El género Tillandsia (Bromeliaceae) en el departamento de Lambayeque, Perú / The genus Tillandsia (Bromeliaceae) in the department of Lambayeque, Peru

Resumen Se presentan 26 taxones de Tillandsia (Bromeliaceae) que ocurren en el departamento de Lambayeque, Perú, de los cuales 12 se registran por primera vez, ocupando distintas ecoregiones y formaciones vegetales, desde el tillandsial de poca elevación hasta la Jalca. Se presentan claves dicotómicas para la determinación de los subgéneros, especies y variedades y además se proporcionan datos actualizados de distribución geográfica, rango altitudinal, usos y de conservación.

Abstract There are 26 taxa of Tillandsia (Bromeliaceae) that occur in the department of Lambayeque, Peru, of which 12 are registered for the first time, occupying different ecoregions and plant formations, from the low tillandsial to the Jalca. Dichotomous keys for the determination of subgenera, species and varieties are presented and updated data on geographical distribution, altitudinal range, uses and conservation are also provided.

The Tillandsia genus: history, uses, chemistry, and biological activity / El género Tillandsia: historia, usos, química y actividad biológica

Tillandsia L. genus comprises 649 species, with different uses at different times. T. usneoides L. uses are reported since the late- archaic and pre-Columbian cultures. In XIX-XX centuries, T. usneoides was used in some manufactured products, as polish and packing fruit. Tillandsia has a favorable reputation as medicine: for leucorrhea, rheumatism, ulcers, hemorrhoid treatment, as an anti-diabetic remedy, emetic, analgesic, purgative, contraceptive, antispasmodic and diuretic. Tillandsia chemical composition includes cycloartane triterpenes and hydroxy-flavonoids, which are present in at least 24 species. Several extracts and compounds from Tillandsia spp. have been reported with pharmacological actions, as anti-neoplasia, hypolipidemic, antifungal, anti-HSV-1, hypoglycemic and microbicide. This review communicates the economic importance, ethnobotany, chemistry composition and biological activities of the Tillandsia genus, and analyze its biological and economic perspective. Tillandsia genus has cultural, economic and pharmacological relevance, with a high potential in many essential aspects of the modern society.

El género Tillandsia L. comprende 649 especies, con diferentes usos en diferentes épocas. T. usneoides L. se han reportado desde el arcáico tardío hasta las culturas precolombinas. En los siglos XIX-XX, T. usneoides se usó en productos manufacturados: como abrasivo y embalaje de fruta. Como medicina tradicional, el género Tillandsia se reporta para leucorrea, reumatismo, úlceras, hemorroides, remedio antidiabético, emético, analgésico, purgante, anticonceptivo, antiespasmódico y diurético. Su composición química incluye triterpenos de tipo ciclo-artano e hidroxi-flavonoides, presentes en al menos 24 especies. Los extractos y compuestos del género Tillandsia se han reportado con propiedades antineoplásicas, hipolipidémicas, antifúngicas, anti-HSV-1, hipoglucemiantes y microbicidas. Esta revisión comunica la importancia económica, etnobotánica, composición química y las actividades biológicas del género Tillandsia, y analiza su perspectiva biológica y potencial económica. Tillandsia tiene importancia cultural, económica y farmacológica, con gran potencial en muchos aspectos esenciales de la sociedad moderna.

Vertebrate florivory of vascular epiphytes: the case of a bromeliad / Florivores vertebrados de epífitas vasculares: o caso de uma bromélia

Abstract The avoidance of vertebrate herbivory is thought to be one of the possible drivers for the evolution of epiphytism. Scarce literature suggests that epiphyte herbivory is mainly related to insect attack on reproductive structures. In a pine-oak forest we observed almost all inflorescences of an epiphytic bromeliad (Tillandsia carlos-hankii) with signs of florivory; the degree of damage suggested that vertebrate herbivores could be involved. To assess the intensity of vertebrate florivory damage we recorded the percentage of damaged individuals in a 500 m2 plots during two flowering seasons. To identify possible vertebrate herbivores, we installed 20 mixed capture stations, 10 photo-traps focused on bromeliads and analyzed stomach contents of captured vertebrates. Florivory was observed on 62% of individuals during the first flowering season and 77% on the second; and average one individual lost 41% of reproductive structures. Vertebrates associated with florivory were a bird, Icterus bullockii (Aves, Passeriformes, Icteridae), a squirrel Sciurus aureogaster (Mammalia, Rodentia, Sciuridae), and mice, Peromyscus gratus, P. levipes and P. aztecus (Mammalia, Rodentia, Cricetidae). Our results suggest that vascular epiphytes are used as opportunistic resources for small vertebrates during seasons when preferred resources are scarce.

