Phlorotannins versus other factors affecting epiphyte abundance on the kelp Ecklonia radiata

We examined factors affecting the abundance and distribution of epiphytes (fouling) on the sublittoral kelp Ecklonia radiata. We first assessed the importance of phlorotannins as chemical defences against epiphytes by (a) correlating epiphyte loads on different parts of the thallus with the phlorotannin content of those tissues, and (b) experimentally testing the effects of variation in phlorotannin concentration against the settlement and growth of gametes of Ulva lactuca, a common epiphyte in the system. Tissue phlorotannin content was, at best, only weakly related to epiphyte loads, with r ² values typically <0.10. Inhibition of Ulva gametes only occurred at concentrations >10 mg l⁻¹, which is 5 orders of magnitude greater than levels of phlorotannins in the water column around beds of E. radiata, and 1–3 orders of magnitude greater than estimated levels in the boundary layer at the surface of the plant. We concluded that phlorotannins have a negligible impact on patterns of epiphytism on E. radiata, and proceeded to investigate other factors influencing the distribution and abundance of epiphytes. In our samples the relative age of different parts of the thallus was strongly correlated with epiphyte abundance, with epiphyte densities greatest on the oldest tissue and least on the youngest. Distal parts of the thalli also had greater epiphyte loads than basal parts. Field experiments in which kelp tissue was suspended at two heights in an E. radiata forest for varying lengths of time confirmed the importance of the length of time that the tissue was in the water, and its height in the water column, to the development of an epiphyte community. Comparison of epiphyte loads on tissue from primary (smooth) and secondary (rough) laminae in these experiments indicated that surface rugosity also affected fouling. Macroherbivores were rare on E. radiata, and abundances of mesofauna and epiphytes were positively related, suggesting that grazers were not important determinants of patterns of epiphyte abundance. Although phlorotannins have been previously suggested to play an important role as defences against epiphytes, we suggest that water-soluble compounds such as phlorotannins are less likely to be effective defences against epiphytes than non-polar metabolites, which can adhere to the surface of the producing organism.     

Non‐consumptive effects of avian predators on fish behavior and cascading indirect interactions in seagrasses

Top–down impacts of avian predators are often overlooked in marine environments despite evidence from other systems that birds significantly impact animal distribution and behavior; instead, birds are typically recognized for the impacts of their nutrient rich guano. This is especially true in shallow seagrass meadows where restoration methods utilize bird perches or stakes to attract birds as a passive fertilizer delivery system that promotes the regrowth of damaged seagrasses. However, this method also increases the local density of avian piscivores that may have multiple unexplored non‐consumptive effects on fish behavior and indirect impacts to seagrass communities. We utilized laboratory and field experiments to investigate whether visual cues of avian predators impacted the behavior of the dominant demersal fish in seagrass habitats, the pinfish Lagodon rhomboides, and promoted cascading interactions on seagrass‐associated fauna and epiphytes. In laboratory mesocosms, pinfish displayed species specific responses to models of avian predators, with herons inducing the greatest avoidance behaviors. Avoidance patterns were confirmed in field seagrass meadows where heron models significantly reduced the number of fish caught in traps. In a long term field experiment, we investigated whether avian predators caused indirect non‐consumptive effects on seagrass communities by monitoring fish abundances, invertebrate epiphyte grazers, and the seagrass epiphytes in response to heron models, bird exclusions, and bird stakes. On average, more fish were recovered under bird exclusions and fewer fish under heron models. However, we found no evidence of cascading effects on invertebrate grazers or epiphytes. Bird stake treatments only displayed a simple nutrient effect where higher bird abundances resulted in higher epiphyte biomass. Our results indicate that although birds and their visual cues can affect fish and epiphyte abundance through non‐consumptive effects and nutrient enrichment, these impacts do not propagate beyond one trophic level, most likely because of dampening by omnivory and larger scale processes.     

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.     

