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.     

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.     

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.     

FACTORS ASSOCIATED WITH HOST SPECIFICITY IN SPOROCLADOPSIS NOVAE-ZELANDIAE (CHLOROPHYTA)1

This study tests the hypotheses that substrate stability and rugose microtopography are the main factors that determine the tissue-specific epiphytism in the chlorophyte Sporocladopsis novae-zelandiae Chapman. Both in vitro and field experiments showed that the epiphyte did not develop on stable algal surfaces nor on artificial (smooth or rugose) substrata. In field experiments, however, other macroalgae settled more abundantly on rugose substrata. Similarly, our field experiments did not support the hypothesis that the physical environment provided by the spatial location of the host in the intertidal or subtidal was a relevant factor associated with specificity. In contrast, our laboratory experiments suggested that some component of the host cell wall might trigger the formation of penetrating rhizoids. These did not develop in isolated cultures of the epiphyte but were detected when the epiphyte contacted its natural host. Also, preliminary results showed that rhizoid-like structures developed, although at very low frequency, only in plants grown in culture medium containing a crude extract of sorus cortical tissue.     

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.     

Seasonal Occurrences of Epiphytic Algae on the Commercially Cultivated Red Alga Kappaphycus Alvarezii (Solieriaceae, Gigartinales, Rhodophyta)

Common problems faced in farming of the red algal genus Kappaphycus/Eucheuma are “ice-ice disease” and the occurrence of epiphytes. Considerable work has been documented on “ice-ice disease” and it's mode of infection but limited information is available on the emergence of epiphytes. The present study addresses the phenomenon of epiphyte infection, its prevalence in commercially cultivated red alga, Kappaphycus alvarezii, and their variability associated with seasonality. Cultured seaweed became susceptible to epiphytes in the dry seasons (1) between March – June and (2) September – November. Findings revealed Neosiphonia savatieri (Hariot) M. S. Kim et I. K. Lee, as the dominant infecting epiphyte, representing up to 80–85% of the epiphyte present during peak seasons. Besides N. savatieri, Neosiphonia apiculata, Ceramium sp., Acanthophora sp. and Centroceras sp. were observed in smaller quantities. SEM (Scanning Electron Microscope) micrographs revealed the epiphyte's attachment to the host. Further histological study showed the extent of penetration of epiphytes into the host's cortex tissues and condition of its surrounding tissues. The outbreak of epiphytic filamentous red algae also correlated with drastic changes in seawater temperature and salinity during March– June and September – November.     

DIATOM EPIPHYTES ON ODONTHALIA FLOCCOSA: THE IMPORTANCE OF EXTENT AND TIMING

The interaction between the epiphytic diatom Isthmia nervosa Kütz and the red alga Odonthalia floccosa (Esp.) Falkenb. was examined in terms of host physiology (photosynthesis and growth), fitness (reproduction and survival), and population dynamics. Most of these characteristics were compared among plants that varied naturally in epiphyte load. Diatom cover was also manipulated in field-deployed containers to study effects on host growth. Increasing levels of epiphytes were correlated with declines in host photosynthesis and growth, and experiments confirmed that reduced growth was directly caused by diatoms. Populations that hosted diatoms throughout the summer contained lower proportions of reproductive plants than did populations colonized in late summer or not epiphytized at all (60% vs. >80%, respectively). Odonthalia with epiphytes experienced greater biomechanical drag when submerged than did unepiphytized individuals of similar size. Despite differences in drag, epiphytized and unepiphytized hosts were equally susceptible to complete mortality (removed with holdfast) relative to partial mortality (shoots broken). In general, higher epiphytism was associated with reduced host performance, and this apparent damage was easier to detect at physiological levels than at population levels. In fact, negative effects of epiphytes were not evident at the population level, and host density and epiphyte load were positively correlated. Based on 4 years of monthly censuses of Odonthalia using marked plants and quadrats, the abundance of Isthmia varies spatially and temporally. Diatoms tend to colonize Odonthalia in late summer, after host growth has stopped and cystocarps and tetraspores have begun to develop. Subsequently, shoots break and plants persist through the winter as shorter perennating basal systems. Life spans were >4 years for some Odonthalia individuals. Many hosts may escape the effects of epiphytes by completing growth and reproduction before diatom colonization.     

