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.     

GENETIC VARIATION IN THE EPIPHYTES TILLANDSIA IONANTHA AND T. RECURVATA (BROMELIACEAE)

The genetic structure of two epiphytic species, Tillandsia ionantha and T. recurvata, was investigated using enzyme electrophoresis. Electrophoretic data suggest that T. ionantha and T. recurvata differ in breeding system, in agreement with predictions based on their strikingly different floral morphologies. Electrophoretic data suggest extremely high levels of inbreeding for T. recurvata, whereas Tillandsia ionantha exhibits characteristics of an outcrossing species. Values of P, H, and mean number of alleles per locus are much higher in T. ionantha than in T. recurvata. The mean value of F_IS for T. ionantha is low (0.056), closely approaching expectations at Hardy-Weinberg equilibrium. In contrast, the mean value of F_IS in T. recurvata (1.000) indicates a complete absence of heterozygotes. The two species also differ in genetic structure. Low F_ST values for T. ionantha indicate little variation in allele frequencies among populations. In contrast, F_ST values are high for T. recurvata, suggesting substantial genetic heterogeneity among populations. In addition, the mean value of I is higher in T. ionantha (0.995) than in T. recurvata (0.931). Population genetic data are in agreement with the suggestion of Benzing (1978), who proposed that extreme epiphytes such as T. recurvata, would be characterized by increased autogamy ensuring high seed set. Due to high chromosome numbers in Bromeliaceae (most taxa have x = 25), the family has been considered polyploid. However, with the exception of an additional isozyme for PGM in T. recurvata, the two species are isozymically diploid.     

Is Allelopathic Activity of Ipomoea murucoides Induced by Xylophage Damage?

Herbivory activates the synthesis of allelochemicals that can mediate plant-plant interactions. There is an inverse relationship between the activity of xylophages and the abundance of epiphytes on Ipomoea murucoides. Xylophagy may modify the branch chemical constitution, which also affects the liberation of allelochemicals with defense and allelopathic properties. We evaluated the bark chemical content and the effect of extracts from branches subjected to treatments of exclusion, mechanical damage and the presence/absence of epiphytes, on the seed germination of the epiphyte Tillandsia recurvata. Principal component analysis showed that branches without any treatment separate from branches subjected to treatments; damaged and excluded branches had similar chemical content but we found no evidence to relate intentional damage with allelopathy; however 1-hexadecanol, a defense volatile compound correlated positively with principal component (PC) 1. The chemical constitution of branches subject to exclusion plus damage or plus epiphytes was similar among them. PC2 indicated that palmitic acid (allelopathic compound) and squalene, a triterpene that attracts herbivore enemies, correlated positively with the inhibition of seed germination of T. recurvata. Inhibition of seed germination of T. recurvata was mainly correlated with the increment of palmitic acid and this compound reached higher concentrations in excluded branches treatments. Then, it is likely that the allelopathic response of I. murucoides would increase to the damage (shade, load) that may be caused by a high load of epiphytes than to damage caused by the xylophages.     

A nurse plant benefits from facilitative interactions through mycorrhizae

• Plant facilitation promotes coexistence by maintaining differences in the regeneration niche because some nurse species recruit under arid conditions, whereas facilitated species recruit under more mesic conditions. In one Mexican community, 95% of species recruit through facilitation; Mimosa luisana being a keystone nurse for many of them. M. luisana individuals manifest greater fitness when growing in association with their facilitated plants than when growing in isolation. This observation suggests that nurses also benefit from their facilitated plants, a benefit thought to be mediated by mycorrhizal fungi.
• Under field conditions, we experimentally tested whether mycorrhizal fungi mediate the increased fitness that M. luisana experiences when growing in association with its facilitated plants. We applied fungicide to the soil for nurse plants growing alone and growing in association with their facilitated plants in order to reduce the mycorrhizal colonisation of roots. We then assessed the quantity and quality of seed production of M. luisana in four treatments (isolated‐control, isolated‐fungicide, associated‐control and associated‐fungicide).
• Fungicide application reduced the percentage root length colonised by mycorrhizae and reduced fitness of M. luisana when growing in association with their facilitated plants but not when growing in isolation. This reduction was reflected in the total number of seeds, number of seeds per pod, seed mass and seed viability.
• These results suggest that nurses benefit from the presence of their facilitated plants through links established by mycorrhizae, indicating that both plants and belowground mutualistic communities are all part of one system, coexisting by means of intrinsically linked interactions.