Spatial ecology of multiple parasitoids of a patchily‐distributed host: implications for species coexistence

1. The coexistence of multiple species sharing similar but spatially fragmented resources (e.g. parasitoids sharing a host species) may depend on their relative competitive and dispersal abilities, or on fine‐scale resource partitioning. Four generalist and one specialist parasitoid species associated with the holly leaf miner, Phytomyza ilicis, in a woodland network of 127 holly trees were investigated. 2. To understand coexistence and persistence of these potential competitors, patterns of occurrence in relation to patch size and isolation, vertical stratum within patches, and incidence and abundance of potential competitors were documented. Field experiments creating empty habitat patches suggested that dispersal rather than local demographic processes determines abundance and incidence. 3. Parasitoids showed species‐specific responses to patch properties, with the incidence of species determined mostly by patch size. Parasitism rates were less clearly related to patch characteristics, but parasitism rates for most species were lower in patches where the numerically dominant parasitoid species, Chrysocharis gemma, was present. No evidence of vertical stratification was found in species composition or abundance within patches, making it unlikely that coexistence is enhanced by fine‐scale resource division. 4. Overall, the patterns detected may be attributed to the distribution of C. gemma and differences in species' ecology other than dispersal ability. The life history of C. gemma may allow it to pre‐emptively exploit a large fraction of the available hosts, avoiding direct competition with other parasitoids. In contrast, direct competition is more likely among the pupal parasitoids Cyrtogaster vulgaris, Chrysocharis pubicornis, and Sphegigaster flavicornis which have a similar biology and phenology. For these species, coexistence may be facilitated by contrasting incidence in relation to patch size and isolation.     

The epiphyte vegetation of Annona glabra on Barro Colorado Island, Panama

Aim Information on the community composition, structure, and dynamics of epiphyte vegetation is scarce. A survey of the epiphytes occurring on all individuals of one particular host tree species in a well-studied neotropical research site allowed us a comparison of the epiphyte flora of this tree with the local epiphyte flora, the analysis of spatial distribution patterns and the use of these patterns as indications for changes in time. In the future, our results can be used as a baseline data-set for the direct observation of the long-term dynamics in epiphyte communities. Location The study was conducted on Barro Colorado Island (BCI), Panama. Methods We recorded all individuals of the vascular epiphytes growing on Annona glabra L., a flood-tolerant, multiple-stemmed tree, which is restricted to the shoreline of BCI. Data on tree biometrics, epiphyte species, and epiphyte abundances were collected for more than 1200 trees. Results In total, we encountered almost 15,000 epiphytic individuals in sixty-eight species, corresponding to more than one third of the entire epiphyte flora of Barro Colorado Island. The component species differed strongly in abundance: the four most important species accounted for >75% of all individuals. In most cases, the same four species were also the first to colonize a tree (=phorophyte). Colonization patterns indicated no replacement of early colonizers by late arrivals. Species richness and epiphyte abundances showed a positive correlation with the size and the density of the host trees. All species showed a highly clumped distribution and the physiognomy of epiphyte communities of individual trees was dominated either by one or several of the four most common species or by a set of frequently co-occurring tank bromeliads. Other species were dominant only in exceptional cases. Most species were always rare. A distance effect on community composition was mostly confined to a local scale with an increased similarity in the species assemblage of stems of a tree v. neighbouring trees. Main conclusions The epiphytes on a single small phorophyte species may encompass a surprisingly large proportion of the local epiphyte flora. The observations that most tree crowns are inhabited by a single or only very few species, and that all epiphyte species show highly clumped distributions suggest a predominance of very local dispersal within a tree crown, which is only infrequently interrupted by successful long-distance dispersal between crowns.     

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.     

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.     

Conservation of Vascular Epiphyte Diversity in Shade Cacao Plantations in the Chocó Region of Ecuador

To assess the contributions of rustic shade cacao plantations to vascular epiphyte conservation, we compared epiphyte species richness, abundance, composition, and vertical distributions on shade trees and in the understories of six plantations and adjacent natural forests. On three phorophytes and three 10 × 10 m understory plots in each of the agroforestry plantations and natural forests, 54 and 77 species were observed, respectively. Individual-based rarefaction curves revealed that epiphyte species richness was significantly higher on forest phorophytes than on cacao farm shade trees; detailed analyses showed that the differences were confined to the inner and outer crown zones of the phorophytes. No differences in epiphyte species richness were found in understories. Araceae, Piperaceae, and Pteridophyta were less species-rich in plantations than in forests, while there were no differences in Orchidaceae and Bromeliaceae. Regression analysis revealed that epiphyte species richness on trunks varied with canopy cover, while abundance was more closely related to soil pH, canopy cover, and phorophyte height. For crown epiphytes, phorophyte diameter at breast height (dbh) explained much of the variation in species richness and abundance. There were also pronounced downward shifts in the vertical distributions of epiphyte species in agroforests relative to natural forests. The results confirm that epiphyte diversity, composition, and vertical distributions are useful indicators of human disturbance and showed that while the studied plantations serve to preserve portions of epiphyte diversity in the landscape, their presence does not fully compensate for the loss of forests.     

