Transient Population Dynamics of Two Epiphytic Orchid Species after Hurricane Ivan: Implications for Management

Populations of epiphytic orchids in disturbance‐prone environments rarely reach stable‐stage equilibrium. We characterized the post‐disturbance, transient dynamics of two epiphytic orchids, Broughtonia cubensis, and the leafless Dendrophylax lindenii, comparing the following indices: reactivity/first‐time attenuation, maximal amplification/attenuation, and amplified/attenuated inertia. We also assessed the effects of reintroducing only seeds or only adults, by examining the elasticity of the inertia on the vital rates. For 2006–2010, the stochastic growth rate of D. lindenii was λ_s = 0.94, or a 6 percent decrease per annum. First‐time step attenuation indicates that in 1 year, the population could decrease by an additional 16 percent, and in the worst‐case scenario could decrease by almost half, relative to the stable‐stage distribution, in 10 years. Broughtonia cubensis had a λ_s = 1.03; reactivity and first‐time step attenuation indicates that in 1 year, the population should not change by more than 14 percent of the estimated stable‐stage distribution. However, the worst‐case scenario projected a reduction of 40 percent relative to the stable‐stage distribution within 8 years. A comparison of reintroduction strategies assessed by elasticity of the population momentum showed that adults performed better when relocated to new habitats.     

Analyzing the Spatial Structure of Broughtonia cubensis (Orchidaceae) Populations in the Dry Forests of Guanahacabibes,Cuba

A current key issue in ecology is the role of spatial effects on population and community dynamics. In this paper, we test several hypotheses related to spatial structures and coexistence of epiphytic tropical orchid species with special emphasis on the endemic species Broughtonia cubensis. More specifically, we explored the spatial structure of orchid–host plant communities at three different levels of organization (occupied vs. nonoccupied host trees, trees with B. cubensis vs. other orchids, and reproductive vs. nonreproductive B. cubensis plants). We mapped all potential host trees and orchids at three 20 × 20 m plots and applied techniques of spatial point pattern analysis such as mark connection and mark correlation functions to evaluate departures from randomized communities. We found spatial aggregation of trees with epiphytic orchids and segregation between trees with and without epiphytic orchids, and that there was an intraspecific spatial aggregation of B. cubensis in relation to the other seven epiphytic orchid species. Furthermore, we found spatial aggregation of reproductive B. cubensis individuals and segregation between reproductive and nonreproductive individuals on their phorophytes. Thus, orchid–host plant communities show hierarchical spatial structuring with aggregation and segregation at different levels of organization. Our results point to an enhancement of local species in the coexistence of tropical epiphytic orchid communities, by reducing competition through niche differentiation.     

Distribution,habitat disturbance and pollination of the endangered orchid Broughtonia cubensis (Epidendrae: Laeliinae)

The geographical distribution, population structure and pollination ecology are key aspects in the conservation and management of rare orchids. Here, we address these aspects and the main threats affecting the endangered Cuban orchid Broughtonia cubensis. This rewardless orchid is self‐compatible, but pollinator dependent. However, seed production can be negatively affected by insect‐mediated selfing. Three species of small bee (genera Ceratina and Lasioglossum) act as pollinators. As in the case of other nectarless orchids, we detected two species of plant producing large amounts of nectar in the area, the floral morphology of which closely resembles that of B. cubensis. The simultaneous flowering of these species could positively affect the reproductive success of B. cubensis. Nonetheless, the fitness of this orchid in natural conditions is low, possibly related to strong pollen limitation. To the problems arising from reduced fitness is added the fact that its historical distribution range has been greatly reduced in recent years. Throughout this study, we have detected dramatic reductions in the population sizes, in some cases as a result of human plundering, but also as a consequence of hurricanes. Based on the results of this study, we propose some guidelines to manage and conserve this orchid. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 345–357.     