     

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.     

Interactions between epiphytes during canopy soil formation: an experiment in a lower montane cloud forest of southeast Mexico

• In several montane forests around the world, epiphytes coexist in mats, sharing the rhizosphere and forming histosol-type soils rich in nutrients. The role of these epiphytes in the formation of canopy soil and the fitness costs that epiphytes face when cohabiting in these mats are unknown.
• In a lower montane cloud forest in central Veracruz, Mexico, a 2-year factorial experiment was carried out with the presence/absence of ramets of Phlebodium areolatum (Polypodiaceae), Tillandsia kirchhoffiana, T. multicaulis and T. punctulata (Bromeliaceae). We examined (i) which epiphyte species contribute to the formation of canopy soil, (ii) the role of epiphyte composition in the soil nutrient composition, and (iii) the fitness costs faced by epiphytes when cohabiting.
• Canopy soil formation highest when P. areolatum is present. Soil nutrient content does not change with epiphyte composition, is influenced by the microbiota, and P content decreases with the presence of epiphytes. The fitness costs show that the species compete, decreasing their survival and growth, but the competitive capacity differs between the species.
• We conclude that P. areolatum is an ecosystem engineer that promotes the creation of canopy soil but is a poor competitor. The results coincide with the model of succession by facilitation. Canopy soil is a slow-created component whose nutrient content does not depend on the epiphytic flora. In epiphyte mats, the dominant interactions are competitive, but there is also facilitation.

     

The nutrient status of epiphytes and their host trees along an elevational gradient in Costa Rica

Vascular epiphytes are a conspicuous and highly diverse group in tropical wet forests; yet, we understand little about their mineral nutrition across sites. In this study, we examined the mineral nutrition of three dominant vascular epiphyte groups: ferns, orchids, and bromeliads, and their host trees from samples collected along a 2600 m elevational gradient in the tropical wet forests of Costa Rica. We predicted that the mineral nutrition of ferns, orchids, and bromeliads would differ because of their putative differences in nutrient acquisition mechanisms and nutrient sources—atmospherically dependent, foliar feeding bromeliads would have lower nitrogen (N) and phosphorous (P) concentrations and more depleted δ¹⁵N values than those in canopy soil-rooted ferns because canopy soil is higher in available N, and more enriched in δ¹⁵N than the atmospheric sources of precipitation and throughfall. We also predicted that epiphyte foliar chemistry would mirror that of host trees because of the likely contribution of host trees to the nutrient cycle of epiphytes via foliar leaching and litter contributions to canopy soil. In the same vein, we predicted that epiphyte and host tree foliar chemistry would vary with elevation reflecting ecosystem-level nutrients—soil N availability increases and P availability decreases with increasing elevation. Our results confirmed that canopy soil-rooted epiphytes had higher N concentrations than atmospheric epiphytes; however, our predictions were not confirmed with respect to P which did not vary among groups indicating fixed P availability within sites. In addition, foliar δ¹⁵N values did not match our prediction in that canopy soil-rooted as well as atmospheric epiphytes had variable signatures. Discriminant function analysis (DFA) on foliar measurements determined that ferns, orchids, and bromeliads are statistically distinct in mineral nutrition. We also found that P concentrations of ferns and orchids, but not bromeliads, were significantly correlated with those of host trees indicating a possible link in their mineral nutrition’s via canopy soil. Interestingly, we did not find any patterns of epiphyte foliar chemistry with elevation. These data indicate that the mineral nutrition of the studied epiphyte groups are distinct and highly variable within sites and the diverse uptake mechanisms of these epiphyte groups enhance resource partitioning which may be a mechanism for species richness maintenance in tropical forest canopies.     

