Epiphyte host preferences and host traits: mechanisms for species-specific interactions

We investigated species-specific relationships among two species of vascular epiphytes and ten host tree species in a coastal plain forest in the southeastern United States. The epiphytes Tillandsia usneoides and Polypodium polypodioides were highly associated with particular host species in the field, but host traits that favored colonization were inadequate to fully explain the epiphyte-host associations for either epiphyte. Field transplant experiments that bypassed epiphyte colonization demonstrated that the growth of epiphytes was significantly higher on host tree species that naturally bore high epiphyte loads than on host species with few or no epiphytes. These species-specific relationships were highly correlated with the water-holding capacity of the host tree's bark. Positive and negative effects of throughfall, light attenuation by the canopy, and bark stability did not explain the overall patterns of host specificity, but did correlate with some epiphyte-host species relationships. The relative importance of particular host traits differed between the "atmospheric epiphyte" Tillandsia, and the fern Polypodium, which roots in the bark of its hosts. Species-specific interactions among plants, such as those described here, suggest that communities are more than individualistic assemblages of co-occurring species.     

Does structural parasitism by epiphytes exist? A case study between Tillandsia recurvata and Parkinsonia praecox

The effects that epiphytes have on their hosts have been poorly explored in an experimentally. Correlational evidence suggests that epiphytes may be either mutualists or structural parasites, as has been proposed for Tillandsia recurvata on Parkinsonia praecox. To test the effect of T. recurvata upon P. praecox, the epiphyte load on branches of P. praecox was measured and two 1‐year experiments were performed to detect the effect of transplantation/removal of epiphytes and shade (0%, 35%. 50% and 80%) on shoot dynamics. If T. recurvata represents a selective pressure for P. praecox, then the frequency of branches carrying large epiphyte loads will be high, and host shoot survival will decrease in the presence of T. recurvata and with increased shade. A weak inverse relationship between epiphyte load and percentage of dead shoots in the host was detected. Shoot survival was independent of epiphyte presence. Shade decreased shoot survival by 35–72%. Results suggest that at the study site, T. recurvata is a commensalist of P. praecox. An alternative hypothesis to explain the correlation between high epiphyte load and branch/tree decay is that older branches carry more epiphytes, receive more shade from neighbouring branches and could be undergoing a natural decline process.     

Community interactions between the filamentous alga Cladophora glomerata (L.) Kuetzing,its epiphytes,and epiphyte grazers

Summary Interactions between epiphytes, epiphyte grazers and the filamentous green alga Cladophora glomerata (L.) Kuetzing were explored with smaples from rivers in Montana. Extracts of C. glomerata lowered photosynthetic rates of Nitzschia fonticola Grunow (an epiphytic diatom). Nutrient enrichment showed that C. glomerata from the Madison River was N deficient and its epiphytes were P deficient on 2 dates and N deficient on one date, while no nutrient deficiencies were detected in samples from 3 other rivers; this implies there was little nutrient competition between the epiphytes and C. glomerata. Epiphytes lowered drag on C. glomerata tufts and current velocity inside the tufts, apparently by decreasing the effective surface area. Lower drag may decrease detachment, but lowering current velocity from 8 to 0 cm s^-1 resulted in a 100 % decrease in photosynthesis. Light absorption by epiphyte pigments may lower photosynthetic rate of C. glomerata when irradiance is below 200–500 E m^-2 s^-1, and protect against photoinhibition above this irradiance range. Invertebrate grazers (predominantly Baetis tricaudatus Dodds, Trycorythodes minutus Traver and Brachycentrus occidentalis Banks) at high densities removed 75% of epiphytes and B. occidentalis grazed on C. glomerata. Invertebrates regenerated a mean of 0.16 mol NH _inf4 ^sup+ individual^-1 d-1 which could have enhanced growth of downstream C. glomerata. Competition and grazing were not the only interactions in the C. glomerata community, positive (mutualistic) interactions were also important.     

