Tillandsia Species Research Articles

Generous-like flowers: nectar production in two epiphytic bromeliads and a meta-analysis of removal effects.

Animal-pollinated angiosperm plants that respond positively to nectar removal by replenishment invest energy that can entail a reproductive cost. We investigated whether or not nectar removal stimulates replenishment in two hummingbird-pollinated bromeliad species. Nectar replenishment rates were also assessed by removing nectar from manually pollinated flowers because pollination events might be used as signals to save energy by preventing allocation to post-pollination nectar production. Then we synthesized the current understanding of nectar removal effects by reviewing existing published studies with a meta-analysis. The magnitude and significance of estimated nectar removal effects and factors associated with variation in size and direction of nectar removal effects were elucidated with the meta-analysis. We found that both Tillandsia species strongly respond to repeated nectar removal by producing >3 times additional nectar. Nectar secretion patterns were not altered by pollination (stigmatic pollen deposition) and we found no evidence of nectar reabsorption. Although the effect size varied widely across systems and/or environmental conditions, the meta-analysis showed that nectar removal had overall a positive effect on nectar replenishment (mainly among species inhabiting wet tropical habitats such as Tillandsia), and a negative effect on the secretion of additional sugar, suggesting that those plants are resource limited and conservative in the secretion of additional sugar.

Andean bear use of the epiphytic bromeliad Tillandsia fendleri at Quebrada el Molino, Venezuela

Epiphytic bromeliads of the genus Tillandsia have been reported as important in the Andean (spectacled) bear diet throughout the bear’s distribution (Mondolfi 1971, 1989; Peyton 1980; Jorgenson and Rodriguez 1986; Rodriguez et al. 1986; Suarez 1989; Goldstein 1990, Rodriguez 1991; Eulert 1995). Epiphytic bromeliads are locally abundant in Andean forests (Gentry and Dodson 1987). Because they are high in soluble carbohydrates, fat, and protein, bromeliads are a nutritious food for bears (Goldstein 1990). Bears often leave sign of their activities in areas where they live; trails, scats, broken branches, claw marks on trees, and beds are common in bear habitats (Burst and Pelton 1983). Andean bears are no exception, leaving abundant sign especially while feeding on epiphytic and terrestrial bromeliads (Peyton 1980, Suarez 1989). In Venezuela, sign of feeding on epiphytic bromeliads by Andean bears has been found in mountain forests between 1,680 and 3,200 m and were the most abundant bear sign found in forests from 2,400 to 2,800 m (Goldstein 1990). Spectacled bears usually produce 2 types of signs at epiphytic bromeliad feeding sites: claw marks on tree bark and piles of bromeliad leaves on the forest floor. Claw marks are visible on tree bark from 40–60 cm above the ground to the branches where the bear fed on the bromeliads. The length and depth of the scars depends on the bark of the tree species. Bears feed on the basal meristematic part of the bromeliads, causing the leaves to drop to the forest floor. Bears have been found to eat 1–10 or more plants at each tree, and Tillandsia fendleri plants have around 50 leaves. Depending on the number of bromeliads eaten, leaves can form a conspicuous carpet on the forest floor. Moreover, usually the central bunch of younger leaves stick together, with bite marks on the white basal meristematic part (Goldstein 1990). Although there are several species of epiphytic bromeliads Tillandsia available in forests between 2,400–3,000 m at Quebrada El Molino (T. compacta, T. complanata, T. tetrantha, T. spiculosa), Andean bears at Quebrada El Molino have been found to only eat the largest species, T. fendleri. They also feed on the fruit of a Lauraceae tree (Beilschmiedia sulcata). In paramo areas, bears feed on the terrestrial bromeliad Puya aristeguietae (Goldstein 1990). T. fendleri plants can reach 2 m in height (including the single central erect inflorescence) and 1 m in diameter (Smith 1971), making them very conspicuous and easily distinguished from smaller Tillandsia species. During work at Quebrada El Molino (Goldstein 1990, 2002), I found that Andean bear T. fendleri feeding sites were mostly located in tall, emergent trees with large canopies and in large trees at the forest–paramo edge, where the abundance of T. fendleri plants was higher (bromeliads/tree x 1⁄4 16.95 [SD 1⁄4 10.89]; diameter at breast height [dbh] x1⁄4 51.84 cm [SD 1⁄4 22.80 cm]; tree height x 1⁄4 10.03 m [SD 1⁄4 3.10]; n 1⁄4 51). Because climbing is energetically costly and the distribution and abundance of bromeliads is clumped, I predicted that Andean bears would maximize expected reward for the effort of climbing trees with T. fendleri, preferentially using trees with the greatest bromeliad loads.

