Different Slopes of a Mountain Can Determine the Structure of Ferns and Lycophytes Communities in a Tropical Forest of Brazil
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Glenda Gabriela Cárdenas Ramírez
ANNALES UNIVERSITATIS TURKUENSIS UNIVERSITATIS ANNALES A II 353 Glenda Gabriea Cárdenas Ramírez EVOLUTIONARY HISTORY OF FERNS AND THE USE OF FERNS AND LYCOPHYTES IN ECOLOGICAL STUDIES Glenda Gabriea Cárdenas Ramírez Painosaama Oy, Turku , Finand 2019 , Finand Turku Oy, Painosaama ISBN 978-951-29-7645-4 (PRINT) TURUN YLIOPISTON JULKAISUJA – ANNALES UNIVERSITATIS TURKUENSIS ISBN 978-951-29-7646-1 (PDF) ISSN 0082-6979 (Print) ISSN 2343-3183 (Online) SARJA - SER. A II OSA - TOM. 353 | BIOLOGICA - GEOGRAPHICA - GEOLOGICA | TURKU 2019 EVOLUTIONARY HISTORY OF FERNS AND THE USE OF FERNS AND LYCOPHYTES IN ECOLOGICAL STUDIES Glenda Gabriela Cárdenas Ramírez TURUN YLIOPISTON JULKAISUJA – ANNALES UNIVERSITATIS TURKUENSIS SARJA - SER. A II OSA – TOM. 353 | BIOLOGICA - GEOGRAPHICA - GEOLOGICA | TURKU 2019 University of Turku Faculty of Science and Engineering Doctoral Programme in Biology, Geography and Geology Department of Biology Supervised by Dr Hanna Tuomisto Dr Samuli Lehtonen Department of Biology Biodiversity Unit FI-20014 University of Turku FI-20014 University of Turku Finland Finland Reviewed by Dr Helena Korpelainen Dr Germinal Rouhan Department of Agricultural Sciences National Museum of Natural History P.O. Box 27 (Latokartanonkaari 5) 57 Rue Cuvier, 75005 Paris 00014 University of Helsinki France Finland Opponent Dr Eric Schuettpelz Smithsonian National Museum of Natural History 10th St. & Constitution Ave. NW, Washington, DC 20560 U.S.A. The originality of this publication has been checked in accordance with the University of Turku quality assurance system using the Turnitin OriginalityCheck service. ISBN 978-951-29-7645-4 (PRINT) ISBN 978-951-29-7646-1 (PDF) ISSN 0082-6979 (Print) ISSN 2343-3183 (Online) Painosalama Oy – Turku, Finland 2019 Para Clara y Ronaldo, En memoria de Pepe Barletti 5 TABLE OF CONTENTS ABSTRACT ........................................................................................................................... -
Author's Personal Copy
Author's personal copy Plant Syst Evol DOI 10.1007/s00606-013-0933-4 ORIGINAL ARTICLE Phylogeny and classification of the Cuban species of Elaphoglossum (Dryopteridaceae), with description of Elaphoglossum sect. Wrightiana sect. nov. Josmaily Lo´riga • Alejandra Vasco • Ledis Regalado • Jochen Heinrichs • Robbin C. Moran Received: 23 May 2013 / Accepted: 12 October 2013 Ó Springer-Verlag Wien 2013 Abstract Although a worldwide phylogeny of the bol- primary hemiepiphytism of E. amygdalifolium, which is bitidoid fern genus Elaphoglossum is now available, little sister to the rest of the genus, was derived independently is known about the phylogenetic position of the 34 Cuban from ancestors that were root climbers. Based on our species. We performed a phylogenetic analysis of a chlo- phylogenetic analysis and morphological investigations, roplast DNA dataset for atpß-rbcL (including a fragment of the species of Cuban Elaphoglossum were found to occur the gene atpß), rps4-trnS, and trnL-trnF. The dataset in E. sects. Elaphoglossum, Lepidoglossa, Polytrichia, included 79 new sequences of Elaphoglossum (67 from Setosa, and Squamipedia. Cuba) and 299 GenBank sequences of Elaphoglossum and its most closely related outgroups, the bolbitidoid genera Keywords Bolbitidoid fern Á Chloroplast DNA Arthrobotrya, Bolbitis, Lomagramma, Mickelia, and Ter- sequences Á Growth habit Á Holoepiphytism Á Primary atophyllum. We obtained a well-resolved phylogeny hemiepiphytism Á Root climber Á Taxonomy including the seven main lineages recovered in previous phylogenetic studies of Elaphoglossum. The Cuban ende- mic E. wrightii was found to be an early diverging lineage Introduction of Elaphoglossum, not a member of E. sect. Squamipedia where it was previously classified. -
Biogeographical Patterns of Species Richness, Range Size And
