Can spatial variation in epiphyte diversity and community structure be predicted from sampling vascular epiphytes alone?

Aim Non‐vascular epiphytes have been largely ignored in studies examining the biotic and abiotic determinants of spatial variation in epiphyte diversity. Our aim was to test whether the spatial patterning of species richness, biomass and community composition across geographic regions, among trees within regions, and among branches within trees is consistent between the vascular and non‐vascular components of the temperate rain forest flora. Location Coastal lowland podocarp‐broadleaved forests on the west coast of the South Island of New Zealand. Methods We collected single samples (30 × 25 cm) from 96 epiphyte assemblages located on the inner branches of 40 northern rata (Metrosideros robusta) trees. For each sample, branch characteristics such as branch height, branch diameter, branch angle, branch aspect, and minimum and maximum epiphyte mat depth were recorded. The biomass for each individual epiphyte species was determined. Results Northern rata was host to a total of 157 species, comprising 32 vascular and 125 non‐vascular species, with liverworts representing 41% of all species. Within epiphyte mats, the average total organic biomass of 3.5 kg m^?2 of branch surface area consisted largely of non‐living biomass and roots. Vascular and non‐vascular epiphytes showed strikingly different spatial patterns in species richness, biomass and composition between sites, among trees within sites, and among branches within trees, which could not be explained by the branch structural characteristics we measured. The two plant groups had no significant association in community composition (r = 0.04, P = 0.08). However, the species richness of vascular plant seedlings was strongly linked to the presence/absence of lichens. Main conclusions Non‐vascular plants contributed substantially to the high species richness and biomass recorded in this study, which was comparable to that of some tropical rain forests. High variability in community composition among epiphyte mats, and very low correlation with any of the environmental factors measured possibly indicate high levels of stochasticity in seed or spore colonization, establishment success or community assembly among branches in these canopy communities. Although we found some evidence that vascular plant seedling establishment was linked to the presence of lichens and the biomass of non‐living components in the epiphyte mats, there was no correlation in the spatial patterning or determinants of species richness between non‐vascular and vascular plants. Consequently, variation in total epiphyte biodiversity could not be predicted from the measurement of vascular plant diversity alone, which highlights the crucial importance of sampling non‐vascular plants when undertaking epiphyte community studies.     

Feeding preferences of two seagrass grazing monacanthid fishes

Two seagrass grazing fishes, Meuschenia freycineti and Meuschenia trachylepis (Monacanthidae), were offered three choices of Posidonia australis seagrass blades of different epiphyte coverage and leaf age to determine whether these fishes exhibit a preference for epiphyte-covered seagrass blades. Both species removed significantly more biomass of the epiphyte-covered blades than of the two other blade types in multiple-choice tests. This clear preference for epiphyte-covered seagrass blades results in a preferred removal of older blades within the seagrass shoot of P. australis .     

Nutrient impacts on epifaunal density and species composition in a subtropical seagrass bed

The capacity of epifauna to control algal proliferation following nutrient input depends on responses of both grazers and upper trophic level consumers to enrichment. We examined the responses of Thalassia testudinum (turtle grass) epifaunal assemblages to nutrient enrichment at two sites in Florida Bay with varying levels of phosphorus limitation. We compared epifaunal density, biomass, and species diversity in 2 m² plots that had either ambient nutrient concentrations or had been enriched with nitrogen and phosphorus for 6 months. At the severely P-limited site, total epifaunal density and biomass were two times higher in enriched than in unenriched plots. Caridean shrimp, grazing isopods, and gammarid amphipods accounted for much of the increase in density; brachyuran crabs, primary predatory fish, and detritivorous sea cucumbers accounted for most of the increase in biomass. At the less P-limited site, total epifaunal density and biomass were not affected by nutrient addition, although there were more caridean shrimp and higher brachyuran crab and pink shrimp biomass in enriched plots. At both sites, some variation in epifaunal density and biomass was explained by features of the macrophyte canopy, such as T. testudinum and Halodule wrightii percent cover, suggesting that enrichment may change the refuge value of the macrophyte canopy for epifauna. Additional variation in epifaunal density and biomass was explained by epiphyte pigment concentrations, suggesting that enrichment may change the microalgal food resources that support grazing epifauna. Increased epifaunal density in enriched plots suggests that grazers may be able to control epiphytic algal proliferation following moderate nutrient input to Florida Bay. Electronic supplementary material Electronic supplementary material is available for this article at and accessible for authorised users.     

