patterns. Material and locations The extant fern taxa occupying Africa south of the Sahara, Madagascar and some islands of the South Atlantic. Methods The number of taxa in each area or operational geographical unit (OGU) was scored, and the correlation between this number and physical and climatic variables analysed by standard pairwise and stepwise multiple regression analysis (SPR and SMR). The effects of biological factors such as dispersal capacity, reproductive biology, genetic features and certain physiological adaptations were evaluated by comparing the number of species in each OGU. Floral affinities among OGUs were analysed using non‐metric multi‐dimensional scaling (NMS) and parsimonic analysis of dispersion (PAD), and compared with β‐turnover and inter‐OGU distances. Results OGU area, elevation and the distance between refuges determined the composition of local floras, but only greater OGU area and the existence of higher maximum elevations increased species richness. The distance between refuges also affected the number of endemic species, especially on islands. The biological features studied only slightly influenced fern distribution. The main climatic predictor of species number was humidity. SPR and SMR revealed three main groups of ferns with different ecological trends. NMS and PAD analyses separated the four areas of highest diversity in Africa, three of which are inhabited by ferns with distinct ecological requirements. The fourth area was Madagascar, which shows an accumulation of endemic and relict diversity that is not easy to explain. Main conclusions The distribution of ferns in Africa has been influenced by refuges. These probably allowed many species to recolonize the neighbouring areas after the extinctions of the Pleistocene. Three major components were detected in the African flora: Guinea‐Congolian thermophilous, cold‐tolerant Afro‐montane, and Southern drought‐tolerant elements. These are related to the three main refuge areas, i.e. the Gulf of Guinea area, the eastern tropical region, and the Cape region. Endemicity in ferns was found to be lower than that of seed plants due to the higher dispersability of fern spores. The distance between OGUs seems to be the main predictor of the number of endemic fern species these areas contain.     

Variation in ploidy level and phenology can result in large and unexpected differences in demography and climatic sensitivity between closely related ferns

? Premise of the study: Current environmental changes may affect the dynamics and viability of plant populations. This environmental sensitivity may differ between species of different ploidy level because polyploidization can influence life history traits. We compared the demography and climatic sensitivity of two closely related ferns: the tetraploid Polystichum aculeatum and one of its diploid parents, Polystichum setiferum. ? Methods: Matrix models were used to assess the effects of life history variation on population dynamics under varying winter conditions. We analyzed the contributions of all key aspects of the fern life cycle to population growth. Our study is the first to also include the gametophyte generation. ? Key results: Projected population growth rate (λ) was much higher for the tetraploid P. aculeatum (1.516) than for P. setiferum (1.071) under normal winter conditions. During a year with harsh winter conditions, population growth of P. aculeatum was strongly reduced. This finding contradicts our expectation that the winter-hardy fronds of this species would allow high survival of harsh winters. Differences in λ between species and between years with different winter conditions were mostly caused by variation in gametophyte-related recruitment rates, a finding that shows the importance of including gametophytes in fern demographic studies. ? Conclusions: Our results indicate that populations of closely related ferns can show large differences in population performance, mainly related to recruitment rates and frond phenology, and that these differences may depend greatly on climatic conditions. Our findings provide a first indication that (allo)polyploidization in ferns can have a significant effect on population dynamics.     

Spatial Scale or Amplitude of Predictors as Determinants of the Relative Importance of Environmental Factors to Plant Community Structure

The literature on tropical rain forest plant‐community relationships with environmental factors usually does not recognize that the relative importance of environmental factors recorded in each study might be due to their amplitude of variation within sites. Geographic scale, however, is recognized as an important modulator of this relative importance. To disentangle the effects of scale and environmental amplitude, ferns and trees in two landscapes of the same size (each 25 km²) with different soil‐fertility amplitudes but similar soil‐texture range were sampled in central Amazonia. We found that major determinants of community structure were the same for ferns and trees. Texture was the main predictor of community structure in the site with homogeneous soil fertility, while availability of exchangeable cations was the main predictor in the site with a wider fertility range. When both sites were analyzed together, soil fertility was the main predictor of community structure and soil texture segregated floristic subgroups within certain ranges of the soil‐fertility gradient. We conclude that: (1) floristic patterns for trees and ferns are congruent; (2) floristic variation depends on the amplitude of the studied gradients, more than on geographical scale; (3) limiting factors are not necessarily the most important predictors of compositional patterns; and (4) communities are structured hierarchically. Therefore, landscape structure (meaning which combinations of environmental factors, their amplitude of variation and which part of the gradient is found within the landscape) affect our perception of the relative importance that environmental factors will have as predictors of species composition.     