Resumo Acredita-se que a prevenção da herbivoria dos vertebrados é um dos possíveis impulsores da evolução da epífita. A literatura escassa sugere que a herbivora em epífitas está relacionada principalmente ao ataque de insetos as estruturas reprodutivas. Em uma floresta de pinheiros observamos que quase da todas as inflorescências de uma bromélia epífita (Tillandsia carlos-hankii) apresentavam sinais de florivoria; o grau de danos sugeria que herbívoros majores (vertebrados) pudessem estar envolvidos. Para avaliar a intensidade do dano de florivoria de vertebrados, registramos a porcentagem de indivíduos danificados em quadrantes de 500 m2 durante duas estações de florescimento. Para identificar possíveis herbívoros vertebrados, instalamos 20 estações de captura mista e analisamos o conteúdo estomacal de vertebrados capturados. Além disso, foram instaladas 10 foto-armadilhas focadas em bromélias. A florivoria foi observada em 62% dos indivíduos durante a primeira estação de floração e 77% na segunda. Os vertebrados associados à florivoria foram pássaro, Icterus bullockii (Aves, Passeriformes, Icteridae), um esquilo Sciurus aureogaster (Mammalia, Rodentia, Sciuridae) e ratos, Peromyscus gratus, P. levipes e P. aztecus (Mammalia, Rodentia, Cricetidae). Assim, nossos resultados sugerem que epífitas vasculares são usadas como recurso facultativo para estes animais durante as estações, quando os recursos preferidos estão escassos.

Vertebrate florivory of vascular epiphytes: the case of a bromeliad

Abstract The avoidance of vertebrate herbivory is thought to be one of the possible drivers for the evolution of epiphytism. Scarce literature suggests that epiphyte herbivory is mainly related to insect attack on reproductive structures. In a pine-oak forest we observed almost all inflorescences of an epiphytic bromeliad (Tillandsia carlos-hankii) with signs of florivory; the degree of damage suggested that vertebrate herbivores could be involved. To assess the intensity of vertebrate florivory damage we recorded the percentage of damaged individuals in a 500 m2 plots during two flowering seasons. To identify possible vertebrate herbivores, we installed 20 mixed capture stations, 10 photo-traps focused on bromeliads and analyzed stomach contents of captured vertebrates. Florivory was observed on 62% of individuals during the first flowering season and 77% on the second; and average one individual lost 41% of reproductive structures. Vertebrates associated with florivory were a bird, Icterus bullockii (Aves, Passeriformes, Icteridae), a squirrel Sciurus aureogaster (Mammalia, Rodentia, Sciuridae), and mice, Peromyscus gratus, P. levipes and P. aztecus (Mammalia, Rodentia, Cricetidae). Our results suggest that vascular epiphytes are used as opportunistic resources for small vertebrates during seasons when preferred resources are scarce.

Resumo Acredita-se que a prevenção da herbivoria dos vertebrados é um dos possíveis impulsores da evolução da epífita. A literatura escassa sugere que a herbivora em epífitas está relacionada principalmente ao ataque de insetos as estruturas reprodutivas. Em uma floresta de pinheiros observamos que quase da todas as inflorescências de uma bromélia epífita (Tillandsia carlos-hankii) apresentavam sinais de florivoria; o grau de danos sugeria que herbívoros majores (vertebrados) pudessem estar envolvidos. Para avaliar a intensidade do dano de florivoria de vertebrados, registramos a porcentagem de indivíduos danificados em quadrantes de 500 m2 durante duas estações de florescimento. Para identificar possíveis herbívoros vertebrados, instalamos 20 estações de captura mista e analisamos o conteúdo estomacal de vertebrados capturados. Além disso, foram instaladas 10 foto-armadilhas focadas em bromélias. A florivoria foi observada em 62% dos indivíduos durante a primeira estação de floração e 77% na segunda. Os vertebrados associados à florivoria foram pássaro, Icterus bullockii (Aves, Passeriformes, Icteridae), um esquilo Sciurus aureogaster (Mammalia, Rodentia, Sciuridae) e ratos, Peromyscus gratus, P. levipes e P. aztecus (Mammalia, Rodentia, Cricetidae). Assim, nossos resultados sugerem que epífitas vasculares são usadas como recurso facultativo para estes animais durante as estações, quando os recursos preferidos estão escassos.