Long‐term changes of the vascular epiphyte assemblage on the palm Socratea exorrhiza in a lowland forest in Panama

Question: What are the qualitative and quantitative long‐term changes in the vascular epiphyte assemblage on a particular host tree species? Location: Lowland rain forest of the San Lorenzo Crane Plot, Republic of Panama. Methods: We followed the fate of the vascular epiphyte assemblage on 99 individuals of the palm Socratea exorrhiza by three censuses over the course of five years. Results: The composition of the epiphyte assemblage changed little during the course of the study. While the similarity of epiphyte vegetation decreased on individual palms through time, the similarity analysed over all palms increased. Even well established epiphyte individuals experienced high mortality with only 46% of the originally mapped individuals surviving the following five years. We found a positive correlation between host tree size and epiphyte richness and detected higher colonization rates of epiphytes per surface area on larger trees. Conclusions Epiphyte assemblages on individual S. exorrhiza trees were highly dynamic while the overall composition of the epiphyte vegetation on the host tree species in the study plot was stable. We suggest that higher recruitment rates, due to localized seed dispersal by already established epiphytes, on larger palms promote the colonization of epiphytes on larger palms. Given the known growth rates and mortality rates of the host tree species, the maximum time available for colonization and reproduction of epiphytes on a given tree is estimated to be ca. 60 years. This time frame will probably be too short to allow assemblages to be ever saturated.     

The effect of substrate abundance in the vertical stratification of bromeliad epiphytes in a tropical dry forest (Mexico)

Diversity of epiphytes is associated with niche partitioning, through vertical strata and host preferences. However, abundance of substrate offered by hosts differs between vertical strata, misleading if epiphytes prefer a stratum or are randomly distributed. In a tropical dry forest of San Andres de la Cal Morelos, central Mexico, we tested the null hypothesis, that epiphytes follow the abundance of the substrate, rather than showing preference for a particular vertical stratum, and tested whether microclimatic variables, seed germination and seedling survival match with observed epiphyte distribution. Our data show that epiphytes could be randomly distributed inside some host; but in some host species, certain structures presented either a deficit or an excess of all, atmospheric, or tank epiphytes. In the hosts Bursera copallifera and Bursera glabrifolia, distribution of epiphytes was biased towards the upper strata, with a deficit of epiphytes in the lower strata. In Conzattia multiflora, Sapium macrocarpum and Ipomoea pauciflora, epiphyte distribution was biased towards the lower strata. Vertical gradation of light, seed germination and seedling survival did not generally match with epiphyte distribution and did not support the notion that the microclimatic gradient governs the vertical distribution of epiphytes. Our data indicate that vertical distribution of epiphytes in such tropical dry forests is mainly driven by the distribution of the structures, which apparently influence dispersion of the seeds and by the lifespan of branches, which allow the concentration of epiphytes in the stratum that optimizes seed capture and the clonal growth of epiphytes.     

Biomass accumulation and shading effects of epiphytes on leaves of the seagrass, Heterozostera tasmanica, in Victoria,Australia

A method is described for estimating the rate of accumulation of epiphyte biomass on leaves of the seagrass, Heterozostera tasmanica (Martens ex Aschers.) den Hartog and for estimating the effect of epiphyte biomass on photosynthesis of the seagrass. Epiphyte biomass was determined by comparison of the weight per unit area of epiphyte-covered and epiphyte-free leaf blades. Epiphyte weight increased as age of the seagrass leaves increased. Linear regression on epiphyte biomass vs. leaf age estimated the rate of biomass accumulation. Rates varied from 5.7 to 104 μg epiphyte dry weight per cm² of leaf surface per day at three sites in Western Port and Port Phillip Bay, Victoria. Rates of accumulation of epiphyte biomass were generally higher during December through March (summer) than in May (autumn), August (winter) or October (Spring). Light attenuation by epiphytes increase linearly with biomass. The rate of biomass accumulation of epiphytes was compared with leaf growth rate, ambient photon flux density in H. tasmanica beds and the photosynthesis—photon flux density curve of H. tasmanica. This comparison demonstrated that epiphyte biomass can accumulate fast enough to shade H. tasmanica leaves and significantly reduce the time (to less than one half of the leaf life span) in which positive net photosynthesis of the leaf blade is possible.     

Is branch damage by xylophages related to the presence of epiphytes?