Survival and Growth of Juvenile Bromeliads in Coffee Plantations and Forests in Central Veracruz,Mexico

Shade coffee plantations can be important refuges for epiphytes, but are not suitable for all species. To test if the performance of early life stages, often the most sensitive phase, is responsible for the species’ ability to colonize coffee plantations, we compared growth and mortality rates of three epiphytic bromeliad species that differ in their ability to colonize secondary arboreal vegetation by transplanting juveniles to trees in forests, and shade trees in old and young coffee plantations in Central Veracruz, Mexico. Growth rates of Tillandsia viridiflora, generally restricted to forests, and Tillandsia juncea, an early colonizer, were related to the pattern of the species occurrence among habitats with growth rates of T. viridiflora being generally higher in forests and growth of T. juncea higher in coffee plantations. Performance of the third species, Tillandsia heterophylla, which is intermediate in habitat preference, was not clearly related to habitat. No difference in growth rates was found between plants transplanted in wet or dry season. In general, mortality in transplanted bromeliads was relatively low (mostly < 5% per month). In coffee plantations herbivory had a severe effect during part of the wet season, when mortality in young coffee plantations reached between 15 and 24 percent per month. Given the substantial contribution of herbivory to the mortality of juvenile plants and the significant differences between habitats, herbivory may be co‐limiting the colonization of young coffee plantations by some epiphytic bromeliads.     

The Canopy Starts at 0.5 m: Predatory Mites (Acari: Mesostigmata) Differ between Rain Forest Floor Soil and Suspended Soil at any Height

Suspended soils in forest canopies are thought to harbor a substantial fraction of canopy biomass and many arboreal specialists, but do forest floor generalist predators with high vagility also use this habitat? We tested the hypothesis of no difference between forest floor and suspended-soil predatory mite faunas (Acari: Mesostigmata) in an Australian rain forest. Our results show that instead of being habitat generalists, many predatory mites partition soil into two main strata: soil suspended aboveground irrespective of height (0.5–20 m) and soil on the ground. Of 53 species of Mesostigmata in suspended soil, 53 percent (28 species) were absent from or rarely found on the ground. This increased to 60 percent (15/25 species) if only common species are considered. Among these 15 ‘suspended-soil specialists’, all but the three least abundant were found throughout the arboreal strata. Moreover, ten species also occurred in litter accumulated on the surface of decaying logs or boulders close to the forest floor. Thus, although the arboreal predatory mite fauna is distinct from that on the forest floor, it is not restricted to the high canopy: even slightly elevated substrate appears acceptable as habitat for these suspended-soil specialists. Our data suggest that a substantial portion of a rain forest's soil and litter fauna is held above the forest floor.     

An ecological perspective of allelochemical interference in land–water interface communities

Allelochemical interactions among aquatic macrophytes and between macrophytes and attached microbial assemblages (epiphyton) influence a number of ecological processes. The ecological importance of these interactions, however, is poorly understood; we hypothesize that paucity has resulted, in part, from (1) a narrow focus on exploration for herbicidal plant products from aquatic macrophytes, (2) the difficulties in distinguishing resource competition from allelopathic interference, and (3) a predominance of approaching aquatic allelopathy from a terrestrial perspective. Based upon recent thorough investigations of allelopathy among aquatic vascular plants, chemical compounds that influence competitive interactions among littoral organisms are amphiphilic compounds that tend to remain near the producing organism (e.g., polyphenolic compounds and volatile fatty acids). Production of these compounds may be influenced by relative availability of nutrients (particularly phosphorus and nitrogen), inorganic carbon, and light. Macrophyte strategies of clonal reproduction, in an effort to persist in these highly productive and competitive habitats, have contributed to reduced reliance upon sexual reproduction that is correlated with allelopathic autotoxicity among several dominant wetland plant species. Although few studies document the importance of allelochemical interactions in the wetland and littoral zones of aquatic ecosystems, abundant evidence supports the potential for significant effects on competition and community structure; effects of altered nutrient ratios and availability on plant chemical composition; and resultant effects on trophic interactions, particularly suppression of herbivory, competitive attached algae and cyanobacteria, and heterotrophic utilization of organic matter by bacteria and fungi.     

Effect of epiphytes on the extent of necrotic injuries of resistant and susceptible poplar clones infected with Dothichiza populea.

Poplar cuttings of a resistant clone, Populus ‘Grandis’, and susceptible clones, Populus nigra ‘Italica’ and Populus ‘Robusta’, were infected with the pathogenic fungus Dothichiza populea alone, or with the pathogen and one of five strains of epiphytes antagonistic towards it (in vitro), isolated from poplar bark. The extent of injury was examined for 28 days after infection by determining the length of necrotic patches and their area as expressed in per cent of the total area of a cutting or the area of necrotic injuries caused by the pathogen alone.

All the poplar cuttings of both the resistant and susceptible clones became diseased when infected with the pathogen alone. Surprisingly enough, however, the least affected clone was the susceptible P. ‘Robusta’, in which necrotic injuries covered 28% of the total area, as against 40% and 70% in the resistant P. ‘Grandis’ and the susceptible P. nigra ‘Italica’, respectively.