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.     

A metapopulation approach to the analysis of long‐term changes in the epiphyte vegetation on the host tree Annona glabra

Question: Do vascular epiphyte species have a metapopulation structure? What are the qualitative and quantitative long‐term changes of the complete vascular epiphyte vegetation in a particular host tree species? Location: Lowland forest on Barro Colorado Island (9° 10’ N, 79°51’ W), Republic of Panama. Methods: In 1994 and 2002 we conducted a census of all vascular epiphytes growing on more than 1000 Annona glabra trees (= patches). Epiphyte species abundances were recorded at the tree level in each census. Results: The number of epiphyte individuals increased from ca. 15 000 to ca. 23 700 individuals during the census interval while the species composition on Annona glabra as a whole was rather stable. There was a strong positive relationship between occurrence in patches and local abundance of the species, and between species richness and host tree stand size. The dynamics of local populations of a given species were uncorrelated to each other; small and large local populations of most species had the same probability to go extinct. The frequency distribution of species on all host trees was not bimodal, but on a subset of heavily colonized host tree stands it was. Numbers of species and individuals were correlated with tree size which was not due to a correlation of tree size and tree age. Conclusions: As far as the most abundant epiphyte species with metapopulation structures are concerned, these species belong to diverse families, e. g. Orchidaceae, Bromeliaceae and Polypodiaceae. Even ca. 80 years after the initial establishment of the host tree species in the study area epiphytes are still in the stage of initial colonization and have not reached a steady state as indicated by the strong increase in individuals and the ongoing colonization of empty trees.     

海南岛热带天然针叶林附生维管植物多样性和分布

作为热带林中一个重要的特征性组分, 附生维管植物对于维持热带森林的物种多样性及其生态系统功能均具有重要作用。该文首次系统地报道了热带天然针叶林中的附生维管植物多样性和分布特征。以海南岛霸王岭国家级自然保护区保存完好的热带天然针叶林(我国唯一较大面积分布的南亚松(Pinus latteri)天然林)中的附生维管植物为研究对象, 通过样带调查(共设置12个10 m × 50 m的样带, 记录每个样带内胸径(DBH) ≥ 5 cm树木上附生维管植物的物种名称、株数及附生高度), 分析附生维管植物的物种多样性和空间分布特征。结果表明: 1)热带针叶林0.6 hm ²面积内共有附生维管植物769株, 分属于7科17属27种, 附生兰科植物和萝摩科植物为优势类群; 2)附生维管植物在水平方向上呈现出聚集分布; 3)附生维管植物在垂直方向上, 在中等高度层次(10-20 m)分布最多, 在下层(0-5 m)也有较多的分布; 4)少数附生维管植物对南亚松表现出一定的选择性, 如华南马尾杉(Phlegmariurus fordii)、玫瑰毛兰(Eria rosea)、眼树莲(Dischidia chinensis)和铁草鞋(Hoya pottsii)等; 5)附生维管植物的物种丰富度及多度与宿主胸径均存在显著的正相关关系。     

Diversity of phlorotannin profiles among sargassasacean species affecting variation and abundance of epiphytes

In general, epiphytes have detrimental effects on the growth of their basiphytes due to competition for light and nutrients. Therefore, basiphyte species must expend energy suppressing epiphytes. Some studies suggest that phlorotannins, i.e. brown algal polyphenols, prevent colonization by epiphytes, whereas others question their allelopathic function because there is not necessarily a negative correlation between epiphyte abundance and the phlorotannin content of the basiphyte algae. Various phlorotannin components are found in brown algal species, thus we hypothesized that the antifouling activities of polyphenolic compounds may differ and that the analysis of phlorotannin profiles could be useful for estimating their ecological functions. We surveyed the epiphyte richness in the apical portions of 373 thalli from 15 sargassacean species, demonstrating that the variation and abundance of epiphyte species differed remarkably among the basiphyte species. However, there was a weak negative correlation between the density and total phlorotannin content of the basiphyte algae in only one of the 18 epiphyte species. The interspecific differences in the phlorotannin profile were characterized by quantitative ¹H nuclear magnetic resonance spectroscopy (qNMR), and four major groups were categorized based on cluster and principal component analyses of polyphenolic signals in the qNMR spectra. The epiphyte Neosiphonia harveyi was more abundant on Sargassum hemiphyllum, S. patens and S. piluliferum than on other basiphyte species, and these three species were similar according to the cluster analysis. These results suggest that some phlorotannin components may be more effective for antifouling; thus interspecific differences in the phlorotannin profile could affect the variation and abundance of epiphytes.     