Soral crypsis: protective mimicry of a coccid on an Indian fern

Herbivory with crypsis is not well documented in ferns. The present record of cryptic coloration of coccid Saissetia filicum Boisduval (Homoptera: Coccidae) to the sori of a fern species Asplenium nidus L. (Aspleniaceae) is unique. Predatory beetles (Jauravia sp., Coleoptera: Coccinellidae) that feed on the coccids, are suggested to be selective pressure for the development of the present homopteran soral crypsis. A higher rate of effective predation is noticed in the vegetative leaves than the fertile leaves. Aggressive ants were found harvesting honeydew secretions from the coccids and defending the trophobionts as well as the host fern from their natural enemies. In addition, a possible three-way mutualistic relationship among the coccids, its host fern and the tending ant is suggested. Differential numbers of coccids on vegetative and fertile leaves is correlated with their phenol content and degree of predation by beetles. Such coloration mimicry by the coccids may enable them to obtain the necessary blend of sorus of the host fern needed to evade beetle detection and attack.     

Stasis and convergence characterize morphological evolution in eupolypod II ferns

Background and Aims

Patterns of morphological evolution at levels above family rank remain underexplored in the ferns. The present study seeks to address this gap through analysis of 79 morphological characters for 81 taxa, including representatives of all ten families of eupolypod II ferns. Recent molecular phylogenetic studies demonstrate that the evolution of the large eupolypod II clade (which includes nearly one-third of extant fern species) features unexpected patterns. The traditional ‘athyrioid’ ferns are scattered across the phylogeny despite their apparent morphological cohesiveness, and mixed among these seemingly conservative taxa are morphologically dissimilar groups that lack any obvious features uniting them with their relatives. Maximum-likelihood and maximum-parsimony character optimizations are used to determine characters that unite the seemingly disparate groups, and to test whether the polyphyly of the traditional athyrioid ferns is due to evolutionary stasis (symplesiomorphy) or convergent evolution. The major events in eupolypod II character evolution are reviewed, and character and character state concepts are reappraised, as a basis for further inquiries into fern morphology.

Methods

Characters were scored from the literature, live plants and herbarium specimens, and optimized using maximum-parsimony and maximum-likelihood, onto a highly supported topology derived from maximum-likelihood and Bayesian analysis of molecular data. Phylogenetic signal of characters were tested for using randomization methods and fitdiscrete.

Key Results

The majority of character state changes within the eupolypod II phylogeny occur at the family level or above. Relative branch lengths for the morphological data resemble those from molecular data and fit an ancient rapid radiation model (long branches subtended by very short backbone internodes), with few characters uniting the morphologically disparate clades. The traditional athyrioid ferns were circumscribed based upon a combination of symplesiomorphic and homoplastic characters. Petiole vasculature consisting of two bundles is ancestral for eupolypods II and a synapomorphy for eupolypods II under deltran optimization. Sori restricted to one side of the vein defines the recently recognized clade comprising Rhachidosoraceae through Aspleniaceae, and sori present on both sides of the vein is a synapomorphy for the Athyriaceae sensu stricto. The results indicate that a chromosome base number of x =41 is synapomorphic for all eupolypods, a clade that includes over two-thirds of extant fern species.

Conclusions

The integrated approach synthesizes morphological studies with current phylogenetic hypotheses and provides explicit statements of character evolution in the eupolypod II fern families. Strong character support is found for previously recognized clades, whereas few characters support previously unrecognized clades. Sorus position appears to be less complicated than previously hypothesized, and linear sori restricted to one side of the vein support the clade comprising Aspleniaceae, Diplaziopsidaceae, Hemidictyaceae and Rachidosoraceae – a lineage only recently identified. Despite x =41 being a frequent number among extant species, to our knowledge it has not previously been demonstrated as the ancestral state. This is the first synapomorphy proposed for the eupolypod clade, a lineage comprising 67 % of extant fern species. This study provides some of the first hypotheses of character evolution at the family level and above in light of recent phylogenetic results, and promotes further study in an area that remains open for original observation.     