When the cuttings were infected simultaneously with Dothichiza populea and its antagonistic epiphytes, the diseased area in the resistant clone diminished by as much as two-thirds, and in the susceptible P. nigra ‘Italica’, by one-third in comparison with the area affected by the pathogen alone. In turn, in the susceptible P. ‘Robusta’ the introduction of three out of five epiphytes stimulated the growth of the pathogenic fungus producing on average a double increase in the necrotic area. The differences in the response of the pathogen to the presence of epiphytes recorded in the susceptible clones indicate a marked influence of the plant on the nature of interactions between its epiphytic microflora and the pathogen.     

Niche Differentiation in Tank and Atmospheric Epiphytic Bromeliads of a Seasonally Dry Forest

Factors influencing the niche differentiation of epiphytes have been determined for the epiphytic bromeliads that coexist in the seasonally dry forest of Chamela, Mexico. Over 40 percent of the bromeliad epiphytes were distributed in only 5 percent of the trees. The occurrence of compound leaves in host trees was highly correlated with abundance of epiphytes, as these allow scattered light to penetrate throughout the canopy. The effect of leaf type overrides the effect of bark type, the main factor determining seedling establishment in moist forests. Eight species had the atmospheric life form, while only two species had tanks, formed by overlapping leaf bases and associated to a lower drought tolerance. Distribution in the canopy is counter to that observed in moist forests, since tank species occur in the upper canopy. Tank life forms showed most annual carbon gain during the rainy season, when the newly leafed out trees provide shade to the lower canopy. Atmospheric species had photosynthetic activity for longer into the dry period, possibly supported by dew and fog events. Leaf angles, orientation, trichome, and stomata densities are discussed in relation to water and light use among the species with contrasting ecological strategies.     

Composition and spatio temporal variability of the epiphytic macroalgal assemblage of Fucus vesiculosus Linnaeus at Clare Island, Mayo, western Ireland

The composition and the patterns of spatial and temporal variability of the epiphytic assemblages of Fucus vesiculosus of Clare Island, on the western coast of Ireland, were investigated for an annual cycle. Specimens of Fucus were collected on seven sampling dates from three sites of the island (Portlea, Kinnacorra and Portnakilly). Data of cover of the most common epiphytic species were collected and analysed by multivariate and univariate techniques. Elachista fucicola, Polysiphonia lanosa, Porphyra umbilicalis, Spongonema tomentosum and Ulva compressa were the most common species. Spatial and temporal variation was detected both for the whole assemblage and for the most abundant species. In general, the assemblage was quantitatively more abundant and diverse in spring-summer than in autumn-winter. In spring-summer, there was a clear differentiation among the assemblage of Portnakilly and the assemblages of the other two sites. Individual species were also generally more abundant in spring-summer and their distribution at the three sites was often not consistent in time; P. lanosa was the only epiphyte for which a consistent effect of site was found. Spatial variation on a scale of meters to tens of meters was the most striking pattern of distribution of the epiphytic assemblage; significant effects related to this spatial scale were detected both for the whole assemblage and for individual species. Phenological patterns of the epiphytic species determining availability of propagules and limited dispersal, leading to aggregated patterns of distribution, are considered the main factors responsible of such patchiness. The importance of the incorporation of small spatial scales in sampling designs analysing the distributional patterns of epiphytic assemblages is discussed.     

Water vapour isotopic exchange by epiphytic bromeliads in tropical dry forests reflects niche differentiation and climatic signals

The 18O signals in leaf water (delta18O(lw)) and organic material were dominated by atmospheric water vapour 18O signals (delta18O(vap)) in tank and atmospheric life forms of epiphytic bromeliads with crassulacean acid metabolism (CAM), from a seasonally dry forest in Mexico. Under field conditions, the mean delta18O(lw) for all species was constant during the course of the day and systematically increased from wet to dry seasons (from 0 to +6 per thousand), when relative water content (RWC) diminished from 70 to 30%. In the greenhouse, progressive enrichment from base to leaf tip was observed at low night-time humidity; under high humidity, the leaf tip equilibrated faster with delta18O(vap) than the other leaf sections. Laboratory manipulations using an isotopically depleted water source showed that delta18O(vap) was more rapidly incorporated than liquid water. Our data were consistent with a Craig-Gordon (C-G) model as modified by Helliker and Griffiths predicting that the influx and exchange of delta18O(vap) control delta18O(lw) in certain epiphytic life forms, despite progressive tissue water loss. We use delta18O(lw) signals to define water-use strategies for the coexisting species which are consistent with habitat preference under natural conditions and life form. Bulk organic matter (delta18O(org)) is used to predict the deltaO18(vap) signal at the time of leaf expansion.