Diversity,distribution and host-species associations of epiphytic orchids in Nepal

Associations between epiphytes and their hosts are among the main factors affecting the biodiversity and distribution of epiphytes. While several previous studies explored the association between epiphyte diversity and host characteristics, very little is known about the generality of such associations at larger spatial scales. We aim to explore the associations between diversity and distribution of epiphytic orchids and host characteristics in different localities in Nepal. Epiphytic orchids and their hosts were recorded along the transects in total of 23,539 host individuals. To describe the diversity of orchids in the different localities, a rarefaction function was used. Univariate and multivariate analyses were carried out to explore the associations with locality, host characteristics, and their interactions with locality. In total, we recorded 141 species of orchids growing on 192 host species. The five localities significantly differed in orchid diversity and abundance. The number of orchid individuals per host species significantly increased with increasing number of host individuals. Species richness, abundances, occupancy of orchid species on host species and composition of orchids varied across the localities. Species richness and abundance was significantly higher on hosts in the higher strata and differed between families of host species. Abundance was higher on evergreen hosts. Composition of orchid communities are also associated with host characteristics, such as habit (shrub/tree/climber), bark texture, nature (deciduous/evergreen) and the plant families of host species. This study revealed a high diversity of epiphytic orchids in the localities studied and strong associations between the orchids and their hosts. Future studies looking at the relationships between epiphyte communities and host characteristics need to identify relationships at a wider scale in order to determine whether they are really general rather than site-specific.     

Consumer identity,abundance and nutrient concentration affect epiphyte diversity in an experimental eelgrass system

Conceptual models predict a unimodal effect of consumer abundance on prey diversity with the highest diversity at intermediate consumer abundance (intermediate disturbance hypothesis). Consumer selectivity and prey productivity are assumed to be further important determinants. Preferential grazing on dominant prey species favoured by high nutrient supply is supposed to increase prey diversity, whereas the effect of consumers on prey diversity may be negative under low nutrient conditions (grazer reversal hypothesis). We tested the effect of four common consumers the isopod Idotea baltica, the amphipod Gammarus oceanicus, and the gastropods Littorina littorea and Rissoa membranacea on diversity and composition of epiphytes growing on eelgrass Zostera marina. Consumer density was manipulated (four levels: grazer free control, low, medium, high) based on abundances observed in eelgrass systems. Additionally, we manipulated nutrient supply (three levels) and the presence of Idotea in a factorial experiment. The impact of consumer abundance on epiphyte diversity varied depending on consumer identity and epiphyte evenness was affected rather than species number in this short‐term experiment. Idotea reduced epiphyte diversity (Shannon‐Wiener index H') and Gammarus increased epiphyte diversity. Littorina had no effect at low and medium abundance, but a negative effect in the high density treatment. Only Rissoa supported the conceptual models as it caused the proposed unimodal pattern in epiphyte diversity. The varying species‐specific selectivity of the studied consumers is likely to explain their diverse impact on epiphyte diversity. Nutrients enhanced epiphyte diversity at medium enrichment, whereas higher nutrient supply reduced epiphyte diversity. The effect of Idotea changed from negative at low nutrient concentration to positive at higher nutrient supply, supporting the grazer reversal hypothesis. This study implies that consumer species identity and nutrient concentrations are important in controlling prey diversity and composition. Different consumer selectivity and changes in selectivity with growing consumer abundance and nutrient concentration are the causal factors for this effect.     

Contribution of peeling host for epiphyte abundance in two tropical dry forests in the “El Cielo Biosphere Reserve”, Mexico

Host traits partly determine the abundance and species richness of epiphytes in tropical forests. It has been proposed that older trees with rough bark and evergreens often house more individuals and more epiphytic species than those with thin, smooth, and peeling bark, which harbor few epiphytes. We hypothesize (i) that epiphytes are more abundant and species-rich in the more shaded forest, which is related to bark roughness, and (ii) that epiphytes are distributed in the middle of the host, where microenvironmental conditions are more favorable to survival. We evaluated abundance, species richness, and vertical distribution of epiphytes in two tropical dry forests, according to the deciduousness and basal area of the trees. Moreover, we selected the most abundant epiphytes to test whether their distribution is related to a specific bark type and examine their vertical distribution in two dry forests. We distinguished a high abundance and species richness of epiphytes in the deciduous forest, although basal area and host species richness were higher in the semi-deciduous forest. In both forests, we found a positive relationship between epiphyte abundance and basal area. Higher abundance of epiphytes was related to the predominance of Tillandsia schiedeana, a drought-adapted species, in both forests. Unexpectedly, epiphytes abundantly colonized Bursera simaruba, a host with peeling bark and a very branched crown, where small individuals of T. schiedeana colonized abundantly toward the top of the crown. Our results show the importance of the tropical dry forest, particularly, B. simaruba, in maintaining epiphyte diversity in terms of T. schiedeana colonization.     