Unearthing the Medicinal Properties of Tillandsia recurvata (Ball Moss): A Mini Review.

Background: More than half of traditional medicines are of natural origin and research has shown that these are associated with fewer side effects than the synthetic ones, since less than 10% of the 350,000 identified plant species have been exposed to some amount of bioactive screens, it is now the aim of researchers to screen more plants and also identify the active ingredients responsible for detected bioactivities. These we believe may provide the foundation for identifying new drug leads that may prove useful against chronic lifestyle diseases. This review takes a look at the work that has currently been conducted on Tillandsia recurvata commonly known as ball moss which is believed to assist with future research. Results: The chloroform, methanol and water extracts of ball moss have shown cytotoxicity against several human cancer cell lines and the methanol extract induces apoptosis in some. Further the chloroform extract was shown to reduce angiogenesis and the methanol extract inhibited particular kinases (CSNK2A2, MEK5, GAK, FLT and DRAK1) of which MEK5 and GAK have been implicated in prostate cancer. The same extract was further shown to display promising anti-diabetic properties via a reduction in fasting blood glucose (P<0.05), fructosamine levels (P<0.05), serum CRP and insulin levels when compared to the control mice. Phytochemical screens identified a novel glycoside and several cycloartanes and dicinnamtes; 1,3-di-O-Cinnamoyl-glycerol and (E)-3-(cinnamoyloxy)-2-hydroxypropyl 3-(3,4-dimethoxyphenyl)acrylate. Further bioactive screens on these isolates showed that cycloartane-3,24,25-triol reduced the viability of prostate PC-3 and DU145 cell lines. This isolate was further shown to inhibit MRCKα kinase implicated in the initiation and progression of prostate cancer. Conclusions: This review confirms the promising efficacy of the T. recurvata plant and so its worth for further research which may prove useful in the pharmaceutical and nutraceutical industries. Such benefits have already begun with the introduction of the alpha prostate formula, now on the market for improved prostate health.

Cycloartanes with Anticancer Activity Demonstrate Promising inhibition of the Mrckα and Mrckβ Kinases.

Aim: The role of Kinases in cancer onset and progression has made kinases a target for the control of some cancers. Recent discoveries that kinases are most effectively inhibited by small molecules have also resulted in an increased search for small molecule kinase inhibitors. Cycloartanes are small molecules found in many medicinal plants including the Jamaican Ball Moss (Tillandsia recurvata). Recent studies on T. recurvata have demonstrated that it possesses anticancer activity. Cycloartane-3,24,25- triol, an analog of a cycloartane identified in Ball moss was also shown to have inhibitory activity against MRCKα kinase. This study was as such set up to determine the MRCKα/β kinase inhibition activity of other cycloartanes in Ball Moss and their analogs. MRCKα/β kinases has been identified as an important kinase implicated in cancer onset and progression and as such a potential drug target. Methodology: Kinase inhibition activity of 6 cycloartanes was investigated using the ligand-kinase binding assay. The WST-1 reagent assay was also used to determine the antiproliferation activity of the cycloartanes against some prostate and breast cancer cell lines. Results: Cycloart-23-ene-3,25-diol (1), Cycloartane-3,24,25-triol (2), Cycloart-25-ene- 3,24-diol (3), 3,23-Dioxo-9,19-cyclolanost-24-en-26-oic acid (4), 24,25- Dihydroxycycloartan-3-one (5) inhibited the MRCKα kinase with Kd of 0.21 μM, 0.25μM, 0.36 μM, 3.0 μM and 2.1 μM respectively. Hydroxycycloart-23-en-3-one,25, (6) showed no inhibition against the MRCKα kinase. Compounds 1, 3, 4, 5 inhibited the MRCKβ kinase with Kd of 4.7 μM, 1.10 μM, 3.2 μM and 9.8 μM, respectively. Three of the six cycloartanes exhibited antiproliferation activity against two prostate and breast cancer cell lines each. Conclusion: Cycloart-23-ene-3,25-diol (1) showed the most promising activity against the MRCKα/β kinase out of the 6 cycloartanes screened demonstrating an interesting structure activity relationship profile when compared with the other molecules. Cycloart- 23-ene-3,25-diol (1) deserves further studies to determine its in vivo activity as well.