Effects of epiphytic lichens on host preference of the vascular epiphyte Tillandsia usneoides

We investigated the potential for nonvascular epiphytic species (primarily lichens) to affect the quality of different host tree species for the vascular epiphyte Tillandsia usneoides in the southeastern USA. Different host tree species had substantially different abundances of Tillandsia , and these abundances were correlated with the composition of nonvascular epiphyte communities. In greenhouse experiments Tillandsia grew significantly faster on the branches of Quercus virginiana (a species with very high natural abundances of Tillandsia ) when the dominant lichen on Q. virginiana was left intact than when the lichen was removed from the branches. In laboratory experiments, extracts from Cryptothecia rubrocincta , a lichen that was 10 times more common on poor host species for Tillandsia than on good host species, reduced Tillandsia seedling survival and growth in comparison to extracts from other species and rainwater. In field experiments, lichens increased the proportion of Tillandsia seeds and vegetative strands that adhered to the trunk of Ilex opaca (a poor Tillandsia host), but lichens did not affect propagule adherence to Q. virginiana . Our results are by no means exhaustive of the possibilities, but they suggest that the structure and diversity of vascular and nonvascular epiphytic communities that grow in different tree species may not be simply the product of host tree characteristics, but may also be influenced by interactions among the epiphytes themselves.     

Disentangling the roles of chance,abiotic factors and biotic interactions among epiphytic bryophyte communities in a tropical rainforest (Yunnan,China)

• Epiphytes offer an appealing framework to disentangle the contributions of chance, biotic and abiotic drivers of species distributions. In the context of the stress-gradient theory, we test the hypotheses that (i) deterministic (i.e., non-random) factors play an increasing role in communities from young to old trees, (ii) negative biotic interactions increase on older trees and towards the tree base, and (iii) positive interactions show the reverse pattern.
• Bryophyte species distributions and abiotic conditions were recorded on a 1.1 ha tropical rainforest canopy crane site. We analysed co-occurrence patterns in a niche modelling framework to disentangle the roles of chance, abiotic factors and putative biotic interactions among species pairs.
• 76% of species pairs resulted from chance. Abiotic factors explained 78% of non-randomly associated species pairs, and co-occurrences prevailed over non-coincidences in the remaining species pairs. Positive and negative interactions mostly involved species pairs from the same versus different communities (mosses versus liverworts) and life forms, respectively. There was an increase in randomly associated pairs from large to small trees. No increase in negative interactions from young to old trees or from the canopy to the base was observed.
• Our results suggest that epiphytic bryophyte community composition is primarily driven by environmental filtering, whose importance increases with niche complexity and diversity. Biotic interactions play a secondary role, with a very marginal contribution of competitive exclusion. Biotic interactions vary among communities (mosses versus liverworts) and life forms, facilitation prevailing among species from the same community and life form, and competition among species from different communities and life forms.

     

Temperature and water requirements for germination and effects of discontinuous hydration on germinated seed survival in Tillandsia recurvata L.

Tillandsia recurvata is an epiphytic bromeliad with a wide distribution in the Americas; however, little is known about the development of its post-seminal adaptations for survival in epiphytic environments. The purpose of this study was to define the temperature and water requirements for the germination of T. recurvata seeds. The absence of radicle emergence in T. recurvata seeds resulted in 2?stages of germination: swollen with broken seed coat (stage-1) and chlorophyllic embryos (stage-2). The effects of partial or discontinuous hydration on germinated seed survival were also assessed. The seeds were collected in a semi-arid shrubland of Mexico City. We explored: (1) whether water vapour can provide a sufficient water source for germination; (2) the temperature required for germination stage-1 and the optimal and critical osmotic potentials for germination in both germination stages; (3) the effect of seed incubation at different osmotic potentials that undergo subsequent dehydration on their survival in stage-2; and (4) the loss of dehydration tolerance during early post-seminal development. In addition, an image of T. recurvata seed anatomy was obtained to illustrate its structures. Germination stage-1 of T. recurvata seeds is rather similar across the tested temperature range. The seeds required to be in contact with liquid water to germinate. The interval of osmotic potential facilitating both germination stages was from 0 to ?0.6?MPa. Although germinated seeds displayed dehydration tolerance, this tolerance decreased in germination stage-2. The osmotic potential during germination affected the tolerance of the chlorophyllic embryos (stage-2) to subsequent dehydration.     