Exotic tree monocultures play a limited role in the conservation of Atlantic Forest epiphytes

The Brazilian Atlantic Forest suffered a severe geographic contraction along the last five centuries that reduced drastically most vascular epiphyte populations. Among the range of man-made matrixes, tree monocultures have the potential to contribute positively to the maintenance of the regional epiphyte diversity. Here, we test the similarity in abundance, richness, and species composition between vascular epiphytic communities established in managed monocultures of exotic and native species with natural communities occurring in neighboring native Araucaria Forest patches. In the São Francisco de Paula National Forest (Rio Grande do Sul state, Brazil), we recorded 62 epiphyte species from 300 phorophytes occurring in 12, one-hectare plots of Araucaria Forest and managed plantations of Pinus, Eucalyptus and Araucaria. Species richness, rarefied richness and abundance were significantly higher in Araucaria Forest in comparison to the exotic stands. Species composition was also substantially differentiated as Araucaria Forest patches harbored a greater number of zoochorous species than those of the exotic stands. Additionally, plantations of Araucaria angustifolia, a native species, sustained more individuals and more species than the exotic plantations. Neither tree height nor DBH explained epiphyte richness; however, both phorophyte diversity and stand age together accounted for 92% of the among-site variation in epiphytic species richness. We conclude that substrate heterogeneity in combination with time available for colonization contribute significantly to beta-diversity of epiphytes in Araucaria forests. However, demographic experimental studies are necessary in order to disentangle the role of substrate quality from metapopulation processes, such as dispersal limitation, at both temporal and spatial scales.     

Host trait preferences and distribution of vascular epiphytes in a warm-temperate forest

To illustrate the ecological factors and process leading to the observed diversity patterns of vascular epiphytes, we examined the effect and importance of host tree traits on epiphyte richness and spatial aggregation of epiphytes. The study was conducted in warm-temperate forest in Japan. The recorded host traits were diameter, height, species, habitat topography, and growth rate, and we analyzed the effects and importance of these traits on three species groups: total epiphytic species, epiphytic orchid species, and epiphytic pteridophyte species. Diameter and species of host trees had the greatest influence on epiphytes and their magnitudes were roughly similar in all species groups. Growth rate and topography were less important than host size and species. Growth rate had a negative effect on all three groups, and topography was important for pteridophytes. Epiphyte richness did not exhibit clear spatial aggregation. Our results suggest that size, stability, and quality of the host are equally important in determining epiphyte colonization.     

Temporal variation of amphipod assemblages associated with Sargassum filipendula (Phaeophyta) and its epiphytes in a subtropical shore

The phytal assemblages change in response to variation in biological and environmental conditions. In the present study, we evaluated the temporal variation of amphipod assemblages associated with a Sargassum filipendula bed in a subtropical shore, in relation to variation of the host alga, its epiphytes and local environmental conditions. Samples of S. filipendula with associated amphipods, water temperature, water movement and suspended solids were obtained monthly from June 2000 to May 2001. We recorded 24 species of amphipods associated with S. filipendula. Species richness varied throughout the year, with maximum values in October 2000 and minimum in April 2001. Total amphipod density gradually increased during the sampling period, with the highest value in March 2001. Amphipod diversity and evenness were both positively influenced by epiphyte load and negatively by temperature, with higher values during summer months. Total density and tube-builder density were also positively influenced by temperature, whereas nestler density was influenced by epiphyte load. Individual amphipod species showed significant density fluctuations over the year. The canonical correspondence analysis performed explained 88.2% of the variation, with a strong correlation of water movement, temperature and suspended solids with the first axis, and a strong effect of epiphyte load on both the first and the second axes. The temporal structural variation of the studied algal bed strongly influenced amphipod diversity and assemblage composition, possibly through direct and indirect effects.     