Population Viability Analysis of the Epiphytic Ghost Orchid (Dendrophylax lindenii) in Cuba

To design feasible conservation and management policies for wild species, it is critical to understand the effects of periodic disturbances, be they natural or anthropogenic. The Caribbean Basin is characterized by high cyclonic activity that has a strong impact on the demography and population dynamics of many taxa, including epiphytic orchids. We conducted a 5‐yr study of rare ghost orchid demography, Dendrophylax lindenii, to assess the stability of a protected population of this species in Cuba. Using both stochastic and deterministic integral projection models, we found that mean annual population growth rates are negative (λ = 0.975). However, we found both population growth rate and extinction risk are highly sensitive to survival rates and reproduction, a difficult to quantify rate for many orchids including our study species. While this species is fairly long‐lived, its relatively slow increase in annual survival with increasing size may reflect the lack of a protected (i.e., subterranean) storage organ—a life‐history trait that may typify other epiphytic species and increase susceptibility to disturbance events. Hurricanes, which are predicted to increase in frequency as a result of climate change, dramatically increase adult mortality. Simulations of these effects indicate that hurricanes and similar disturbances could result in near certain extinction in short time horizons (25 yr) if their annual probability of occurrence exceeds 14 percent. These results suggest a need to better quantify recruitment rates, as well as the sensitivity of population dynamics of this and other orchid species to hurricanes and other periodic disturbances.     

Effects of fire regimen on two species of epiphytic orchids in tropical savannas of the Northern Territory

The effect of four fire regimens (early dry season annual, late dry season annual, early dry season biennial, unburnt) on the numbers of epiphytic orchids in a savanna community in Kakadu National Park was investigated. These fire regimens had been maintained on I ha plots for 16 years. Two species of epiphytic orchid were present — Cymbidium canaliculatum and Dendrobium affine. The numbers of Cymbidium plants were insufficient to enable statistical analysis, but plants were present in the unburnt, early annual and early biennial fire treatments. Dendrobium occurred in substantial numbers irrespective of fire treatment, but the extent of colonization of all available host trees was greater in the unburnt than the burnt treatments. The lower numbers of Dendrobium plants in burnt treatments was due to a lower proportion of potential host trees being colonized and a lower number of orchids on each colonized tree. The proportion of small trees colonized was markedly lower in burnt treatments. Frequent fires late in the dry season may prevent recruitment of the main host species. Eucalyptus tectifica. The survival of orchids despite frequent fires depended on the availability of relatively protected sites for colonization and the ability of the orchids to withstand some fire damage.     

FLORAL ANTHOCYANINS IN SPECIES AND HYBRIDS OF BROUGHTONIA,BRASSAVOLA, AND CATTLEYOPSIS (ORCHIDACEAE)

Chromatographic and spectrophotometric methods were used to identify the floral anthocyanins of Broughtonia domingensis, B. negrilensis, B. sanguinea, Brassotonia ‘John Miller,’ Diabroughtonia, Brassavola nodosa, Cattleyopsis lindenii, and Cattleytonia ‘Rosy Jewell’ for the purpose of establishing chemotaxonomic relationships and inheritance patterns. Flowers of Brassavola nodosa were found to contain raphanusin C, an anthocyanin not present in Cattleyopsis lindenii or in any of the Broughtonia species and hybrids. Pelanin is present in flowers of Broughtonia domingensis and B. negrilensis, but not in those of B. sanguinea. Raphanusin B, raphanusin D, and pelargonidin-3-glucoside, 5-sophoroside which are present in B. sanguinea were also found in its intergeneric hybrids with Diacrium and Brassavola. These findings suggest that Cattleyopsis lindenii should not be transferred to Broughtonia and that B. domingensis, B. negrilensis, and B. sanguinea are separate species.     

Host tree utilization by epiphytic orchids in different land-use intensities in Kathmandu Valley, Nepal