Host identity influences the guilds (epiphytes and xylophages) that interact within canopies. Host species can be categorized as either limiting or preferred hosts based on epiphyte load. It is possible that, depending on the host category (limiting or preferred), galleries bored by xylophages would affect the quality and availability of space for epiphytes. The objective of this study was to determine, among and within limiting and preferred hosts, the relationship between the damage inflicted by insects to branches and epiphytic bromeliads. We collected two branches each (with and without epiphytes, respectively) from limiting hosts (Bursera fagaroides, Ipomoea murucoides and I. pauciflora) and preferred hosts (Bursera copallifera and B. glabrifolia). The variables measured were: number and species of epiphytes, number of holes, number and taxonomical group of insects, percentage of epiphyte cover and percentage of area damaged by insects. These variables were compared among and within hosts and the significant correlations, where present, determined. We identified five bromeliad species and six taxonomical groups of insects. I. murucoides showed a higher proportion of damage and a larger number of insects. For the three limiting hosts, there was a negative relationship between (1) epiphyte cover and damaged area, (2) number of epiphyte individuals and number of xylophages and (3) number of epiphyte individuals and damaged area. Within species, B. copallifera, B. glabrifolia and I. pauciflora had more holes in branches that supported epiphytes than in branches without. We hypothesized that, inter-specifically, xylophages would interfere with the establishment of epiphytes by facilitating the release of allelopathics, but this possibility needs to be examined in more detail. Intra-specifically, it is possible that holes made by xylophages do not cause sufficient damage in hosts and, consequently, a possible repercussion on epiphytes is not reflected.     

Epiphytes on the rocky intertidal red alga RhodomelaLarix (Turner) C. Agardh: Negative effects on the host and food for herbivores?

The diversity of epiflora and fauna associated with a dominant turf-forming alga was examined in intertidal communities on the central Oregon coast. Epiphytes associated with the red alga, Rhodomelalarix (Turner) C. Agardh, were examined by surveying intertidal areas for the presence of epiphytes, and by following changes in epiphyte cover in marked quadrats of R. larix. The alga is host for at least 17 species of sessile plants and animals. To determine the role of some of the larger epiphytes in the community, Rhodomela plants were marked and monitored and herbivore feeding was examined. Data suggest that epiphytes decrease the growth rate of their host, increase the probability of axis breakage and decrease reproductive output. Epiphytes provide food for littorine snails and gammarid amphipods that live in the beds of the plant. Amphipods were found to decrease epiphyte cover on R. larix in laboratory tanks, suggesting that these herbivores may have beneficial effects on the host plant.     

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.     

ROLE OF SURFACE WOUNDS AND BROWN ALGAL EPIPHYTES IN THE COLONIZATION OF ASCOPHYLLUM NODOSUM (PHAEOPHYCEAE) FRONDS BY VERTEBRATA LANOSA (RHODOPHYTA)1

Ascophyllum nodosum (L.) Le Jol. forms extensive beds in wave‐sheltered, rocky intertidal habitats on the northwestern Atlantic coast. This fucoid seaweed is host to an obligate red algal epiphyte, Vertebrata lanosa (L.) T. A. Chr. [=Polysiphonia lanosa (L.) Tandy], and two facultative brown algal epiphytes, Elachista fucicola (Velley) Aresch. and Pylaiella littoralis (L.) Kjellm. Although V. lanosa can occur throughout most of the length of host fronds, it largely predominates in midfrond segments. The two brown algal epiphytes are restricted to distal segments. Through field experiments conducted in Nova Scotia, Canada, we tested the hypothesis that surface wounds are required for the colonization of distal segments of host fronds by V. lanosa. Distal tissues normally have a smooth surface because of their young age (A. nodosum fronds grow apically). By creating small wounds that mimicked grazing wounds distributed elsewhere on host fronds, we demonstrated that V. lanosa can colonize distal frond segments during the growth and reproductive season (summer and autumn). Approximately half of the artificial wounds were colonized by V. lanosa during this time. The experimental exclusion of both brown algal epiphytes from distal frond segments did not affect colonization by V. lanosa. Thus, we conclude that the absence of surface irregularities on distal segments of host fronds, specifically small wounds, is the main factor explaining the absence of V. lanosa there. We propose that further experimental work clarifying epiphyte distribution in host beds will enhance our ability to understand the functional role of epiphytes in intertidal ecosystems.     