Habitat isolation changes the beta diversity of the vascular epiphyte community in lower montane forest,Veracruz, Mexico

Habitat isolation is one of the most important factors endangering the biodiversity, but little research has been done with vascular epiphytes. In order to understand the effect of isolation on the epiphyte community, we studied epiphyte diversity on three plots in a forest fragment, two riparian forest fragments, and in isolated pastureland trees. We found 118 vascular epiphyte species. On forest plots, both epiphyte richness per tree (S_tree) and species turnover rate within trees (β_tree) registered the highest values, although the lowest S_tree diversity was also found there; additionally inside the forest were host species with clearly different epiphyte community. S_tree and β_tree diversities of riparian fragments behaved similarly to those of the forest. Isolated trees had the second highest S_tree diversity, although their β_tree diversity was the lowest. In the forest plots were both, the highest and lowest expected accumulated richness (α diversity); on riparian fragments it was intermediate, and the second lowest α diversity was registered for isolated trees. Species turnover rate among plots (β) was high and was associated with both, isolation and a distance gradient from permanent water sources. The epiphyte community on isolated trees was clearly different to the other habitats. Results suggest that deforestation eliminated dry areas and specific hosts that were important for the maintenance of epiphyte species richness. In pastureland trees the epiphyte β_tree diversity diminished, suggesting a simplification of the environment for epiphytes and causing a low α diversity.     

Unique distribution of epiphytic Neosiphonia harveyi (Rhodomelaceae,Rhodophyta) along sargassacean hosts

Although epiphytism is generally heaviest on older portions of the host thallus and both variation and abundance of epiphytes decrease with proximity to meristematic apices, Neosiphonia harveyi (Bailey) Kim, Choi, Guiry and Saunders was frequently found on upper parts of sargassacean hosts. This study compared density, thallus length and fertility of epiphytic N. harveyi among different regions of Sargassum patens C. Agardh thalli to reveal a unique distribution pattern of this epiphyte. The majority of epiphytic N. harveyi was observed on the upper part of host thalli and was seldom seen near the basal part. Over a 3‐month period, mean epiphyte density was greater on host apices while thallus length and fertility were greater on subapical portions of host thalli, suggesting the colonization by this epiphyte near the apical part of the host. Distribution patterns of this epiphyte were similar among S. patens growing at different depths. Apical portions of S. patens appear to be more suitable substrata for N. harveyi settlement and colonization compared with other portions of the host thallus, regardless of depth.     

Factors influencing epiphytic bryophyte and lichen species richness at different spatial scales in managed temperate forests

The effect of management related factors on species richness of epiphytic bryophytes and lichens was studied in managed deciduous-coniferous mixed forests in Western-Hungary. At the stand level, the potential explanatory variables were tree species composition, stand structure, microclimate and light conditions, landscape and historical variables; while at tree level host tree species, tree size and light were studied. Species richness of the two epiphyte groups was positively correlated. Both for lichen and bryophyte plot level richness, the composition and diversity of tree species and the abundance of shrub layer were the most influential positive factors. Besides, for bryophytes the presence of large trees, while for lichens amount and heterogeneity of light were important. Tree level richness was mainly determined by host tree species for both groups. For bryophytes oaks, while for lichens oaks and hornbeam turned out the most favourable hosts. Tree size generally increased tree level species richness, except on pine for bryophytes and on hornbeam for lichens. The key variables for epiphytic diversity of the region were directly influenced by recent forest management; historical and landscape variables were not influential. Forest management oriented to the conservation of epiphytes should focus on: (i) the maintenance of tree species diversity in mixed stands; (ii) increment the proportion of deciduous trees (mainly oaks); (iii) conserving large trees within the stands; (iv) providing the presence of shrub and regeneration layer; (v) creating heterogeneous light conditions. For these purposes tree selection and selective cutting management seem more appropriate than shelterwood system.     

Richness, structure and functioning in metazoan parasite communities

Ecosystem functioning, characterized by components such as productivity and stability, has been extensively linked with diversity in recent years, mainly in plant ecology. The aim of our study was thus to quantify general relationships between diversity, community structure and ecosystem functions in metazoan parasite communities. We used data on parasite communities from 15 species of marine fish hosts from coastal Chile. The volumetric abundance (volume of all parasite species per individual host, in mm³) was used as a surrogate for productivity. Species diversity was measured using both species richness and evenness, while community structure was estimated using the co‐occurrence indices V‐ratio, C‐score and a new C‐score_s index standardized for the number of host replicates. After correcting for fish size, 47% of host species show no relationship, 13% show a hump shaped curve and 40% show positive monotonic relationships between productivity and parasite richness across all host individuals in a sample. We obtained a logarithmically decreasing relationship between evenness and productivity for all fish species, and propose a ‘dominance‐resistance’ hypothesis based on immunity to explain this pattern. The stability of the parasite community, measured as the coefficient of variation in productivity among individual hosts, was strongly and positively related to mean species richness across the 15 host species. The C‐score_s index, based on the number of checkerboard units in the host‐parasite presence/absence matrix, increases linearly with mean productivity across the 15 host species, suggesting that parasite communities tend to be more structured when they are more productive. This is the likely reason why linear relationships between richness and productivity were not observed consistently in all fish species. Parasite communities provide some clear patterns for the diversity–ecosystem functioning debate in ecology, although other factors, such as the history of community assembly, may also influence these patterns.     