Epiphytic bromeliads in a changing world: the effect of elevated CO2 and varying water supply on growth and nutrient relations

• Global climate change is likely to impact all plant life. Vascular epiphytes represent a life form that may be affected more than any other by possible changes in precipitation leading to water shortage, but negative effects of drought may be mitigated through increasing levels of atmospheric CO₂.
• We studied the response of three epiphytic Aechmea species to different CO₂ and watering levels in a full‐factorial climate chamber study over 100 days. All species use crassulacean acid metabolism (CAM). Response variables were relative growth rate (RGR), nocturnal acidification and foliar nutrient levels (N, P, K, Mg).
• Both elevated CO₂ and increased water supply stimulated RGR, but the interaction of the two factors was not significant. Nocturnal acidification was not affected by these factors, indicating that the increase in growth in these CAM species was due to higher assimilation in the light. Mass‐based foliar nutrient contents were consistently lower under elevated CO₂, but most differences disappeared when expressed on an area basis.
• Compared to previous studies with epiphytes, in which doubling of CO₂ increased RGR, on average, by only 14%, these Aechmea species showed a relatively strong growth stimulation of up to +61%. Consistent with earlier findings with other bromeliads, elevated CO₂ did not mitigate the effect of water shortage.

     

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

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

Microsatellites in the bromeliads Tillandsia fasciculata and Guzmania monostachya

We developed microsatellites for the bromeliads Tillandsia fasciculata and Guzmania monostachya, epiphytes of Central‐American montane rain forests. Fragments obtained using amplified fragment length polymorphism (AFLP) pre‐amplification were enriched by hybridization with biotin‐labelled repeat sequences, and subsequently cloned and sequenced. Primers were designed for 15 loci; seven produced interpretable and repeatable amplification products and five were polymorphic in one or both species. We will use these primers to study the population structure of Bromeliaceae in relation to their breeding system as well as to their role in (re‐)colonization of secondary rain forests.     

External water transport is more important than vascular transport in the extreme atmospheric epiphyte Tillandsia usneoides (Spanish moss)

Most epiphytic bromeliads, especially those in the genus Tillandsia, lack functional roots and rely on the absorption of water and nutrients by large, multicellular trichomes on the epidermal surfaces of leaves and stems. Another important function of these structures is the spread of water over the epidermal surface by capillary action between trichome “wings” and epidermal surface. Although critical for the ultimate absorption by these plants, understanding of this function of trichomes is primarily based on light microscope observations. To better understand this phenomenon, the distribution of water was followed by its attenuation of cold neutrons following application of H₂O to the cut end of Tillandsia usneoides shoots. Experiments confirmed the spread of added water on the external surfaces of this “atmospheric” epiphyte. In a morphologically and physiologically similar plant lacking epidermal trichomes, water added to the cut end of a shoot clearly moved via its internal xylem and not on its epidermis. Thus, in T. usneoides, water moves primarily by capillarity among the overlapping trichomes forming a dense indumentum on shoot surfaces, while internal vascular water movement is less likely. T. usneoides, occupying xeric microhabitats, benefits from reduction of water losses by low‐shoot xylem hydraulic conductivities.     

Microgeographic variation in recruitment under adult trees: arrival of new genotypes or perpetuation of the existing ones?