COMPLEX PALEOZOIC FILICALES IN THE EVOLUTIONARY RADIATION OF FERNS

Filicalean frond segments bearing indusiate sori with gradate development have been discovered in Pennsylvanian deposits of eastern North America. The specimens combine characters of Schizaeaceae, Gleicheniaceae, Cyatheaceae and Hymenophyllaceae, and together with data from other sources, indicate that the Filicales were in the midst of an evolutionary radiation during the late Paleozoic. Evidence from fossil and extant species prompts the hypothesis that filicalean ferns originated near the base of the Carboniferous, and have undergone three major evolutionary radiations.     

Fern richness,tree species surrogacy,and fragment complementarity in a Mexican tropical montane cloud forest

We related pteridophytes versus tree species composition to identify surrogate measures of diversity, and complementarity of seven cloud forest fragments. Forest structure, and fern and tree composition were determined in 70 (2 × 50 m) transects. Fern density (10,150–25,080 individuals/ha) differed among sites. We recorded 83 fern species in the transects. Nonparametric richness estimators indicated that more sampling effort was needed to complete fern inventories (14 more species). However, ferns recorded outside of the transects increased richness to 103 species (six more species than predicted). Twenty-eight species were unique and rare due to special habitat requirements (Diplazium expansum, Hymenophyllum hirsutum, Melpomene leptostoma, Terpsichore asplenifolia), or were at a geographical distribution edge (Diplazium plantaginifolium, Lycopodium thyoides, Pecluma consimilis, Polypodium puberulum). Correlations between fern richness and tree richness and density were not significant, but were significant between fern richness and fern density, between epiphytic fern density and tree richness and density. Tree richness is not a good surrogate for fern diversity. Only three species were recorded in all fragments (Polypodium lepidotrichum, P. longepinnulatum, P. plebeium); thus fragments pteridophytes compositions are highly complementary, but more similar for ferns than for trees. A regional conservation approach which includes many small reserves needs to focus supplementarity on patterns of tree and fern species richness.     

Ashicaulis woolfei n. sp.: additional evidence for the antiquity of osmundaceous ferns from the Triassic of Antarctica

Numerous small fern trunks and dispersed osmundaceous frond fragments occur within a Middle Triassic silicified peat near Fremouw Peak in the Transantarctic Mountains of Antarctica. These specimens form the basis of a new species of osmundaceous ferns that further helps to characterize the early Mesozoic vegetation of high latitude Gondwana. Ashicaulis woolfei n. sp. consists of small, upright trunks with a persistent armor of frond bases, adventitious roots, and vegetative frond parts. In cross section the trunks are ～2.5 cm in diameter and include up to 45 frond bases. Stems range from 5 to 8 mm in diameter with a xylem cylinder of 8-9 xylem segments separated by leaf gaps. Phyllotaxy is variable, approaching 2/5 or 3/8, with 10-12 frond traces in the cortex. Stipes have parenchymatous, stipular wings that are usually devoid of sclerenchyma; fronds are pinnate with alternate-subopposite pinnatifid pinnules. Although the absence of fertile pinnules and sporangia precludes assigning the fossils to a living genus, this species demonstrates that ferns with stelar architecture and histology similar to Osmunda subgenus Osmundastrum (Osmundaceae) were present in the Southern Hemisphere by the mid-Triassic.     

Seasonal and interspecific variation in the biochemical composition of some British fern species and their effects on Spodoptera littoralis larvae