Interaction mechanisms between Gracilaria chilensis (Rhodophyta) and epiphytes

Epiphytes can have a considerable effect on Gracilaria production, and Ulva is one of the commonest algal species identified as an epiphyte, reaching loads of 60% (g of epiphytes per g of Gracilaria) in the intertidal cultures of southern Chile. This study evaluates the relative importance of light reduction, addition of weight to the host thalli and nutrient depletion, as mechanisms determining the interaction effects of Ulva epiphytes on Gracilaria cultivation. Using field experiments undertaken during the main Gracilaria growth season (spring), we evaluate the mechanisms of epiphyte-host algae interaction by manipulating artificial epiphytes. The results indicate that Ulva can significantly depress Gracilaria biomass production and that the addition of weight to the host algae and the consequent dislodgement increase, appear to be the main mechanisms involved in the Ulva-Gracilaria interaction. However, the light reduction caused by the epiphytes can also partially explain the reduction in Gracilaria production. Nutrients depletion would not appear to fully account for the detrimental effects of Ulva over Gracilaria in intertidal farming areas of southern Chile.     

Distribution and development of some Manx epiphyte populations

The success ofElachista fucicola as an epiphyte ofFucus vesiculosus is partly due to its ability to grow on most parts of the host thallus. Its ability to reach reproductive maturity quickly ensures its fitness, even on temporary structures such as receptacles. Epiphyte distribution may also vary with host morphology.Himanthalia elongata is a basiphyte for several tissue-specific epiphytes:Ectocarpus fasciculatus, Elachista scutulata, Herponema velutinum andMyriactula areschougii are exclusive, or almost so, to receptacles. On growing receptacles their specificity extends to conceptacles, which also provide refuges from grazing. Epiphyte abundance in summer is greatest at the upper and lower limits ofHimanthalia distribution, suggesting that basiphytes living under suboptimal conditions are more prone to infestation by epiphytes. 5–10% ofHimanthalia receptacles are thought to persist over winter and so to facilitate the transmission of epiphytes to the following year's crop of receptacles. The distribution ofEctocarpus fasciculatus onLaminaria digitata differs in pattern from that onHimanthalia. The twoEctocarpus populations also differ morphologically, but both have reproductive and developmental, characteristics that are in accord with the ecological cycles of their basiphytes. This characteristic is likely to be an important factor in determining the success of an epiphyte. Success may also be facilitated by epiphyte variability or by possession of a fixed but well-adapted phenotype. The very high incidence of brown algal epiphytes on growingHimanthalia receptacles does not reflect the composition of the surrounding algal vegetation. The possibility that some host-recognition mechanism operates cannot be discounted. Dedicated to Dr. Dr. h. c. Peter Kornmann on the occasion of his eightieth birthday.     

Epiphytism in a subtidal natural bed of Gracilaria gracilis of southwestern Atlantic coast (Chubut, Argentina)

The diversity of epiphytes, their temporal abundance variation and the anatomical structure of host–epiphyte interfaces were studied in the agarophyte Gracilaria gracilis from a natural bed in Bahía Bustamante, Chubut Province, Argentina. Twenty-nine algal species were recorded as epiphytes during 2 years of monthly sampling (March 2006–February 2008). Total epiphyte density ranged between 0.037 ind. cm^?2 (November 2006) and 39.37 ind. cm^?2 (April 2007), with higher density values throughout the second sampling year. Ceramiales species were the most abundant epiphytes. The density of Ceramium rubrum ranged from 0.09 ind. cm^?2 (52 % of the total amount) in September 2006 to 17.4 ind. cm^?2 (44.18 % of the total amount) in April 2007. Epiphyte infection was more dependent on spore recruitment and sporeling development, especially on thalli derived from fragmentation, than on seasonal environmental variations. The different infections were analysed, taking into account the epiphytic attachment strength and invasiveness and the degree of damage inflicted on the host. Calothrix confervicola was one of the most abundant species. This epiphyte, weakly attached to the host surface, generated no host tissue damage. In contrast, C. rubrum, Polysiphonia abscissa and other Ceramiales were the species that caused more damage to the host because their rhizoids penetrated the cortical portion of the host thallus, sometimes reaching the medullary tissue. Some generalisations and characterisations of the different epiphyte groups in relation to their consequences to Gracilaria spp. are presented.     