Tillandsia recurvata L. (Bromeliaceae): aspectos farmacognósticos / Tillandsia recurvata L. (Bromeliaceae): a pharmacognostic study

A cobertura superior das florestas tropicais, formada pelas copas das árvores, constitui um ambiente de extrema diversidade vegetal. Contribuindo com a grande riqueza dessas florestas encontram-se as espécies botânicas epífitas, cuja importância pode ser observada do ponto de vista ecológico, faunístico, etnobotânico e até mesmo farmacológico. Dentre as espécies adaptadas à vida epifítica podemos citar as bromeliáceas, que compõem uma das famílias mais representativas dessa flora com elevada variabilidade genética. Tillandsia recurvata é uma espécie epífita cosmopolita, nativa, adaptada a habitats áridos, utilizada para o tratamento de diversas doenças, mas pouco se tem publicado a respeito das comprovações científicas de suas propriedades. O presente trabalho revisa os aspectos etnofarmacológicos, atividades biológicas e compostos químicos relacionados à espécie em questão, pertencente à Bromeliaceae. A partir do levantamento de dados realizado, observa-se que esta se trata de uma espécie de conhecido uso popular no tratamento de diferentes distúrbios, com alguns estudos farmacológicos que comprovam suas propriedades terapêuticas. Do ponto de vista químico observa-se a presença de terpenos, flavonoides e derivados cinâmicos em sua constituição, em que alguns compostos isolados ilustram a potencialidade desta espécie como fonte de biomoléculas de interesse. Todos estes aspectos considerados nesta revisão contribuem para o conhecimento a respeito da importância biológica e o potencial terapêutico acerca dos metabólitos desta promissora espécie.

The canopy of tropical forests, formed by the crowns of trees, constitutes a habitat of extreme plant diversity. Contributing to the great richness of these forests are epiphytes, which are of outstanding importance from the ecological, faunal, ethnobotanical and even pharmacological standpoints. Among the species adapted to the epiphytic life (growing on other plants) are the bromeliads, one of the most representative families of that flora, possessing high genetic variability. Tillandsia recurvate is a cosmopolitan epiphyte adapted to arid habitats, traditionally used to treat various diseases, though little has been published on scientific evidence for its properties. This paper reviews its ethnopharmacology and the biological activities and chemical compounds related to this member of the Bromeliaceae. According to the data collected, this species is known for its popular use in the treatment of various disorders, with some pharmacological studies that demonstrate its therapeutic properties. From the chemical point of view, terpenes, flavonoids and cinnamic acid derivatives are found in its constitution and some of the compounds isolated illustrate the potential of this species as a source of interesting biomolecules. All these features reviewed here contribute to what is known about the biological importance and therapeutic potential of metabolites from this promising species.

In vitro and In vivo anti-cancer effects of Tillandsia recurvata (Ball Moss) from Jamaica / Efectos Anticancerosos In vitro e In vivo de la Tillandsia recurvata (Bola de Musgo) de Jamaica

OBJECTIVE: Tillandsia recurvata, also commonly known as Ball Moss, is endemic to Jamaica and some parts of the Caribbean and South America. The plant, despite being reported to be used in folk medicine, had not previously been evaluated for its anti-cancer potential. The aim of this study was to evaluate the anti-cancer activity of Ball Moss. METHODS: The anti-proliferation activity of the crude methanolic extract of the T recurvata was evaluated in vitro in five different histogenic cancer cell lines (prostate cancer - PC-3, breast cancer, Kaposi sarcoma, B-16 melanoma and a B-cell lymphoma from a transgenic mouse strain) using the trypan blue assay. The crude extract was also evaluated in vivo in tumour-bearing mice. Immunohistochemistry staining with Apoptag was used for histology and determination of apoptosis. RESULTS: The crude methanolic extract of T recurvata demonstrated anti-proliferation activity against all the cell lines, killing > 50% of the cells at a concentration of 2.5 µg/ml. Kaposi sarcoma xenograft tumours were inhibited by up to 75% compared to control in the in vivo study (p < 0.05). There was evidence of DNA fragmentation and a decrease in cell viability on histological studies. The methanolic extract showed no toxic effect in the mice at a dose of 200 mg/kg. CONCLUSIONS: Our data suggest that T recurvata has great potential as an anti-cancer agent and that one of its mechanisms of cell kill and tumour inhibition is by the induction of apoptosis.