Site specific distribution of the mud shore crab <Emphasis Type="Italic">Helograpsus haswellianus</Emphasis> in temperate wetlands

An important aspect in determining the ecology of a species is an understanding of its abundance and distribution. The population structure and distribution of Helograpsus haswellianus (Brachyura, Grapsidae) were evaluated over 12 months at four intertidal wetland sites that differed in vegetation and inundation. Surface activity of H. haswellianus varied over time with no consistent seasonal pattern. Sex ratios were always significantly skewed towards males, and crabs of both sexes were distributed evenly across habitats at all sites. Recruitment patterns and size frequency distributions varied significantly between sites, independent of habitats. Crabs were smaller in wetter, more sheltered areas, but size ranges overlapped between habitats. The site-specific difference in population structure and distribution of H. haswellianus indicates that the functional role of this burrowing crab may vary across wetlands.     

Abundance dynamics and functional role of predaceous <Emphasis Type="Italic">Leptodora kindtii</Emphasis> in the Curonian Lagoon

The abundance and distribution of predatory cladoceran Leptodora kindtii was investigated in the estuarine lagoon (Curonian Lagoon, SE Baltic Sea). Three hydrodynamically different parts of the lagoon were selected, representing transitory oligohaline, intermediate freshwater and stagnant freshwater sites. L. kindtii was least abundant at the oligohaline site, never occurring at salinities greater than 4 psu. At the two freshwater sites, the abundance of L. kindtii varied from a low of <0.1 up to 2.2 indv. L⁻¹ during peak abundance. Two peaks of L. kindtii abundance were observed with timing differences between stations: at the stagnant site the population of L. kindtii peaked two weeks earlier relative to the more hydrodynamically active sites, likely due to a 2°C higher May temperature. The small body size of L. kindtii in the lagoon (seasonal mean 2.68±0.6 mm) shows high fish predation pressure and predicts small cladocerans, juvenile copepods and rotifers being in the preferred prey size range. The calculated L. kindtii daily consumption during the population peak was as high as 100% of the daily zooplankton production, which implies high potential of this predator to shape the grazing zooplankton community in the lagoon.     

Local coexistence and niche differences between the Lusitanian and Mediterranean pine voles (<Emphasis Type="Italic">Microtus lusitanicus</Emphasis> and <Emphasis Type="Italic">M. duodecimcostatus</Emphasis>)

In the present study, we analyzed the coexistence pattern of the Lusitanian pine vole (Microtus lusitanicus) and the Mediterranean pine vole (Microtus duodecimcostatus) in a potential area of sympatry in a Mediterranean landscape (Portugal). We also determined the relative contribution of local, landscape, and spatial factors explaining the differences in the distribution patterns of the two species in the region. Using a kriging interpolation method, we obtained a map of sympatric and allopatric areas of species occurrence. The estimated sympatry area corresponded to a northwest–southeast belt representing 11.3% of the study area. Habitat niche differences were assessed with binomial GLMs followed by a variance partitioning. At a local scale, higher altitude, higher cover of shrubs, lower clay content in the soil, and lower cover of tree canopy were the most important factors distinguishing M. lusitanicus presence sites from those with M. duodecimcostatus. At a larger scale, the presence of forest landscape units and the low abundance of “montado” units were the most influencing landscape factors in the identification of M. lusitanicus occurrence sites when compared to M. duodecimcostatus. Our results suggested that local coexistence of M. lusitanicus and M. duodecimcostatus in the field is a rare event. The differences in distribution patterns of the two pine vole species were mostly explained by fine-scale environmental factors and by shared spatial effects.     