Effect of epiphytism on reproductive and vegetative lateral formation in the brown, intertidal seaweed Ascophyllum nodosum (Phaeophyceae)

In the brown alga Ascophyllum nodosum (L) Le Jolis, a common species on sheltered Northern temperate rocky shores, gametes are produced in receptacles that emerge from small depressions (lateral pits) along the branched frond. These lateral pits are also the preferred settling site for the obligate epiphyte Polysiphonia lanosa (L) Tandy. Therefore, epiphytism can be expected to interfere with host reproductive output. The present study investigated the potential impact of the epiphyte on A. nodosum in two series of laboratory experiments that measured: (i) the direct shading of the host plant underneath an epiphyte canopy; and (ii) the development of receptacles in clean and epiphytised A. nodosum segments (excised from individual fronds) over a 6 month period. These experiments showed that light reaching emerged fronds underneath a dense epiphyte cover was reduced by 40%, and this was independent of the degree of desiccation the epiphyte experienced. Concurrently, in the growth study with epiphytised A. nodosum segments (segments with one clean and one epiphytised lateral pit) total receptacle biomass per epiphytised fragment was significantly reduced compared with clean segments (0.52 g and 1.25 g per gram of frond segment, respectively), although this effect was only significant in A. nodosum from sheltered shores. However, expressed as biomass per lateral pit, receptacle biomass in the remaining clean lateral pits in epiphytised segments was significantly increased in segments from both shores, demonstrating that A. nodosum can at least partially compensate for the loss of production resulting from epiphytism.     

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.     

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.     

Reti trofiche e flussi di energia nei sistemi a fanerogame marine

Abstract

Food webs and energy flow in seagrass ecosystems. A review on the pathways in the food webs of seagrass ecosystems, both tropical and temperate, with a particular emphasis to Mediterranean Posidonia oceanica meadows is given. Three main pathways of energy transfer from primary producers (host plant and algal epiphytes) were identified: i) the plant itself through photosynthetic tissue; ii) the leaf detritus which in some species forms a litter compartment; iii) the algal epiphytes of leaf blades. The detritus and epiphyte ways are the most common, but they can be differently important according to the season and the spatial patterns of the meadows.     

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.     

THE RED ALGAL EPIPHYTES MICROCLADIA COULTERI AND M. CALIFORNICA (RHODOPHYCEAE,CERAMIACEAE). II. BASIPHYTE SPECIFICITY1

The phenomenon of basiphyte specificity in the settlement and growth of the red algal epiphytes Microcladia californica Farl. and M. coulteri Harv. was examined by studying the interface with their respective basiphytes and by cross-inoculation experiments. Microcladia californica attaches only to the surface of its single basiphyte Egregia menziesii (Aresch.) Turn. whereas M. coulteri penetrates the tissue of a wide range of basiphytes. The pattern of primary rhizoid development in both epiphytes determines the mode of attachment and may influence the range of basiphytes possible for each epiphyte. Cross-inoculation experiments show that Microcladia californica is not able to colonize the basiphytes of M. coulteri, Iridaea and Prionitis, or Ulva. The mechanisms by which these algae restrict the growth of epiphytes include short life-span, “cuticle peeling” and chemical defense. Microcladia coulteri, which naturally colonizes Iridaea and Prionitis, has evolved mechanisms to counteract the antifouling effects of those basiphytes. The question of why Egregia is the exclusive substratum for M. californica remains undetermined. However, Egregia may provide the appropriate ecological conditions and a surface topography conductive to M. californica spore settlement and growth.     