Two species of Halimeda, a calcifying genus of tropical macroalgae, are robust to epiphytism by cyanobacteria

Cyanobacteria, an increasingly important epiphyte on macroalgae and seagrass, have been shown to have strong effects on its hosts; this association has been identified as a driving mechanism that maintains algal blooms on coral reefs. We examined both the costs and benefits of epiphytism on 2 algal congeners of Halimeda (H. tuna and H. opuntia), both of which are abundant members of tropical reef communities in the Caribbean. To evaluate potential benefits of an associational defense as well as costs to growth, we manipulated herbivore access to (uncaged/caged) and cyanobacteria presence on (epiphytized/cleaned) 2 species of Halimeda on shallow patch reefs in Belize and measured change in branch length and segment number after 10 (H. tuna) and 5 (H. opuntia) days. Cyanobacterial epiphytes did not serve as an associational defense from herbivory as there were no differences between caged and uncaged treatments for either response variable. The presence of cyanobacterial epiphytes did not affect the growth of branches or net generation of new segments, demonstrating there was also no cost to growth. The robustness of both species of Halimeda to epiphytism contrasts strongly with recent research that found strong effects of epiphytes on several other species of tropical algae. Our results may be attributed to the unique characteristics of Halimeda, a heavily physically and chemically defended algal genus, and the shallow nature of the patch reefs reducing the potential for significant light limitation. These findings suggest that close interactions such as epiphytism may not be as generalizable as originally assumed; studies must consider differences among host species, as this may lead to a better understanding of community-wide effects.     

Temporal dynamics of algal epiphytes on Leathesia marina and Colpomenia sinuosa macrothalli (Phaeophyceae)

This is the first study on the temporal dynamics of the epiphyte communities of Leathesia marina and Colpomenia sinuosa, in terms of richness, diversity and abundance, and their relationships between the hosts and the environmental factors. A total of 31 epiphyte taxa were recorded, with the two hosts sharing 64.5% of the epiphytic species found. The red algal epiphytic group had the highest abundance on both hosts and the diatoms presented the highest species richness. The epiphyte species on L. marina showed a progressive increase in abundance during the warm seasons that was related to the end of the macroscopic life stage of the host. They also showed a greater specific richness on L. marina than on C. sinuosa, but a lower diversity and evenness, and a clear dominance of the species Urospora penicilliformis and Erythrotrichia carnea. Low abundance of epiphytes was found on C. sinuosa, without temporal variation, but high diversity and evenness and without any dominant species. In addition, a clear zonation was observed on this host, where epiphytic abundance decreased on the lateral and upper part of the thallus.     

Successional Sequences of Microbial Colonization on Three Species of Rhodophycean Macroalgae

The successional sequences of microbial colonization of Centrocerasclavulatum, Bryocladia cuspidata, and Gelidium crinale wereobserved by SEM. Colonization was initiated by filamentous andsmaller rod or coccoid bacteria, and these microbes were replacedby diatom populations in a successional pattern on Centrocerasand Bryocladia. Gelidium was colonized primarily by bacteria.The spatial fouling patterns on each host plant could be correlatedwith plant shape. Differences in epiphyte attachment morphologiescould be correlated in some species with the host plant or withthe position of the epiphyte on the host plant. Diatoms, epiphytes, fouling, microbial colonization, periphyton     

DISTRIBUTION OF ALGAL EPIPHYTES ACROSS ENVIRONMENTAL GRADIENTS AT DIFFERENT SCALES: INTERTIDAL ELEVATION,HOST CANOPIES,AND HOST FRONDS1

Understanding epiphyte distribution in coastal communities is important because these organisms affect many others directly or indirectly. Yet, their distribution has been considerably less studied than that of their hosts and other primary‐space holders. Identifying major sources of variation in epiphyte abundance is thus still a need. Environmental gradients help predict species distribution and are pervasive on marine shores. In this study, we test the notion that environmental gradients across intertidal elevation, throughout host canopies, and along host fronds explain a large variation in the abundance of sympatric epiphytes. Our model system was the assemblage of Ascophyllum nodosum (L.) Le Jol. and its epiphytes Vertebrata lanosa (L.) T. A. Chr. [= Polysiphonia lanosa (L.) Tandy], Elachista fucicola (Velley) Aresch., and Pylaiella littoralis (L.) Kjellm. On the coast of Nova Scotia (Canada), we found evidence of a spatial segregation among these species at almost all scales. While the red epiphyte V. lanosa was more common at high‐ and midintertidal elevations (peaking at midelevations) and on middle segments of host fronds, the brown epiphytes E. fucicola and P. littoralis were more common at low elevations and restricted to distal segments of host fronds. Canopy habitat affected abundance only for V. lanosa, which was more common within the host canopy than on its periphery at midelevations. Since the studied gradients are related to predictable changes in abiotic factors, the identification of likely causes behind the observed patterns was facilitated. Our study ends by proposing abiotic and biotic factors that deserve priority in the experimental testing of the forces structuring this assemblage.     