Tissue culture enhances the propagation potential of some Tillandsioideae

Seeds of three Tillandsia species were germinated in vitro and multiple shoots produced within 13 months in culture. Different Murashige and Skoog media; full- strength, half-strength, and half-strength macro/full micro salts plus 25 mg l − 1 NaFeEDTA, were used for germination. For multiple shoots the best germination medium was used with different plant growth regulators added (0.5/0.5 mg l- 1 or 1.0/0.5 mg l- 1 BA/NAA).

Size‐Dependent Variation of Carbon and Nitrogen Isotope Abundances in Epiphytic Bromeliads

Abstract: While atmospheric species of bromeliads have narrow leaves, densely covered with water‐absorbing trichomes throughout their life cycles, many tank bromeliads with broad leaves, forming phytotelmata, go through an atmospheric juvenile phase. The effect of the different habits and the phase change in tank‐forming bromeliads on water and nutrient relations was investigated by analysing the relationship between plant size, C/N ratios and the natural abundance of 13C and 15N in five epiphytic bromeliad species or morphospecies of a humid montane forest in Xalapa, Mexico. The atmospheric species Tillandsia juncea and T. butzii exhibited full crassulacean acid metabolism, with δ13C values (mean ‐ 15.3 ‰ and ‐ 14.7 ‰, respectively) independent of size. In Tillandsia species with C3 photosynthesis, δ13C decreased with increasing plant size, indicating stronger drought stress in juveniles. The increase of the C/N ratio with size suggests that, at least in heteroblastic bromeliads, the availability of water is more limiting during early growth, and that limitations of nitrogen supply become more important later on, when water stored in the tank helps to bridge dry periods, reducing water shortage. δ15N values of the two atmospheric species were very negative (‐ 12.6 ‰ and ‐ 12.2 ‰, respectively) and did not change with plant size. Tank‐forming bromeliads had less negative δ15N values (c ‐ 6 ‰), and, in species with atmospheric juveniles and tank‐forming adults, δ15N values increased significantly with plant size. These differences do not appear to be an effect of the isotopic composition of N sources, but rather reflect N availability and limitation and stress‐induced changes in 15N discrimination.

A New Species of Tillandsia (Bromeliaceae) from the Mexican Yucatan Peninsula

A new species of Tillandsia is pro- posed. It is similar to Tillandsia brachycaulos Schlechtendal and Tillandsia balbisiana Schultes f., two sympatric species. The new species is most similar to T. brachycaulos in flower and bract color, but the elongated inflorescences resemble those of T. balbisiana, although the branches are much shorter.

INTERSPECIFIC AND INTERGENERIC FERTILIZATION BARRIERS IN BROMELIACEAE

Bromeliaceae are tropical ornamentals with a variety of colours and inflorescence shapes. To remain the interest of the public, breeders create new cultivars using interspecific and intergeneric hand-pollinations. In our breeding studies, 4 genera were used: Aechmea (2 species), Guzmania (3 species), Tillandsia (1 species) and Vriesea (3 species). In preparation of crossing experiments, two important flower characteristics were studied: pollen quality and stigma receptivity. Pollen germination of different flower stages was determined by an in vitro germination test. All species, except two of the Guzmania species, possessed viable pollen and germination was always highest at anthesis. Stigma receptivity was studied by pollinating flowers before, at and after anthesis. Pollen germination and pollentube growth on stigma were visualised with fluorescence microscopy (aniline blue staining). All species showed optimal stigma receptivity and ovule penetration at anthesis. Interspecific and intergeneric hand-pollinations were performed with pollen and stigma at anthesis stage. Pollen germination on stigma, pollentube growth in style and ovary and fertilization of ovules was evaluated using fluorescence microscopy. Five species were self-compatible, making emasculation necessary. Half of the interspecific and all of the intergeneric crosses were classified as incongruent because no ovules were fertilised (prefertilization barriers) or no seedlings developed (postfertilization barriers). Prefertilization barriers occurred in the style, mostly the lower part of the style. Only in crosses with the self-incompatible Aechmea species as mother or the extremely short-styled Tillandsia species as father, pollentube growth stopped in the upper part of the style. To understand more about these style barriers, the morphology of the pistil was studied. Styles of all species consisted of a hollow channel, covered with protein-rich cells. These ‘feeder’-cells also appeared on the placenta and the ovule funiculus. In this way, pollen tubes could be guided towards the micropyles. Further, Ca-oxalate and starch grains were observed in the style.