We studied the influence of site conditions on epiphytic orchids under a subtropical climate in the Kathmandu Valley, Nepal. We analysed 96 systematically distributed grid points situated in Kathmandu Valley across a land-use intensity gradient (national park to urbanised city area). Geographical Information System (GIS) and remote sensing were used for classification of land-use types. We identified 23 species of epiphytic orchids, within 13 genera, from 42 different host tree species. Host preference is obvious for some orchid species (e.g., Dendrobium nobile), with certain tree species (e.g., Schima wallichii, Ficus religiosa) hosting more orchid species than others. The orchid Rhynchostylis retusa was the most common species found on many different host tree species across the land-use intensity gradient. Host species and host bark characteristics (e.g., rugosity, pH and exposure to wind) played a vital role for orchid distribution, with lower abundance in areas of higher impact. Under strong human impact (urban city area), F. religiosa was the dominant host tree, with large individual trees (mean diameter in breast height, dbh?=?1.3?m) providing the habitat for considerable populations of R. retusa individuals. In general, epiphytic orchids were found on larger host trees in urban areas than in areas of lower human impact. We found that some hosts are more likely to harbour orchid species, especially native host species. Older larger trees with rougher bark, low pH, exposed to wind and reduced human impact provided better habitats for orchids. We suggest these characteristics should be considered in urban planning to reduce human impact on the associated orchid epiphytic community.     

Effects of Multiple Disturbances (Fire and Hurricane) on Epiphyte Community Dynamics in a Subtropical Forest,Florida, U.S.A.1

We addressed the interacting effects of a natural large–scale fire and a subsequent major hurricane on relative positions of epiphytes in a subtropical forest. In Everglades National Park, subtropical hammocks (hardwood tree “islands”; burned and unburned) during the Ingraham Fire (1989) were surveyed for trees and epiphytic bromeliads (Tillandsia spp.) one year before, as well as one and five years after, Hurricane Andrew (1992). We measured trees (species, diameter, and status [alive/dead]) and epiphytes (species, height, host tree characteristics, substrate life status, and density). The fire decreased the height of epiphytes during the hurricane because branches and bark of trees killed by the fire were unstable epiphyte substrates in the high winds. Proportions of epiphytes on Quercus virginiana were equally increased after the hurricane in both unburned and burned hammocks; the large size and bark characteristics resulted in greater proportional survival of epiphytes on this species. During the five years following the hurricane, changes in the distributions of epiphytes generally were toward pre–hurricane distributions, but recovery was faster in unburned than burned hammocks. We conclude that disturbances that kill trees are likely to amplify the vertical reduction of epiphytes during a subsequent hurricane and that effects of a single disturbance on plant populations can be influenced by the disturbance history of the system, including different types of disturbances.     

Exodermis structure controls fungal invasion in the leafless epiphytic orchid Dendrophylax lindenii (Lindl.) Benth. ex Rolfe

Leafless and shootless epiphytic orchids rely essentially on CAM photosynthesis in roots for carbon gain. However, it is believed that a proportion of carbon is obtained by endomycorrhizal associations. In this study, we show that Dendrophylax lindenii possesses a dimorphic exodermis with smaller, thin-walled passage cells that are depleted in flavonoids. No hyphae succeeded in penetrating into the cortex from a non-passage cell, but 20% of the hyphae in contact with a passage cell managed to penetrate into the cortex. The passage cells represent 40% of the amount of cells in the centre of the side that touches the substrate, but no passage cells are observed in the upper side of the root. This distribution and density of exodermal passage cells define a strategy for controlling the extent and location of fungal invasion in the orchid root. This strategy provides a mechanism for restricting fungal growth to the lower cortex and thus maximising carbon gain from photosynthesis while enabling further trophic exchanges from mycorrhizal associations.     

The impact of life form on the architecture of orchid mycorrhizal networks in tropical forest

Understanding the processes that determine the architecture of interaction networks represents a major challenge in ecology and evolutionary biology. One of the most important interactions involving plants is the interaction between plants and mycorrhizal fungi. While there is a mounting body of research that has studied the architecture of plant–fungus interaction networks, less is known about the potential factors that drive network architecture. In this study, we described the architecture of the network of interactions between mycorrhizal fungi and 44 orchid species that represented different life forms and co‐occurred in tropical forest and assessed the relative importance of ecological, evolutionary and co‐evolutionary mechanisms determining network architecture. We found 87 different fungal operational taxonomic units (OTUs), most of which were members of the Tulasnellaceae. Most orchid species associated with multiple fungi simultaneously, indicating that extreme host selectivity was rare. However, an increasing specificity towards Tulasnellaceae fungal associates from terrestrial to epiphytic and lithophytic orchids was observed. The network of interactions showed an association pattern that was significantly modular (M = 0.7389, M_random = 0.6998) and nested (NODF = 5.53, p < 0.05). Terrestrial orchids had almost no links to modules containing epiphytic or lithophytic orchids, while modules containing epiphytic orchids also contained lithophytic orchids. Within each life form several modules were observed, suggesting that the processes that organize orchid–fungus interactions are independent of life form. The overall phylogenetic signal for both partners in the interaction network was very weak. Overall, these results indicate that tropical orchids associate with a wide number of mycorrhizal fungi and that ecological rather than phylogenetic constraints determine network architecture.     