Species composition, comparative size and abundance of the genus Littoraria (Gastropoda: Littorinidae) from different mangrove strata along the East African coast

Variation in the abundance, distribution and size of four species of mangrove littorinid gastropods (genus Littoraria) was investigated using a nested sampling design at different spatial scales along the East African coast, from Tanzania to South Africa. Littorinid abundance and diversity decreased abruptly south of Inhaca Island at the southern end of the study area. All species presented a large-scale spatial variation in abundance, with L. subvittata showing the greatest abundance while L. intermedia was rare. Littoraria scabra and L. intermedia were found mainly at the seaward edge of the forests. Littoraria subvittata increased in abundance in the middle of the forest and towards the landward side. Littoraria pallescens occurred mainly at the seaward edge and in the middle areas in the Rhizophora zone. These small-scale variations show contrasting specific distribution patterns within the mangrove, likely reflecting different tolerances to physical factors and biological interactions. All species appeared decreased in shell height from north to south. Littoraria scabra was always significantly larger than other species at all mangrove study sites. Handling editor: P. Viaroli     

Effects of seasonality on the species–area relationship: a case study with birds in urban parks

Aim To examine how parameters of the species–area curve and factors determining bird species richness are affected by seasonality. Location One hundred and thirty urban parks of the city of Valencia (Spain) ranging between 0.03 and 18.6 ha. Methods Bird censuses were conducted monthly during 1998 and 2004. For each park, 27 variables were recorded as measures of size, structure and isolation. Linear regression was used to test for patterns in species richness relative to independent variables. The results of each season were compared. Results Curves corresponding to the breeding period (spring–summer) had significantly higher intercepts, but slopes did not change across seasons. Turnover rates of resident species were dependent on patch size and isolation considering spring, but not autumn, from different years. Additionally, turnover rates in spring were lower in parks in close proximity to others than in isolated ones. In all seasons, the number of bird species showed a strong relationship with park area, which explained c. 60% of the variation in species numbers. Habitat variables and park isolation explained a small amount of the variation in species numbers. The data did not give support to the random placement hypothesis. Main conclusions The overall conclusions of this study give little support to the possible influence of seasonality on species–area relationship or on the three main hypotheses explaining it. This lack of influence of seasonality could be explained by the seasonal stability of the bird populations of the parks within the study area as a consequence of different factors, mainly the characteristics of the species pool, similarity in the features of the parks irrespective of their size, and stability of resources and conditions across seasons. The question of whether this lack of influence of seasonality on the species–area relationship occurs in those ecological systems that have a larger seasonal variation than the urban parks of the study area remains unresolved. Closer examination of the seasonal patterns reported in this study is likely to be useful in increasing our understanding of the species–area relationship.     

Intrashoot distributions of leaf dwelling harpacticoid copepods on the seagrass Zostera marina L.: implications for sampling design

The distribution of leaf dwelling harpacticoid copepods within seagrass (Zostera marina L.) shoots was investigated on four dates in 1986 and 1987. Copepods were found to be non-uniformly distributed on shoots, with higher abundances observed on older leaves. Patterns of abundance within shoots could not be explained by the surface area of individual leaves except on the sampling date with the highest copepod densities. It is suggested that harpacticoid copepod distributions on seagrass shoots are primarily determined by the pattern of epiphytic biomass. However, at high population densities, habitable surface area may be the limiting factor. Increased habitat complexity at high epiphyte loads does not seem to be the cause of the copepod distributions observed in this study. An accurate method for estimating seagrass copepod abundance per unit sediment area using intrashoot distributions is described and compared to existing methodology.     

Development of a diverse epiphyte community in response to phosphorus fertilization

The role of terrestrial soil nutrient supply in determining the composition and productivity of epiphyte communities has been little investigated. In a montane Hawaiian rainforest, we documented dramatic increases in the abundance and species richness of canopy epiphytes in a forest that had been fertilized annually with phosphorus (P) for 15 years; there was no response in forest that had been fertilized with nitrogen (N) or other nutrients. The response of N-fixing lichens to P fertilization was particularly strong, although mosses and non-N-fixing lichens also increased in abundance and diversity. We show that enhancement of canopy P availability is the most likely factor driving the bloom in epiphytes. These results provide strong evidence that terrestrial soil fertility may structure epiphyte communities, and in particular that the abundance of N-fixing lichens – a functionally important epiphyte group – may be particularly sensitive to ecosystem P availability.