OBJETIVO: La Tillandsia recurvata, también conocida como bola de musgo, es endémica en Jamaica, así como en algunas partes del Caribe y América del sur. Si bien se había reportado su uso como parte de la medicina popular, esta planta no había sido evaluada previamente en relación con su potencial para la lucha contra el cáncer. El objetivo de este estudio fue evaluar la actividad anticancerígena de la bola de musgo. MÉTODOS: La actividad antiproliferativa del extracto metanólico crudo de la T recurvata, fue evaluada in vitro en cinco líneas celulares diferentes de cáncer histogenético (cáncer de próstata - PC-3, cáncer de mama, sarcoma de Kaposi, melanoma B-16 y un linfoma de células B de una cepa de ratón transgénico) usando el ensayo con azul de tripano. El extracto crudo también se evaluó in vivo en ratones portadores de tumor. La tinción inmunohistoquímica con ApopTag fue utilizada para la histología y determinación de la apoptosis. RESULTADOS: El extracto metanólico crudo de T recurvata demostró la actividad proliferativa frente a todas las líneas celulares, matando > 50% de las células a una concentración de 2,5 µg/ml. Los tumores de xenoinjerto de sarcoma de Kaposi fueron inhibidos hasta un 75% en comparación con el control en el estudio in vivo (p < 0.05). Hubo evidencia de fragmentación de DNA y una disminución en la viabilidad celular en los estudios histológicos. El extracto metanólico no mostró ningún efecto tóxico en los ratones a dosis de 200 mg/kg. CONCLUSIONES: Nuestros datos sugieren que la T recurvata tiene gran potencial como agente anticanceroso, y que uno de sus mecanismos de inhibición de tumores y muerte de las células tiene lugar mediante la inducción de la apoptosis.

Uso de bromeliáceas em biomonitoramento atmosférico / Bromeliads use in atmospheric biomonitoring

Tillandsia streptocarpa Baker e Tillandsia pohliana Mez. (Bromeliaceae) são plantas epifíticas neotropicais capazes de absorver água e nutrientes diretamente do ar. Por essa razão, foram selecionados essas espécies para realizar um estudo de monitoramento da qualidade do ar na cidade de Umuarama (PR, Brasil). Os conteúdos de, Ni, Cr, K, Zn, Cu e Pb foram determinados por espectrometria de absorção atômica. Os resultados obtidos mostraram uma acumulação mais acentuada dos elementos K e Fe nas espécies T. streptocarpa e T. pohliana. Resultados indicam ainda um potencial promissor dessas espécies como biomonitores de poluição do ar. Desse modo, recomenda-se o uso de Tillandsiaem locais onde espécies destas epífitas são frequentes.

Tillandsia streptocarpa Baker e Tillandsia pohliana Mez. (Bromeliaceae) are an epiphytic neotropical plants able to absorb water and nutrients directly from the air. For this reason we have selected these species to carry out a monitoring study of air quality in Umuarama city (PR, Brazil). The contents of Fe, Ni, Cr, K, Zn, Cu and Pb were determined by Atomic Absorption Spectrometry. The results obtained showed a more accentuated accumulation of K and Fe elements in T. streptocarpa e T. pohliana.. The results indicated a promising potential of these species as biomonitors of air pollution. We recommend the use of Tillandsia in sites where some species of these epiphytes are very widespread.