Composition and ecology of vascular epiphyte communities along an altitudinal gradient in central Veracruz,Mexico

Abstract. Vascular epiphytes were studied in forests at altitudes from 720 to 2370 m on the Atlantic slope of central Veracruz, Mexico. The biomass of all trees of each species > 10 cm diameter at breast height within plots between 625 and 1500 m² was estimated. The number of species per plot ranged between 22 and 53, and biomass between 9 and 249 g dry weight/m². The highest values, both of species and biomass, were found at an intermediate altitude (1430 m). Habitat diversity may contribute to epiphyte diversity in humid forests, but the importance of this effect could not be distinguished from the influence of climate. A remarkably high number of bromeliads and orchids grew in relatively dry forests at low altitudes. In wet upper montane forests, bromeliads were replaced by ferns, while orchids were numerous at all sites, except for a pine forest. The number of epiphytic species and their biomass on a tree of a given site were closely related to tree size. According to Canonical Correspondence Analysis, the factor determining the composition of the epiphytic vegetation of a tree was altitude and to some extent tree size, whereas tree species had practically no influence. The only trees which had an evidently negative effect on epiphytes were pines, which were particularly hostile to orchids and to a lesser degree to ferns, and Bursera simaruba, which generally had few epiphytes due to its smooth and defoliating bark.     

Genetic inference of epiphytic orchid colonization; it may only take one

Colonization of vacant habitat is a fundamental ecological process that affects the ability of species to persist and undergo range modifications in continually shifting landscapes. Thus, understanding factors that affect and limit colonization has important ecological and conservation implications. Epiphytic orchids are increasingly threatened by various factors, including anthropogenic habitat disturbance. As cleared areas (e.g. pastures) are recolonized by suitable host trees, the establishment and genetic composition of epiphytic orchid populations are likely a function of their colonization patterns. We used genetic analyses to infer the prevailing colonization pattern of the epiphytic orchid, Brassavola nodosa. Samples from three populations (i.e. individuals within a tree) from each of five pastures in the dry forest of Costa Rica were genotyped with neutral nuclear and chloroplast markers. Spatial autocorrelation and hierarchical genetic structure analyses were used to assess the relatedness of individuals within populations, among populations within pastures and among populations in different pastures. The results showed significant relatedness within populations (mean r = 0.166) and significant but lower relatedness among populations within a pasture (mean r = 0.058). Our data suggest that colonization of available habitats is by few individuals with subsequent population expansion resulting from in situ reproduction, and that individuals within a tree are not a random sample of the regional seed pool. Furthermore, populations within a pasture were likely colonized by seeds produced by founders of a neighbouring population within that pasture. These results have important ramifications for understanding conservation measures needed for this species and other epiphytic orchids.     

Host associations of the strangler fig Ficus watkinsiana in a subtropical Queensland rain forest

Abstract The distribution of epiphytic organisms is limited by the availability of, and dispersal to suitable hosts. We examined the distribution of a hemi‐epiphytic strangler fig, Ficus watkinsiana (Moraceae) in Cooloola National Park (Queensland, Australia), in order to determine whether this species exhibits a preference for certain host species and why. We assessed host bark roughness and flakiness, fruit type, and size to explain the observed distribution of F. watkinsiana. We surveyed over 1900 potential host trees of the 30 most common forest canopy species and found that host size measured by diameter at breast height accounted for most variation in fig prevalence (Binary Logistic Regression log‐likelihood = ?588.178, G = 314.494, d.f. = 1, P < 0.005). After controlling for host size, F. watkinsiana prevalence still differed significantly between host species (χ² = 54.612, d.f. = 24, P < 0.005), a difference that was only partly explained by variation in the bark roughness of host trunks. These results suggest that variation in the rate at which tree species host strangler figs are primarily related to individual tree size – figs may simply be more likely to colonize and thrive upon host species that grow larger.     