Mutagenesis and in vitro culture of Tillandsia fasciculata Swartz var. fasciculata (Bromeliaceae)

The genus Tillandsia (Bromeliaceae) has very few variegated species, and cultivars with chlorophyll-deficient variegation are especially rare. With the objective of inducing chlorophyll-deficient leaf variegation, seeds of Tillandsia fasciculata var. fasciculata were treated with gamma radiation, combined gamma and thermal neutron radiation or by chemical mutagenesis with ethyl methanesulfonate (EMS). Wild type, albino, yellow, yellowish-green and variegated phenotypes were obtained in the subsequent M 0 generation. These variegated seedlings were either sectorial or mericlinal chimeras, consequently the variegation of these seedlings was lost as they grew older. Gamma radiation at 21 kR and 27 kR produced the highest percentage of variegated seedlings (4.4%). The highest percentage of seedlings with chlorophyll-deficient leaves was 8.4% with 27 kR gamma radiation, and 15.8% with 1.2% EMS. Radiation and chemical mutagenesis caused chlorophyll-deficiency mutations in one or more of the histogenic layers: LI, LII, LIII. Wild types had greater total chlorophyll a, b and total chlorophyll than mutant phenotypes. Most of the yellow and yellowish-green seedlings multiplied in a solid half-strength MS medium with equimolar 0.3 or 0.5 μM BA and IBA. The yellowish-green seedlings were able to grow photoautotrophically while the yellow ones were not. This is one of the first reports on the mutagenesis of a Tillandsia species. Stable periclinal chlorophyll-deficient chimeras of Tillandsia species can likely be obtained via mutagenesis if large numbers of seeds are treated with a suitable mutagen.

The Use of 6-Benzylaminopurine for Rapid Multiplication of Tillandsias

Some species of the genus Tillandsia (Bromeliaceae) are highly endangered in their natural habitats. This report introduces a method for induced lateral shoot development in atmospheric Tillandsias using the cytokinin BAP. Spraying or dipping the plants with BAP-solutions induced many lateral shoots. Even difficult-to-propagate species like T. cacticola L.B.SM developed up to 30 times more usable shoots than the control. Applying low concentrations (1 and 5 mg·L-1) over a longer period of time (10 weeks, 3 times a week) produced results similar to those using higher concentrations (50 mg·L-1) for shorter periods (2 weeks, 3 times a week). Chemical name used: 6-benzylaminopurine (BAP).

Cytological aspects of the hypocotyl correlated to the behavior of the embryo radicle ofTillandsiaatmospheric species

SUMMARYGenerally in Bromeliaceae, as in all monocotyledons, upon germination of the seed, the embryo radicle grows very little. In the atmospheric species of Tillandsia, not even this minimal growth occurs. In the adult stage, these species use special hairs as their sole absorbing organ. The present work analyses the morphological and cytochemical events that are responsible for the lack of growth in the embryo radicle. Ultrastructural analysis and specific stainings for light microscopy highlighted a special area (that we have called the «constriction zone») in the median region of the hypocotyl-root axis of embryos of T. flabellata Back. and T. schiedeana Steud., both atmospheric species. During the final phase of seed maturation, the cells of the aforesaid zone thickened their walls by means of materials that, at least in part, consisted of acid polysaccharydes and neutral lipids. These materials were derived from the fused contents of characteristic cytoplasmic vesicles. Due to the presence of hydrop...

Effects of drought stress on CO 2 exchange and water relations in the CAM epiphyte Tillandsia utriculata (Bromeliaceae)

Summary Water was withheld from plants of Tillandsia utriculata L. for up to two months in a growth chamber. Nocturnal CO2 uptake declined substantially in the first week without water, then declined slowly between 7 and 31 d of the drought treatment. Net CO2 exchange after two months of desiccation was nearly zero, although substantial nocturnal increases in malic acid concentrations were still measureable. Thus, the degree of CO2 recycled internally via CAM increased dramatically throughout the drought treatment. In spite of the slow, yet substantial declines in nighttime CO2 uptake observed throughout the drought treatment, leaf water potentials remained relatively high (above −1.5 MPa) and declined by only 0.5 MPa after two months without water. Based on pressure-volume analyses, the leaf bulk elastic modulus was approximately 3.3 MPa, a very low value indicative of highly elastic cell walls. Thus, maintenance of high turgor pressures as a result of elastic cell walls may constitute an important adaptation of this species and presumably other epiphytic species of Tillandsia that reduces the negative effects of drought stress. This, as well as other adaptations to drought stress such as the preferential utilization of water from water-storage parenchyma, osmotic adjustment, and low rates of transpiration, should contribute to survival during drought in these unusual plants.