Community structure of amphipods on shallow <Emphasis Type="Italic">Posidonia oceanica</Emphasis> meadows off Tunisian coasts

The structure, diversity and spatial distribution of the amphipod fauna associated with Posidonia oceanica meadows were studied along the Tunisian coasts in 2007. Samples were collected in nine different meadows at 2 m depth. A total of 44 species belonging to 12 families were collected. The most common species in terms of abundance were Ampithoe helleri, Hyale camptonyx and Ericthonius punctatus. The highest values of abundance and species richness and the lowest values of diversity and equitability were found in meadows with high epiphyte biomass. Multivariate analyses of data indicated that epiphyte biomass and geographical position were major determinants of the distribution and composition of amphipod assemblages along Tunisian coasts. The presence of two lessepsian amphipod species in one of the southern Posidonia oceanica meadows modified the structure of assemblage.     

Host abundance,durability, basidiome form and phylogenetic isolation determine fungivore species richness

Species with close associations to a specific host species, such as parasites and phytophages, make immense contributions to biodiversity. Hence, factors determining the variation in species richness among hosts are a main focus of ecological research. Investigations of determining factors of fungivorous species among host species are still scarce. Based on ecological patterns of parasites and phytophages, we hypothesized that the species richness of tree‐fungus beetles of the family Ciidae (Coleoptera) would increase with increasing basidiome size, niche diversity of the growth form, durability, increasing abundance and decreasing phylogenetic isolation of the host fungus. Our generalized least‐squares model, controlled by host phylogeny, revealed that Ciidae species richness increases with host abundance, but decreases with host phylogenetic isolation. In contrast with our prediction, Ciidae species richness was higher in annual basidiomes than in perennials. Pileate basidiomes revealed higher species richness than resupinate and stipitate basidiomes, which may be interpreted as being a result of their higher host niche diversity. The importance of host abundance, measured on the landscape scale, corroborates that fungivore species richness among macrofungal hosts is determined by factors similar to those that determine parasite and phytophage species richness among their hosts. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 699–708.     

Relationships between arthropod richness,evenness, and diversity are altered by complementarity among plant genotypes

Biodiversity is quantified via richness (e.g., the number of species), evenness (the relative abundance distribution of those species), or proportional diversity (a combination of richness and evenness, such as the Shannon index, H′). While empirical studies show no consistent relationship between these aspects of biodiversity within communities, the mechanisms leading to inconsistent relationships have received little attention. Here, using common evening primrose (Oenothera biennis) and its associated arthropod community, we show that relationships between arthropod richness, evenness, and proportional diversity are altered by plant genotypic richness. Arthropod richness increased with O. biennis genotypic richness due to an abundance-driven accumulation of species in response to greater plant biomass. Arthropod evenness and proportional diversity decreased with plant genotypic richness due to a nonadditive increase in abundance of a dominant arthropod, the generalist florivore/omnivore Plagiognathas politus (Miridae). The greater quantity of flowers and buds produced in polycultures—which resulted from positive complementarity among O. biennis genotypes—increased the abundance of this dominant insect. Using choice bioassays, we show that floral quality did not change in plant genotypic mixtures. These results elucidate mechanisms for how plant genotypic richness can modify relationships between arthropod richness, evenness, and proportional diversity. More broadly, our results suggest that trophic interactions may be a previously underappreciated factor controlling relationships between these different aspects of biodiversity.     