西双版纳片段化石灰岩森林附生兰科植物多样性研究

西双版纳石灰岩地区拥有丰富的兰科植物资源,但近年来随着橡胶树的大面积种植,使得该地区很多石灰岩森林呈片段化。为了解片段化石灰岩森林中附生兰科植物多样性状况以及片段化对附生兰科植物的影响,该研究选取了生境片段化的青岩寨和曼纳览,以及连续生境的绿石林和巴卡新寨等4个样地的29个样方进行多样性调查和对比研究。结果表明:4个石灰岩森林样共记录到附生兰科植物34属76种1 528株(丛)。通过对坡向、坡度、海拔、地形、郁闭度和森林类型等6个环境因子与附生兰科植物丰富度进行CCA分析,发现石灰岩地区附生兰科植物的分布主要受海拔和森林类型2个因素的影响,在海拔较高的青岩寨和巴卡新寨附生兰科植物物种丰富度高于低海拔的绿石林和曼纳览,而片段化对附生兰科植物的物种多样性和多度均无显著影响,这可能与片段化的历史较短有关。虽然目前来看生境片段化对石灰岩地区的附生兰科植物多样性无显著影响,但生境脆弱的石灰岩森林植被的保护对于兰科植物多样性保护则更具重要性。     

Vertical distribution of Hymenophyllaceae species among host tree microhabitats in a temperate rain forest in Southern Chile

Question: Are differences in microhabitat preferences of co‐occurring epiphytic Hymenophyllaceae species (filmy ferns) correlated with differences in ecophysiological responses to light availability and humidity in the host tree? Location: The Andean foothills in south‐central Chile. Methods: We evaluated the distribution pattern of nine filmy fern species in microhabitats that differ in light availability and humidity in four host tree species. A DCA was developed to assess Hymenophyllaceae species microhabitat preference in terms of canopy openness (CO) and relative humidity. We assessed whether differences in chlorophyll content, maximum photochemical efficiency (Fv/Fm), photosynthetic capacity (A_max), evapotranspiration (E) and instantaneous water use efficiency (WUE) are consistent with any pattern. Results: CO and relative humidity differed significantly with height in the host trees. While CO increased with height in a host tree, relative humidity decreased. DCA analysis showed that filmy fern species distribution within and among trees was mainly explained by the relative humidity of the microhabitat. Chlorophyll content, chlorophyll a/b ratio, A_max and E differed significantly among filmy fern species. A_max and E were correlated with axis 1 scores from the DCA analysis. Conclusions: The vertical distribution and abundance of filmy fern species in Chilean temperate rain forest seems to be closely related to the different microhabitats offered by host trees. This pattern may reflect interspecific differences in ecophysiological traits related both to light availability and humidity. Our results suggest that humidity is the main environmental factor driving functional responses and habitat preferences of these filmy fern species.     

Bromeliad‐associated Reductions in Host Herbivory: Do Epiphytic Bromeliads Act as Commensalists or Mutualists?

Many members of the family Bromeliacae are able to adopt epiphytic lifestyles and colonize trees throughout the Neotropics. Bromeliacae do not extract nutrients from their hosts and confer relatively minor costs on their host plants. We suggest that bromeliads, however, may benefit their hosts by providing habitat for predators of host plant herbivores. We report a correlation between bromeliad presence and a reduction in herbivore damage in orange trees, an effect that is increased when bromeliads are colonized by ants. Our results may have important implications for agricultural systems in the Neotropics, where bromeliads are often removed in the belief they are parasitic. We instead demonstrate that bromeliads may impart a benefit to their hosts, and speculate that under particular circumstances they may be part of a three‐species mutualism.     