Leaf Anatomy and CO<sub>2</sub> Recycling during Crassulacean Acid Metabolism in Twelve Epiphytic Species of Tillandsia (Bromeliaceae)

The relationship between leaf anatomy, specifically the percent of leaf volume occupied by waterstorage parenchyma (hydrenchyma), and the contribution of respiratory CO₂ during Crassulacean acid metabolism (CAM) was investigated in 12 epiphytic species of Tillandsia. It has been postulated that the hydrenchyma, which contributes to CO₂ exchange through respiration only, may be causally related to the recently observed phenomenon of CO₂ recycling during CAM. Among the 12 species of Tillandsia, leaves of T. usneoides and T. bergeri exhibited 0% hydrenchyma, while the hydrenchyma in the other species ranged from 2.9% to 53% of leaf cross-sectional area. Diurnal malate fluctuation and nighttime atmospheric CO₂ uptake were measured in at least four individuals of each species. A significant excess of diurnal malate fluctuation as compared with atmospheric CO₂ absorbed overnight was observed only in T. schiedeana. This species had an intermediate proportion (30%) of hydrenchyma in its leaves. Results of this study do not support the hypothesis that CO₂ recycling during CAM may reflect respiratory contributions of CO₂ from the tissue hydrenchyma.

Host Tree Utilization by Vascular Epiphytes in a Seasonally Inundated Forest (Tintal) in Mexico

We studied fifteen species of vascular epiphytes in a 1200 m 2 sample of seasonally inundated forest (tintal) near Cancun, Mexico. The community of epiphytes occupied 14 species of host tree. While certain species of host tree tended to accommodate more epiphytes than others, no host tree specificity was exhibited by any of the epiphyte species. There was some evidence of canopy partitioning by branch size among the mote common epiphytes (8 species with >10 individuals in the sample area). Four species of Tillandsia (Bromeliaceae) tended to occupy the smallest diameter supports (mostly 2.5 cm (...)

Role of the Nitrogen-Fixing Bacterial Microflora in the Epiphytism of Tillandsia (Bromeliaceae)

Nitrogen-fixing activity in the phyllosphere of 12 species of Tillandsia from different Mexican habitats was evaluated by the acetylene reduction assay, and nitrogen-fixing microorganisms were isolated and characterized. The leaves from eight of the 12 Tillandsia species examined exhibited nitrogenase activity in enrichment cultures. Among the microorganisms implicated—Agrobacterium, Bacillus, Erwinia, Pseudomonas, Rahnella, Vibrio, and Xanthomonas—only Bacillus megatherium reduced acetylene in pure culture. Our findings suggest that nitrogen fixation in the phyllosphere of the sampled epiphytes occurs under suitable conditions and that most of the bacteria involved are primarily soil and water inhabitants. The results also suggest a relationship between the composition of the nitrogen-fixing microbial communities grown on the leaf and the different development of the leaf area in Tillandsia due to the aerial components (wings) of the trichomes.

ROLE OF THE NITROGEN‐FIXING BACTERIAL MICROFLORA IN THE EPIPHYTISM OF TILLANDSIA (BROMELIACEAE)

Nitrogen‐fixing activity in the phyllosphere of 12 species of Tillandsia from different Mexican habitats was evaluated by the acetylene reduction assay, and nitrogen‐fixing microorganisms were isolated and characterized. The leaves from eight of the 12 Tillandsia species examined exhibited nitrogenase activity in enrichment cultures. Among the microorganisms implicated—Agrobacterium, Bacillus, Erwinia, Pseudomonas, Rahnella, Vibrio, and Xanthomonas—only Bacillus megatherium reduced acetylene in pure culture. Our findings suggest that nitrogen fixation in the phyllosphere of the sampled epiphytes occurs under suitable conditions and that most of the bacteria involved are primarily soil and water inhabitants. The results also suggest a relationship between the composition of the nitrogen‐fixing microbial communities grown on the leaf and the different development of the leaf area in Tillandsia due to the aerial components (wings) of the trichomes.