Sponge–seaweed associations in species of Ptilophora (Gelidiaceae, Rhodophyta)

Sponge–seaweed associations in the seaweed genus Ptilophora are poorly understood; therefore, 94 specimens, representing all 17 species of Ptilophora, were examined to detail this phenomenon. All but 2 Ptilophora species were shown to produce surface proliferations, with 13 species found to have sponge associations. Evidence for facultative sponge epiphytism was found with species–specific interactions being unlikely. Results show that surface proliferations are not induced by sponge epiphytes, as they often occur in the absence of sponge epiphytes, and vice versa. The significant number of proliferate thalli found with sponge epiphytes suggests that there is a likely relationship between the presence of surface proliferations and sponge infestation. Sponge epiphytes and Ptilophora species appeared structurally related in that the sponge probably exploits a niche habitat provided by the alga, for which surface proliferations might aid the sponge in bonding to the alga.     

Ceramialean epiphytism in an intertidal Gracilaria chilensis (Rhodophyta) bed in southern Chile

The cultivation of the agarophytic red alga Gracilaria has become an activity of major importance in several parts of the world. However, Gracilaria cultivation in Chile still faces problems such as epiphytism. We report ceramialean epiphytism abundance, recruitment patterns and the microscopic stages fixed on the Gracilaria thalli in an intertidal bed (Metri bay) of southern Chile. Using a factorial field experiment, we analyze the effect of intertidal elevation and the use of epiphyte free inoculum on the abundance of ceramialean epiphytic algae. This evidence is used to provide management and control recommendations for ceramialean epiphytes. The results show a summer recruitment and increase in abundance of ceramialean epiphytes. A seasonal and spatial pattern of abundance of microscopic stages of this epiphytic algae was also found, showing a significant increase from the apical (new tissues) to the central parts of the thalli (older tissues). Apical tissues are free of epiphytic propagules. The epiphyte-free tissue zone decreases significantly from winter to summer. Also, a significantly higher density of microscopic stages of ceramialean epiphytes was found on the thalli collected in the farming are a during the summer at low tidal levels. The experimental results indicated that the abundance of epiphytes was greater at the lower tidal levels than at the higher distribution limits of Gracilaria and it was not possible to control epiphytism by manual cleaning. This revised version was published online in September 2006 with corrections to the Cover Date.     

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.     

Epiphytism,anatomy and regressive evolution in trichomanoid filmy ferns (Hymenophyllaceae)

The few studies on the evolution of epiphytism in ferns have mostly focused on xerophytic and humus‐collecting strategies, neglecting hygrophytes that are abundant in rainforests, such as the trichomanoids (Hymenophyllaceae). Using a phylogenetic approach, we studied the acquisition of epiphytism in this lineage, with the aim of identifying ecological anatomical adaptations and verifying the regressive epiphytic ‘bryophyte‐like’ strategy previously suggested for the group. Inferred evolution of anatomy and morphology, regression and ecology (more particularly colonial epiphytism) were analysed and compared using a maximum likelihood approach. Regressive evolution of anatomy and morphology is revealed in the three clades of colonial epiphytes, probably linked to the selection of water acquisition by blades rather than by regressed roots. However, the ‘bryophyte‐like’ strategy is restricted to some taxa (especially Didymoglossum). Furthermore, a relationship is revealed between large metaxylem and climbing habit. Diversification of colonial epiphytes (and some individual epiphytes) and hemi‐epiphytism would have occurred in the upper Cretaceous and Tertiary, in accordance with the timing of diversification of modern ferns and the evolution of epiphytism in other fern families in the first angiosperm‐dominated forests. This was here performed by selecting hygrophilous strategies that are unique in vascular plants. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 573–593.