Biodiversity of vespid wasps in spatial and temporal dimensions in northern Zanjan Province of Iran

Vespidae is one of the major families of Hymenoptera with more than 115000 described species. In the present paper, Simpson and Shannon-Wiener diversity indices, Simpson,Camargo, Smith and Wilson, and modified Nee evenness indices, and richness index based on rarefaction method were adopted to study the biodiversity of vespid wasps in spatial and temporal dimensions in northern Zanjan Province of Iran. In spatial dimension, Zanjan and Gilvan showed the highest and the lowest species diversity, evenness, and richness, respectively; while in tem-poral dimension, 16 July-6 August showed the highest species diversity and richness, and 18 May -27 May and 23 August -2 September showed the highest and the lowest species evenness,respectively. Significant differences in species abundance between localities of study area were observed by Kruskal-Wallis nonparametric test.     

Neotropical flowering epiphyte diversity: local composition and geographic affinities

Worldwide, the highest diversity of vascular epiphytic plants resides in the Neotropics. The general pattern of taxonomic composition of the neotropical epiphytic flora has been described, but information regarding the magnitude and geographic distribution of species richness is lacking. In this paper, we carried out a regional scale analysis in order to provide an overview of the richness, composition and geographic affinities among several neotropical epiphyte floras. Our database comprised 7,524 flowering epiphyte species (48 % of the estimated total of neotropical epiphytes) from eight representative localities with politically-defined boundaries. The epiphyte quotient (e.g., percentage of epiphyte species in the total flora) per locality ranged from 5.2–27.7 % (mean: 17.5 %) of the flowering plants. Ecuador represented the most species diverse locality (4,247 spp.), followed by Costa Rica (2,611 spp.). At the family level, the epiphyte composition among localities was roughly homogeneous and dominated by emblematic epiphytic groups: Orchidaceae, Bromeliaceae, and Araceae. However, the generic composition of the epiphytic floras was more heterogeneous. Ecuador and Cuba showed the highest epiphyte endemism (37 and 26 %, respectively), and in some cases vascular epiphytes represented 30 % or more of the total endemic flowering plants at a particular locality. From the available information, four main regional epiphyte floras were identified: the northwestern Andean region, the northern and southern Mesoamerican regions; Cuba as a representative of the Caribbean region remained as a separated group, though weakly related to the Mesoamerican groups. This study identified important geographic localities as targets for the conservation of neotropical epiphyte diversity.     

Do cloud forest tree species differ in their suitability as a substrate for epiphytic bromeliads?

There is evidence for the existence of varying degrees of host preference in vascular epiphytes; certain tree species can be positively, neutrally, or negatively associated with epiphytes. The objective of this study was to evaluate whether tree species of the cloud forest differ in their suitability as a substrate for epiphytic bromeliads. To evaluate the association between epiphytic bromeliad cover and host tree species, we sampled 62 plots (each of 200 m²) in four cloud forest fragments in Veracruz, Mexico. For all trees ≥10 cm in diameter at breast height (DBH), we recorded species name, DBH, and percentage cover of bromeliads in categories of tree coverage. In total, 587 trees belonging to 52 species were recorded. All of the 10 tree species used to assess differences in epiphyte cover (each with a minimum of nine individuals) supported bromeliads, but mean bromeliad cover differed significantly among the tree species. The tree species that concentrated the highest bromeliad cover were Quercus sartorii (29.86%) and Liquidambar styraciflua (21.72%). Our results indicate that, while none of the tree species analyzed was a limiting host for epiphytic bromeliads in general, varying levels of bromeliad cover occur depending on the host species in tropical montane cloud forest fragments suggesting that certain tree species are better hosts than others. The implications for conservation efforts of differential tree species suitability as epiphyte hosts are discussed.     

Epiphytic (including hemiepiphytes) diversity in three timber species in the southwestern Amazon, Brazil

Forestry managers have been searching for ways to reduce the impacts of logging on Amazonian biodiversity, but some basic factors are still not considered in native forestry operations, among them the diversity of epiphytes associated with the logged trees. Our goals in this study were to determine the floristic composition, quantify the species richness, and characterize the species diversity of the vascular epiphytic community present in three timber tree species in Acre State, Brazil. We collected and identified all epiphytes in 30 randomly selected trees ≥35 cm DBH of each of three important timber species, Tabebuia serratifolia, Manilkara inundata and Couratari macrosperma. We also documented the epiphyte diversity in 120 randomly selected trees ≥35 cm DBH of 56 other species to determine whether the three timber species have different epiphyte diversity than the tree community at large. The epiphyte samples in the three timber species showed 77 species, 13 of which were new records for the flora of Acre state. The epiphyte community in the randomly selected trees presented a total of 56 species. The timber species phorophytes hosted on average three times more epiphyte species per tree than the other 120 randomly selected trees. These results show that a substantial portion of local floristic richness can be lost during logging activity due if not properly managed by rescuing epiphytes after felling the trees. Although these epiphytes could contribute positively to forestry sustainability due to their ornamental value, increasing the economic yield per hectare, there are no local initiatives for economic use of epiphytes.     