• Investigating spatial variation in the relative importance of sexual reproduction and clonal propagation is critical to obtain more accurate estimates of future effective population sizes and genetic diversity, as well as to identify ecological correlates of clonality.
• We combined a stratified sampling scheme with microsatellite genetic analyses to estimate variation in the proportion of sexual versus clonal recruits among saplings in five populations of the tree Pyrus bourgaeana. Using a likelihood framework, we identified clones among the genotypes analysed and examined variation among populations regarding the proportion of saplings coming from clonal propagation. We also examined the relationship between the relative abundance of clonal shoots across the studied populations and their herbivory levels.
• Our results revealed that one third of the saplings examined (N = 225 saplings) had a probability above 0.9 of being clones of nearby (<10 m) trees, with the ratio between clonal propagation and sexual recruitment varying up to eight‐fold among populations. A small portion of these putative clonal shoots reached sexual maturity. Relative abundance of clonal shoots did not significantly relate to the herbivory by ungulates.
• Our results call into question optimistic expectations of previous studies reporting sufficient levels of recruitment under parental trees without animal seed dispersal services. Nevertheless, given that some of these clonal shoots reach sexual maturity, clonal propagation can ultimately facilitate the long‐term persistence of populations during adverse periods (e.g. environmental stress, impoverished pollinator communities, seed dispersal limitation).

     

Behavioural responses to predation may explain shifts in community structure

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Ecological longevity of Polaskia chende (Cactaceae) seeds in the soil seed bank,seedling emergence and survival

• Soil seed banks are essential elements of plant population dynamics, enabling species to maintain genetic variability, withstand periods of adversity and persist over time, including for cactus species. However knowledge of the soil seed bank in cacti is scanty. In this study, over a 5‐year period we studied the seed bank dynamics, seedling emergence and nurse plant facilitation of Polaskia chende, an endemic columnar cactus of central Mexico.
• P. chende seeds were collected for a wild population in Puebla, Mexico. Freshly collected seeds were sown at 25 °C and 12‐h photoperiod under white light, far‐red light and darkness. The collected seeds were divided in two lots, the first was stored in the laboratory and the second was use to bury seeds in open areas and beneath a shrub canopy. Seeds were exhumed periodically over 5 years. At the same time seeds were sown in open areas and beneath shrub canopies; seedling emergence and survival were recorded over different periods of time for 5 years.
• The species forms long‐term persistent soil seed banks. The timing of seedling emergence via germination in the field was regulated by interaction between light, temperature and soil moisture. Seeds entered secondary dormancy at specific times according to the expression of environmental factors, demonstrating irregular dormancy cycling.
• Seedling survival of P. chende was improved under Acacia constricta nurse plants. Finally, plant facilitation affected the soil seed bank dynamics as it promoted the formation of a soil seed bank, but not its persistence.

     

FOLIAR TRICHOMES OF PLEUROTHALLIDINAE (ORCHIDACEAE): FUNCTIONAL SIGNIFICANCE

Foliar surfaces of nine pleurothallid orchids (Cryptophoranthus lepidotus L. O. Wms., Octomeria sp., Pleurothallis pidax Luer, P. poeppigii Lindl., P. revoluta (Ruiz & Pav.) Garay, Restrepiella ophiocephala (Lindl.) Garay & Dunsterv., Restrepia muscifera Rchb. f. ex Lindl., Scaphosepalum rapax Luer, Stelis endresii Rchb. f.), four nonpleurothallid orchids (Cyrtopodium punctatum (L.) Lindl., Encyclia cochleata (L.) Lemee, E. tampensis (Lindl.) Small, Paphiope-dilum hybrid) and five tillandsioid bromeliads (Catopsis nutans (Sw.) Griseb., Tillandsia fas-ciculata Sw., T. streptophylla Scheidw., T. stricta Soland., T. tectorum E. Morr.) were assayed for permeability to Ca, S and P ions. Compared to leaves of the atmospheric bromeliads, those of the orchids proved less permeable to Ca and S. Moisture exchange profiles were determined for two of the bromeliads and three orchids; bromeliad leaves were able to rehydrate completely, orchid leaves only partially. Absorptive capacities of trichomes borne by five taxa were tested by autoradiographic analysis. Unlike bromeliad trichomes, those of the orchids exhibited no capacity to accumulate ³H-leucine. These results and other considerations described here suggest that, unlike tillandsioid bromeliads, neither the pleurothallid nor the nonpleurothallid orchids examined in this survey rely heavily on shoots for mineral and moisture procurement.     

It is about time: genetic variation in the timing of leaf‐litter inputs influences aquatic ecosystems

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Positive co‐occurrence between feeding‐associative savannah fishes depends on species and habitat

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