A new quantitative classification of ecological types in the bromeliad genus Tillandsia (Bromeliaceae) based on trichomes

Using collection specimens, we measured the density and wing area of trichomes in 37 species of the bromeliad genus Tillandsia, specifically the abaxial proximal, abaxial distal, adaxial proximal and adaxial distal parts of the leaf. The product of the trichome "wing" area by the number of trichomes (means) produced a pure number (T) that was correlated to ecological features. The correlation was positive with respect to arid environments (xeric Tillands) and negative with respect to humid environments (mesic Tillands). Bulbous, and particularly myrmecophytic species and species with tanks, represented particular categories. Other intermediate types were identified based on the T number, totalling five ecological types. In comparison with other systems of ecological typification for Tillands and other Bromeliaceae, the present system offers measurable data whose analysis is reproducible.

Medimos el número por milímetro cuadrado y el área del “ala” (parte móvil) de los tricomas en las partes adaxial próxima y distal, y adaxial próxima y distal, de la hoja de 37 especies de bromelias del género Tillandsia. El producto del área del ala para el número de los tricomas (promedio) produjo un número puro (T). Hallamos que T se correlaciona con las características ecológicas de las tilandsias investigadas. La correlación es positiva con respecto a ambientes áridos (especies xéricas) y negativa con respecto a los ambientes húmedos (especies mésicas). Las especies con bulbo, y particularmente las asociadas con hormigas y especies con de tanque representan categorías particulares. Identificamos otros tipos intermedios, agradando así cinco tipos ecológicos. En comparación con otros sistemas de tipificación ecológica, este sistema ofrece la ventaja de ser reproducible y cuantitativo.

Potencial abortivo de Tillandsia usneoides L. (Barba-de-pau) em coelhas gestantes - nota prévia / Abortive potential of Tillandsia usneoides L. in pregnant rabbits - preliminary report / Potencial abortivo de tillandsia usneoides l. en conejas gestantes ­ relato preliminar

Tillandsia usneoides L., também conhecida como barba-de-pau ou barba-de-velho, é uma Bromeliaceae presente em grande parte do território brasileiro, e em áreas úmidas desde o sudoeste dos EUA até o Chile e centro da Argentina. Relatos de criadores de animais relacionam essa planta à ocorrência de abortos em éguas e vacas. Os objetivos deste trabalho foram a análise fitoquímica de partes totais de Tillandsia usneoides e a avaliação preliminar do potencial abortivo de seu extrato hidroalcoólico em coelhas. Na análise fitoquímica observou-se a presença de flavonóides, esteróides, antraquinonas, cumarinas, micotoxinas, antocianinas, saponinas, taninos e aminogrupos. Os resultados da administração do extrato hidroalcoólico liofilizado a coelhas gestantes foram inconclusivos, em função da ocorrência de pseudogestação e da alta variação do número de filhotes nascidos dentro dos grupos de estudo.

Tillandsia usneoides L., also known as Spanish moss, is a Bromeliaceae present to a large extent of the Brasilian territory, and in humid areas from southeastern U.S.A. to Chile and central Argentina. Farmers report a relation of this plant to the occurrence of abortions in mares and cows. The objectives of this study were the phytochemical analysis of total parts of Tillandsia usneoides and the evaluation of the abortive potential of its hydroalcoholic extract in female rabbits. The phytochemical analysis showed fl avonoids, steroids, antraquinons, coumarins, micotoxins, antocianins, saponins, tannins, and aminogroups. The results of administration of the lyophilized hidroalcoolic extract to pregnant rabbits were not conclusive due the occurrence of pseudopregnancy and the high variation in number of younglings born within the studied groups.

Tillandsia usneoides L. es una Bromeliaceae presente en gran parte del territorio brasileño, y en áreas húmedas del sudeste de los EUA hasta Chile y centro de Argentina. Granjeros mencionan una relación de esta planta con la ocurrencia de abortos en yeguas y vacas. Los objetivos de este estudio fueron el análisis fi toquímico de partes totales de Tillandsia usneoides y la evaluación del potencial abortivo de su extracto hidroalcohólico en conejas. En el análisis fi toquímico se observaron fl avonoides, esteroides, antraquinonas, cumarinas, micotoxinas, antocianinas, saponinas, taninos y aminogrupos. Los resultados de la administración del extracto hidroalcohólico liofi lizado a conejas preñes no fueron conclusivos debido la ocurrencia de pseudopreñez y a la alta variación en el número de conejitos nacidos dentro de los grupos estudiados.