Upstream effects of a reservoir on fish assemblages 45 years following impoundment

Fish assemblage structure, rarefied species richness, species diversity and evenness of assemblages upstream of a reservoir in Oklahoma, U.S.A., were compared pre and post‐impoundment as well as in contemporary collections from streams above and below the reservoir. There were significant shifts in assemblage structure between historical and contemporary collections above the reservoir but not between contemporary assemblages above and below the impoundment. Indicator species analysis revealed that the sand shiner Notropis stramineus and fathead minnow Pimephales promelas have declined, whereas largemouth bass Micropterus salmoides and western mosquitofish Gambusia affinis have increased in relative abundance in assemblages upstream of the impoundment. Species richness was lower in contemporary assemblages compared with historical assemblages. Furthermore, contemporary assemblages below the dam had lower species richness, diversity and evenness compared with contemporary collections above the dam. These results highlight the spatial and temporal extent of reservoirs altering fish assemblages upstream of impoundments.     

Effects of plant species richness and composition on epifaunal colonization in brackish water angiosperm communities

Studies conducted in terrestrial ecosystems have shown that increasing plant diversity enhances ecosystem processes such as primary production. In marine systems, knowledge of how plant diversity influences ecosystem processes and higher trophic levels is still limited. To examine how plant richness and composition influence recruitment and colonization processes, defaunated eelgrass (Zostera marina), sago pondweed (Potamogeton pectinatus) and perfoliate pondweed (Potamogeton perfoliatus) were planted on an unvegetated, sandy bottom in all possible combinations in July and August. Both field experiments lasted 1 week. Our results showed that in these plant assemblages plant richness had a negative effect on faunal diversity (H′) and evenness (J′), while total abundance showed strong temporal variation with weak, positive relationships with plant species richness in both August and July. Plant species composition had strong effects on amphipods (Gammarus spp.), thus both the abundance and biomass were higher in treatments containing P. perfoliatus. The colonization process was significantly influenced by the numerical dominance of a few faunal species, e.g. the settlement of lagoon cockles (Cerastoderma glaucum), and by the timing of the experiments. The results indicate that faunal colonization in these communities is rapid and significantly influenced by the traits of particular plant species.     

Epiphytes from a deep-water characean meadow in an oligotrophic New Zealand lake: species composition,biomass and photosynthesis

1. The epiphytic flora of a characean meadow in Lake Coleridge, a deep, oligotrophic lake on the South Island of New Zealand, was dominated by diatoms, particularly Eunotia pectinalis and Achnanthes minutissima. The meadows occupied a depth range from 5 to 30 m. Adnate taxa predominated at all depths below 5 m, while increased taxonomic diversity at 5 m resulted from an increased abundance of erect taxa, including chlorophytes and stalked diatoms. 2. Seasonal changes in epiphyte biomass were followed using artificial substrata and by estimating epiphyte chlorophyll a concentration on host plants. The latter required development of a novel technique utilizing the consistent relationship between fucoxanthin and chlorophyll a concentrations in the epiphyton. Epiphyte chlorophyll a on host plants varied with depth and host species between 0.1 and 0.3 mg g^–1 dry weight. Maximum epiphyte biomass was at 10–15 m depth. At depths of 15 m and less, epiphyte chlorophyll a reached a maximum of ≈ 200–300 mg m^–2 in mid-summer, while at greater depths maximum biomass was less and coincided with a period of clear water in spring. 3. Photosynthetic carbon fixation was estimated from photosynthesis–radiation curves and estimates of radiation flux at sampling depths. At depths greater than 10 m, variability of the vertical extinction coefficient of lake water rather than seasonal fluctuations in incident radiation were responsible for determining the temporal pattern of production. Chlorophyll a-specific photosynthesis was estimated to peak in summer at 5 m (8 mg mg^–1 day^–1), and in spring at all other depths. 4. Annual epiphyte production was estimated as 27 g C m^–2 year^–1 at 5 m depth, falling to 15 g C m^–2 year^–1 at 15 m and 1 g C m^–2 year^–1 at 30 m. Areal biomass changes tended to be temporally but not quantitatively coupled to estimated in situ photosynthesis, and we hypothesize that epiphyte biomass may have been controlled by grazing gastropod snails.