Fern-feeding insects in Britain are mainly found on mature fronds in late summer. Six fern species (Dtyopteris filix-mas, D. dilatata, D. borreri, Phyllitis scolopendrium, Polyslkhum setiferum and Polypodium vulgare) were analysed for fibre, lignin, cellulose, tannins, cyanogenesis and thiaminase activity in an attempt to determine the biochemical basis for this seasonal pattern of attack. A bioassay was also carried out, using frond material incorporated in the diet of Spodoptera littoralis, to determine the effects of seasonal changes in fern toxins on a non-adapted, insect herbivore. The young fronds of all six species had a high protein content and low fibre-lignocellulose; protein levels decreased and cell wall materials increased as the fronds matured. Tannin concentrations and thiaminase activity showed a less consistent seasonal pattern. Tannins were present in the highest concentration in the young fronds of D. borreri, Polystichum and Polypodium and decreased with season; D. dilatata, Polystichum and Polypodium showed a similar pattern of thiaminase activity. The other species showed no strong seasonality in the levels of either tannins or thiaminase. No cyanogenic activity was detected in any of the fern species. Frond material from all six species decreased survivorship and growth rates of Spodoptera larvae. Larval performance and growth on diets containing fern material from young immature fronds was generally poorer than on diets containing material from mature fronds, collected later in the season. Neither tannin concentrations nor thiaminase activity levels closely corresponded to the observed interspecific and seasonal patterns of larval development and mortality, and other toxins must be involved. It is concluded that ferns are highly toxic to non-adapted herbivorous insects but the effects on adapted species are unknown. The restricted seasonal occurrence of a small number of insect species exhibiting specialized feeding habits suggests, however, that biochemical properties of the ferns determine this pattern of attack and limit the number of species which are able to exploit ferns as food resources.     

Environmental and neighbourhood effects on tree fern distributions in a neotropical lowland rain forest

Questions: To what extent are the distributions of tropical rain forest tree ferns (Cyatheaceae) related to environmental variation, and is habitat specialization likely to play a role in their local coexistence? Location: Lowland rain forest at La Selva Biological Station, Costa Rica. Methods: Generalized linear (GLM) and generalized additive (GAM) logistic regression were used to model the incidence of four tree fern species in relation to environmental and neighbourhood variables in 1154 inventory plots regularly distributed across 6 km² of old‐growth forest. Small and large size classes of the two most abundant species were modelled separately to see whether habitat associations change with ontogeny. Results: GLM and GAM model results were similar. All species had significant distributional biases with respect to micro‐habitat. Environmental variables describing soil variation were included in the models most often, followed by topographic and forest structural variables. The distributions of small individuals were more strongly related to environmental variation than those of larger individuals. Significant neighbourhood effects (spatial autocorrelation in intraspecific distributions and non‐random overlaps in the distributions of certain species pairs) were also identified. Overlaps between congeners did not differ from random, but there was a highly significant overlap in the distributions of the two most common species. Conclusions: Our results support the view that habitat specialization is an important determinant of where on the rain forest landscape tree ferns grow, especially for juvenile plants. However, other factors, such as dispersal limitation, may also contribute to their local coexistence.     

Contrasting Predictors of Fern versus Angiosperm Decomposition in a Common Garden

We evaluated differences in the rates and correlates of decomposition among 32 fern and angiosperm litter types collected in Hawai'i. Leptosporangiate ferns were separated into groups based on phylogeny: 'polypod' ferns, a monophyletic clade of ferns that diversified in the Cretaceous, and all other ('non-polypod') ferns that diversified earlier. We measured initial litter chemistry (nutrients and carbon chemistry), and mass loss and nitrogen (N), phosphorus (P), and calcium (Ca) of litter tissue during a 1-yr incubation in a common garden. Nutrient concentrations and carbon (C) chemistry differed significantly among litter types, and litter turnover ( k -values) ranged from 0.29 to 8.31. Decomposition rates were more closely correlated with nutrient concentration than is typically observed. Lignin:N was the best predictor of decomposition across all litter types combined; however, among plant groups different predictors of decomposition were important. Nitrogen and P concentrations best predicted fern decomposition, whereas C chemistry, particularly lignin concentration, was more important for angiosperm (monocot and dicot) decomposition. Among native plants, non-polypod ferns decomposed significantly more slowly than both polypod ferns and angiosperms. Contrary to our hypothesis, fern litter did not decompose more slowly than angiosperm litter overall. Nutrient dynamics in litter were affected by initial litter concentration more than phylogeny; low-nutrient litter immobilized more nutrients than high-nutrient litter. Systematic differences in rates of decomposition, and the importance of nutrients in predicting fern decomposition, imply that changes in species composition within ferns and between ferns and angiosperms could influence the functioning of ecosystems where ferns are important forest components.