THE BIOLOGY OF THE EPIPHYTIC BROMELIAD TILLANDSIA CIRCINATA SCHLECHT. I. THE NUTRIENT STATUS OF POPULATIONS IN SOUTH FLORIDA

The atmospheric epiphyte Tillandsia circinata (Bromeliaceae) was observed and collected from six diverse habitats in south Florida. Calcium, K, N, Na, Mg, and P levels were determined in prefruiting, fruiting, and postfruiting rosettes from these plants. The mineral composition varied considerably among growth stages at a site and among sites within growth stages. By weight, Na tended to predominate while the other five elements were usually present at much lower levels. Potassium and P showed pronounced positive correlations with this epiphyte's vigor and reproductive capacity whereas the other elements did not show such correlations. Although T. circinata seems able to tolerate very low nutrient levels, these data suggest that its distribution and vigor are in part determined by nutritional factors which vary with habitat and host type. Plants growing on hosts with a large leaf surface area and where ant nidification is relatively common tend to be more vigorous and fruitful than those occurring in depauperate forests which support a sparse canopy. These latter hosts, however, often harbor numerous small epiphytes which produce a few small capsules per shoot.     

Effect of nutrient availability, grazer assemblage and seagrass source population on the interaction between Thalassia testudinum (turtle grass) and its algal epiphytes

Seagrass leaves are often densely covered by epiphytic algae which can suppress seagrass productivity and has been implicated in declines of seagrass meadows worldwide. The net effect of epiphytes on seagrass growth and morphology depends on the independent and interactive effects of a variety of factors, including nutrient availability and the intensity of grazing on epiphytes. Here I report the results of a mesocosm experiment designed to test the effects of nutrient addition and within-functional group variation (grazer species composition and the source population of seagrass) on the strength of the interactions among grazers, epiphytes, and turtle grass (Thalassia testudinum). Turtle grass ramets from two sites in the northern Gulf of Mexico were cleared of epiphytes and transplanted into common-garden mesocosms. Replicate ramets were grown in a split-split plot design with two levels of dissolved nutrients and four different grazer species combinations (Tozeuma carolinense alone, Pagurus maclaughlinae alone, both species together, and no grazers present). As expected, grazers had a significant negative effect on epiphyte biomass/leaf area and a significant positive effect on turtle grass growth in the mesocosms. The two species were more similar in their direct effects on epiphyte biomass than in their indirect effects on turtle grass growth; this may reflect differences in epiphyte community composition under different grazer treatments. The effect of nutrient addition on turtle grass growth depended critically on the intensity of grazing: in the presence of grazers, turtle grass tended to produce a greater biomass of new leaf tissue in the tanks with nutrients added than in the control tanks. However, when grazers were absent, the direction of the effect was reversed, and plants with nutrients added grew less than the control plants. The two source populations of turtle grass differed significantly in epiphyte biomass/leaf area accrued in the mesocosms as well as in the strength of the effect of grazers on turtle grass growth. This suggests that population differentiation in seagrass interactions with epiphytes, as well as spatial and temporal variation in resources and grazer community composition, can greatly effect the role of epiphytes in limiting seagrass productivity.     

Epiphytism of <Emphasis Type="Italic">Gracilaria cliftonii</Emphasis> (Withell,Millar &; Kraft) from Western Australia