Comparative biology of Neotropical epiphytic and saxicolous Tillandsia species: population structure

ABSTRACTIn comparing the population structure of epiphytic and saxicolous Tillandsia species I found no significant differences between polycarpic species. Epiphytic T. ionochroma populations had a higher seedling:adult ratio than saxicolous T. ionochroma populations, but the difference was not statistically significant. Tillandsia ultriculala, a monocarpic epiphyte, had a significantly higher seedling:adult ratio than the polycarpic species. T. utriculata's higher ratio may result from its greater seed production and lack of vegetative propagation. Unlike the polycarpic species, all adult-size rosettes grow from seeds in T. utriculata. Tillandsia sphaerocephala, an obligate saxicole, occurred in larger clusters (i.e. more rosettes per genet) than T. ionochroma. The increased tendency for vegetative reproduction in T. sphaerocephala suggests that spatial patchiness is less of a constraint than with epiphytes.

Observations on some Mexican species of the genus Tillandsia L. (Bromeliaceae)

Summary On Mexican soil, during a scientific expedition the aim of which was to investigate the biology of Bromeliads, the authors gathered samples of six species of Tillandsia genus. From these saples the authors made taxonomic profiles integrated with the information derived from odservations made in the field regarding: the relationship between the epiphyte and the host; the cohabitation among the epiphytes and the microenvironment of each species. Original drawings accompany the text.

Contribution to the knowledge of the Mexican species of the genus «Tillandsia» (Bromeliaceae)

Summary In monographic cards, we have put together taxonomic, ecologic and phenologic informations of four species of Tillandsia from Mexico: T. ionantha, T. macdougallii, T. schiedeana, T. seleriana. Considerations are added of biological nature connected by the different water avalaibilities of the environment and observations on the autogamic mechanism in the genus Tillandsia.

OBSERVATIONS ON THE MICROBODIES IN THE GENUS TILLANDSIA (BROMELIACEAE)

Organelles morphologically similar to microbodies have been found in several tissues of atmospheric species of Tillandsia from different habitats. The presence of catalase was demonstrated by the DAB reaction thus confirming the microbody nature of these organelles. They are a feature of the Tillandsia species with normal photosynthetic carbon fixation and with CAM. Their size is consistently small.The nucleoid observed in the microbodies shows a characteristic morphology which has not been reported before within other plant microbodies. This nucleoid is composed of minute tubular structures, for which the authors here propose a three‐dimensional arrangement.

Anatomic and Ultrastructural Study of the Septal Nectary in SomeTillandsia(Bromeliaceae) Species

SUMMARYThe anatomy of the septal nectary and the ultrastructural features of the secretory mechanism of several species of Tillandsia are examined. The results are discussed with special reference to the particular ecology of this epiphitic genus. Although the basic anatomy of the secretory epithelium and of the underlying tissue in the nectary shows no outstanding differences from what reported by previous authors as regards other Bromeliaceae, a new fact is worth mentioning: nectariferous tissue is found also in the receptacle although the ovary of Tillandsia is tipically hy- pogynous. The ultrastructural morphology of the epithelium in flower buds near the anthesis and in open flowers in two successive moments of the anthesis indicates that the epithelial cells play an active role in nectar secretion, and that sugars are discharged to the outside mainly by means of a granulocrine process. The post-secretory stage is cytologically characterized by a consistent vacuolization of the cytoplasm and by the thickening of the wall. Therefore the ultrastructure of the septal nectary of Tillandsia shows no significant differences from the Bromeliaceae investigated by other authors. There is, however, a particular aspect in Tillandsia worth underlining, that is the mechanism by which the occlusion of the nectariferous slits is attained once the epithelium has become inactive. This mechanism does not only involve the approaching of the epithelium tangential cell walls, which at this stage are considerably thick, but it is due mainly to the production of a cementing substance made of acid polysaccharides.

Ursachen der Verbreitungsmuster einiger Bromeliaceae in Rio Grande do Sul (Südbrasilien)

Summary The areas of some Bromeliaceae from Rio Grande do Sul are described and discussed in relation to the topography of this state and to the there existing vegetation types. By comparation of the total range of some species and with help of palaeogeographical data it was possible to suggest the time of immigration of various genera and species. The genus Dyckia is the oldest element of Bromeliaceae which invaded in this southern area. The Bromelioideae with berries are the most recent elements of this family which came to Rio Grande do Sul. For the Tillandsioideae nine andine elements could be prooved. The immigration of Bromeliaceae during Holocene went through the coastal gate in NE of the country and in NW in the region of Alto Uruguai. All these species don t show any speciation in Rio Grande do Sul. They invaded for various distances into the araucarian forest. The genus Vriesea and the East-Brazilian representants of Tillandsia came with the evergreen tropical rain forest through the coastal gate. The Paraguayan immigration is documentated by three species of Tillandsia. The independent development of CAM-mecanism is coupled in terrestrial species with succulent leaves and in epiphytic species with a special design of leafe surface, respectively with the movableness of the trichomes.