The effects of hurricanes on the stochastic population growth of the endemic epiphytic orchid Broughtonia cubensis living in Cuba

We carried out a posthurricane evaluation of Broughtonia cubensis (Lindl.) Cogn., an endemic Cuban epiphytic orchid, after Hurricane Ivan (2004). We studied the transient responses in the stochastic dynamics of the species at three different sites over 13 successive years (2006–2019), monitored plot inventories (464 individuals in 10 transects) and built stochastic population models. The deterministic stochastic growth rate values () did not significantly differ (F = 2.76; p > 0.076) among the three sites over the 2006–2019 period. The long-term stochastic growth rate was 0.973 [0.932, 1.034]. The matrix elements that had the largest effect on were the transition to and stasis within the largest size class. Transient responses explained an average of 86% of the variation in the observed population growth rates , compared to 4% of the variation in the vital rates . Because transient dynamics are dependent on the population size composition, we ran extinction risk analyses under two scenarios: a population composed mainly of juveniles and another composed mainly of adults. There was little risk of falling below the quasi-extinction threshold before 25 year for both juveniles and adults. However, the risk of quasi-extinction was almost certain for both size classes by 80 year. We also simulated the effect of increasing the hurricane occurrence probability over 80 year on the population. There was little risk of extinction before 20 year in the baseline model, but there was a significant risk of extinction within 5 year when 90% of the individuals were affected by a new hurricane event.     

Resurgence of cucurbit downy mildew in the United States: Insights from comparative genomic analysis of Pseudoperonospora cubensis

Pseudoperonospora cubensis, the causal agent of cucurbit downy mildew (CDM), is known to exhibit host specialization. The virulence of different isolates of the pathogen can be classified into pathotypes based on their compatibility with a differential set composed of specific cucurbit host types. However, the genetic basis of host specialization within P. cubensis is not yet known. Total genomic DNA extracted from nine isolates of P. cubensis collected from 2008 to 2013 from diverse cucurbit host types (Cucumis sativus, C. melo var. reticulatus, Cucurbita maxima, C. moschata, C. pepo, and Citrullus lanatus) in the United States were subjected to whole‐genome sequencing. Comparative analysis of these nine genomes confirmed the presence of two distinct evolutionary lineages (lineages I and II) of P. cubensis. Many fixed polymorphisms separated lineage I comprising isolates from Cucurbita pepo, C. moschata, and Citrullus lanatus from lineage II comprising isolates from Cucumis spp. and Cucurbita maxima. Phenotypic characterization showed that lineage II isolates were of the A1 mating type and belonged to pathotypes 1 and 3 that were not known to be present in the United States prior to the resurgence of CDM in 2004. The association of lineage II isolates with the new pathotypes and a lack of genetic diversity among these isolates suggest that lineage II of P. cubensis is associated with the resurgence of CDM on cucumber in the United States.     

Endophytic fungi assemblages from 10 <Emphasis Type="Italic">Dendrobium</Emphasis> medicinal plants (Orchidaceae)

Dendrobium is the largest genus of tropical epiphytic orchid, some of which are traditional Chinese medicinal plants. The therapeutic components varied significantly among species. Endophytic microbes (fungi) hidden in medicinal plants may play an important effect on the overall quality of herb. Investigation of fungal composition in host plants is the first step toward elucidating the relationship endophyte-therapeutic content of herbal medicine. In this study, 401 culturable fungal endophytes were isolated and identified from 10 species of medicinal Dendrobium based on morphological and molecular techniques. The results showed that endophytic fungi from Dendrobium plants exhibited high biodiversity (37 genera, about 80 species). Acremonium, Alternaria, Ampelomyces, Bionectria, Cladosporium, Colletotrichum, Fusarium, Verticillium and Xylaria were the dominant fungal endophytes. Tropical epiphytic orchids appear to vary in degree of host specificity in their endophytic fungi.     

Gas exchange of terrestrial and epiphytic orchids from Brazilian Atlantic Rainforest

Leaf gas exchange of terrestrial and epiphytic orchids from the Atlantic Rainforest in northeast Brazil was investigated under artificial growth conditions. The terrestrial orchids showed higher values of all photosynthetic parameters in comparison to epiphytic ones. There was a close relationship between P _N and g _s for both terrestrial and epiphytic orchids. Taken together, our results demonstrated that the photosynthetic parameters were related to the specific growth habits of the orchids under study.