Biogeographical patterns of species richness, range size and phylogenetic diversity of ferns along elevational-latitudinal gradients in the tropics and its transition zone Kumulative Dissertation zur Erlangung als Doktorgrades der Naturwissenschaften (Dr.rer.nat.) dem Fachbereich Geographie der Philipps-Universität Marburg vorgelegt von Adriana Carolina Hernández Rojas aus Xalapa, Veracruz, Mexiko Marburg/Lahn, September 2020 Vom Fachbereich Geographie der Philipps-Universität Marburg als Dissertation am 10.09.2020 angenommen. Erstgutachter: Prof. Dr. Georg Miehe (Marburg) Zweitgutachterin: Prof. Dr. Maaike Bader (Marburg) Tag der mündlichen Prüfung: 27.10.2020 “An overwhelming body of evidence supports the conclusion that every organism alive today and all those who have ever lived are members of a shared heritage that extends back to the origin of life 3.8 billion years ago”. This sentence is an invitation to reflect about our non- independence as a living beins. We are part of something bigger! "Eine überwältigende Anzahl von Beweisen stützt die Schlussfolgerung, dass jeder heute lebende Organismus und alle, die jemals gelebt haben, Mitglieder eines gemeinsamen Erbes sind, das bis zum Ursprung des Lebens vor 3,8 Milliarden Jahren zurückreicht." Dieser Satz ist eine Einladung, über unsere Nichtunabhängigkeit als Lebende Wesen zu reflektieren. Wir sind Teil von etwas Größerem! PREFACE All doors were opened to start this travel, beginning for the many magical pristine forest of Ecuador, Sierra de Juárez Oaxaca and los Tuxtlas in Veracruz, some of the most biodiverse zones in the planet, were I had the honor to put my feet, contemplate their beauty and perfection and work in their mystical forest. It was a dream into reality! The collaboration with the German counterpart started at the beginning of my academic career and I never imagine that this will be continued to bring this research that summarizes the efforts of many researchers that worked hardly in the overwhelming and incredible biodiverse tropics. -
Rafaelcruzteseoriginal.Pdf
Rafael Cruz Desenvolvimento de folhas em samambaias sob a visão contínua de Agnes Arber em morfologia vegetal Development of leaves in ferns under the Agnes Arber’s continuum view of plant morphology Tese apresentada ao Instituto de Biociências da Universidade de São Paulo para a obtenção do título de Doutor em Ciências Biológicas, na área de Botânica. Orientação: Profa. Dra. Gladys Flávia de Albuquerque Melo de Pinna Coorientação: Prof. Dr. Jefferson Prado São Paulo 2018 Ficha Catalográfica: Cruz, Rafael Desenvolvimento de folhas em samambaias sob a visão contínua de Agnes Arber em morfologia vegetal. São Paulo, 2018. 104 páginas. Tese (Doutorado) – Instituto de Biociências da Universidade de São Paulo. Departamento de Botânica. 1. Desenvolvimento; 2. Anatomia Vegetal; 3. Expressão Gênica. I. Universidade de São Paulo. Instituto de Biociências. Departamento de Botânica. Comissão Julgadora: __________________________________ __________________________________ Prof(a). Dr(a). Prof(a). Dr(a). __________________________________ __________________________________ Prof(a). Dr(a). Prof(a). Dr(a). __________________________________ Profa. Dra. Gladys Flávia A. Melo de Pinna Orientadora Aos meus queridos amigos. “'The different branches [of biology] should not, indeed, be regarded as so many fragments which, pieced together, make up a mosaic called biology, but as so many microcosms, each of which, in its own individual way, reflects the macrocosm of the whole subject.” Agnes Robertson Arber The Natural Philosophy of Plant Form (1950) Agradecimentos À minha querida orientadora, Profa. Dra. Gladys Flávia Melo-de-Pinna, por mais de dez anos de orientação. Pela liberdade que me deu em propor ideias e pelo apoio que me deu em executá-las. Agradeço não só pela orientação, mas por ter se tornado uma grande amiga, e por me resgatar em todo momento que acho que fazer pesquisa poderia ser uma má ideia. -
Class I KNOX Is Related to Determinacy During the Leaf Development of the Fern Mickelia Scandens (Dryopteridaceae)
International Journal of Molecular Sciences Article Class I KNOX Is Related to Determinacy during the Leaf Development of the Fern Mickelia scandens (Dryopteridaceae) Rafael Cruz 1,2,* , Gladys F. A. Melo-de-Pinna 2, Alejandra Vasco 3 , Jefferson Prado 1,4 and Barbara A. Ambrose 5 1 Instituto de Botânica, Av. Miguel Estéfano 3687, São Paulo (SP) CEP 04301-902, Brazil; [email protected] 2 Instituto de Biociências, Universidade de São Paulo, Rua do Matão 277, São Paulo (SP) CEP 05422-971, Brazil; [email protected] 3 Botanical Research Institute of Texas, 1700 University Drive, Fort Worth, TX 76107-3400, USA; [email protected] 4 UNESP, IBILCE, Depto. de Zoologia e Botânica, Rua Cristóvão Colombo, 2265, São José do Rio Preto (SP) CEP 15054-000, Brazil 5 The New York Botanical Garden, 2900 Southern Blvd, Bronx, NY 10458-5126, USA; [email protected] * Correspondence: [email protected] Received: 2 May 2020; Accepted: 12 June 2020; Published: 16 