Epiphytism in Gracilaria is a common phenomena observed in natural populations and under culture conditions. Generally, epiphytes are attached superficially to the surface of the host however, genera such as Polysiphonia spp. and Ceramium spp. can penetrate into the host tissue affecting its growth and hence productivity. The present paper aims to identify and quantify epiphyte abundance and characterise their attachment and penetration on the natural populations of Gracilaria cliftonii collected from Shoalwater Marine Park, Perth, Western Australia. G. cliftonii samples were collected monthly for 1 year and their epiphytes were identified. Histological studies of G. cliftonii were also performed on epiphyte attachment sites. Twenty-four species of macroalgae epiphytes were recorded, 21 belonging to Rhodophyta and 3 to Chlorophyta. Hypnea episcopalis and Polysiphonia forfex were the dominant (p < 0.05) species during the sampling period. The maximum epiphytic load was observed in October (40%), however, the maximum diversity of epiphytes was recorded in September and February (n = 14). Light and scanning electron microscopy studies of the epiphyte attachment sites revealed that P. forfex and Ceramium isogonum penetrated into the cortex and outer medulla of G. cliftonii, while Bryopsis plumosa and Laurencia clavata altered the cellular structure of the cell wall of G. cliftonii.     

The impact of Dictyota spp. on Halimeda populations of Conch Reef, Florida Keys

Species of the brown alga Dictyota dominate the reef tract in the Florida Keys. In surveys during summer and fall months between 1994 and 2001, Dictyota occupied as much as 70% of the benthos on Conch Reef. Dictyota spp. were found growing epiphytically on Halimeda tuna, Halimeda opuntia, Lobophora variegata, Galaxura sp., fire coral, hard corals, soft corals, bryozoans and a variety of sponges on Conch Reef. From 1994 to 2001, the percent coverage of Halimeda spp. declined from 15% to 3% on the same reef. In Aug. 1999, 2000 and 2001, on average, 56% of two Halimeda species on Conch Reef had >50% of their thalli covered by Dictyota menstrualis and Dictyota pulchella. To address the impact of Dictyota on Halimeda, short-term growth of unepiphytized and heavily epiphytized (>50% Dictyota cover) H. tuna were compared with unepiphytized H. tuna to which a Dictyota mimic was attached. The number of new segments per plant ranged from 1 to 174 over 9 days. Halimeda thalli with >50% Dicytota cover and thalli covered with Dictyota mimic grew significantly slower than unepiphytized thalli. A second short-term experiment addressed the impact of neighboring Dictyota on the growth and metabolism of unepiphytized H. tuna. Augmenting or clearing epilithic Dictyota around but not in contact with H. tuna had no impact on growth or metabolism of H. tuna. Unepiphytized and heavily epiphytized H. tuna were also collected for studies of metabolism. This work indicated that epiphytic Dictyota negatively impacts metabolic rates of H. tuna in part by shading H. tuna thalli. This negative impact was also in part chemically mediated, as exposure to Dictyota-conditioned water elevated respiration rates in the same manner as seen in the metabolic studies of naturally epiphytized H. tuna.     

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.     

Patterns in the diversity and distribution of epiphytes and vines in a New Zealand forest

Abstract Plants that rely on other plants for support (i.e. epiphytes and vines) are common in many forest ecosystems. However, they are poorly understood relative to terrestrial plants, especially in the Southern Hemisphere. To help bridge this gap, we evaluated the diversity and distribution of vascular epiphytes and vines on seven common tree species in a conifer‐broadleaf forest on New Zealand's North Island. Ground‐based surveys of 274 host trees were used to test whether epiphyte and vine diversity increased with tree diameter, and whether diversity‐diameter relationships differed among host tree species. Occurrence patterns of individual epiphyte and vine species were also assessed. We first evaluated the accuracy of ground‐based inventories by comparing surveys of trees made from the ground to those made from a canopy walkway. On average, 1 in 10 species of epiphytes and vines were unseen from the ground. However, sampling accuracy did not differ among the three host tree species growing along the walkway, suggesting unbiased comparisons could be made between hosts. Results from ground‐based surveys showed that species diversity of epiphytes and vines increased with host tree diameter. However, epiphytes showed stronger diversity‐diameter relationships than vines. Epiphyte diversity increased markedly in four host species and less strongly in the remaining three host species. Conversely, vines showed weak diversity‐diameter relationships in all host species. Occurrence patterns of individual species helped to explain diversity‐diameter relationships. All common epiphyte species occurred more frequently on large trees, regardless of host species, but occurrence patterns in most vine species were unrelated to tree size. Rather, the vines often showed strong host ‘preferences’. Overall results illustrate a rich diversity of distributional patterns in New Zealand's epiphytes and vines, and suggest that a similarly diverse set of ecological and evolutionary processes are responsible for them.