June 2020 Abstract: Unlike seed plants, ferns leaves are considered to be structures with delayed determinacy, with a leaf apical meristem similar to the shoot apical meristems. To better understand the meristematic organization during leaf development and determinacy control, we analyzed the cell divisions and expression of Class I KNOX genes in Mickelia scandens, a fern that produces larger leaves with more pinnae in its climbing form than in its terrestrial form. We performed anatomical, in situ hybridization, and qRT-PCR experiments with histone H4 (cell division marker) and Class I KNOX genes. We found that Class I KNOX genes are expressed in shoot apical meristems, leaf apical meristems, and pinnae primordia. -
Phylogenetic Analyses Place the Monotypic Dryopolystichum Within Lomariopsidaceae
A peer-reviewed open-access journal PhytoKeysPhylogenetic 78: 83–107 (2017) analyses place the monotypic Dryopolystichum within Lomariopsidaceae 83 doi: 10.3897/phytokeys.78.12040 RESEARCH ARTICLE http://phytokeys.pensoft.net Launched to accelerate biodiversity research Phylogenetic analyses place the monotypic Dryopolystichum within Lomariopsidaceae Cheng-Wei Chen1,*, Michael Sundue2,*, Li-Yaung Kuo3, Wei-Chih Teng4, Yao-Moan Huang1 1 Division of Silviculture, Taiwan Forestry Research Institute, 53 Nan-Hai Rd., Taipei 100, Taiwan 2 The Pringle Herbarium, Department of Plant Biology, The University of Vermont, 27 Colchester Ave., Burlington, VT 05405, USA 3 Institute of Ecology and Evolutionary Biology, National Taiwan University, No. 1, Sec. 4, Roosevelt Road, Taipei, 10617, Taiwan 4 Natural photographer, 664, Hu-Shan Rd., Caotun Township, Nantou 54265, Taiwan Corresponding author: Yao-Moan Huang ([email protected]) Academic editor: T. Almeida | Received 1 February 2017 | Accepted 23 March 2017 | Published 7 April 2017 Citation: Chen C-W, Sundue M, Kuo L-Y, Teng W-C, Huang Y-M (2017) Phylogenetic analyses place the monotypic Dryopolystichum within Lomariopsidaceae. PhytoKeys 78: 83–107. https://doi.org/10.3897/phytokeys.78.12040 Abstract The monotypic fern genusDryopolystichum Copel. combines a unique assortment of characters that ob- scures its relationship to other ferns. Its thin-walled sporangium with a vertical and interrupted annulus, round sorus with peltate indusium, and petiole with several vascular bundles place it in suborder Poly- podiineae, but more precise placement has eluded previous authors. Here we investigate its phylogenetic position using three plastid DNA markers, rbcL, rps4-trnS, and trnL-F, and a broad sampling of Polypodi- ineae. -
Criptógamos Do Parque Estadual Das Fontes Do Ipiranga, São Paulo, SP, Brasil
Hoehnea 39(4): 555-564, 1 fig., 2012 Criptógamos do Parque Estadual das Fontes do Ipiranga, São Paulo, SP, Brasil. Pteridophyta: 7. Dryopteridaceae e 11. Lomariopsidaceae Regina Yoshie Hirai1,2 e Jefferson Prado1 Recebido: 18.07.2012; aceito: 6.11.2012 ABSTRACT - (Cryptogams of Parque Estadual das Fontes do Ipiranga, São Paulo, São Paulo State, Brazil. Pteridophyta: 7. Dryopteridaceae and 11. Lomariopsidaceae). The data of the floristic survey of the families Dryopteridaceae and Lomariopsidaceae in Parque Estadual das Fontes do Ipiranga (PEFI) are presented. Five genera and nine species were found in the area. Dryopteridaceae is represented by two genera (Polybotrya and Rumohra) and four species: Polybotrya cylindrica Kaulf., P. semipinnata Fée, P. speciosa Schott, and Rumohra adiantiformis (G. Forst.) Ching, while Lomariopsidaceae is represented by three genera (Elaphoglossum, Mickelia, and Lomariopsis) and five species:Elaphoglossum ornatum (Mett. ex Kuhn) Christ, E. nigrescens (Hook.) T. Moore ex Diels, E. macrophyllum (Mett. ex Kuhn) Christ, Mickelia scandens (Raddi) R.C. Moran et al., and Lomariopsis marginata (Schrad.) Kuhn. Identification keys for genera and species, as well as descriptions, geographical distribution, comments, and illustrations for some studied taxa are presented. Key words: Elaphoglossum, Lomariopsis, Mickelia, Polybotrya, Rumohra RESUMO - (Criptógamos do Parque Estadual das Fontes do Ipiranga, São Paulo, SP, Brasil. Pteridophyta: 7. Dryopteridaceae e 11. Lomariopsidaceae). Neste trabalho são apresentados os dados referentes ao levantamento florístico das famílias Dryopteridaceae e Lomariopsidaceae no Parque Estadual das Fontes do Ipiranga (PEFI). No total foram encontrados na área cinco gêneros e nove espécies, sendo que Dryopteridaceae está representada por dois gêneros (Polybotrya e Rumohra) e quatro espécies: Polybotrya cylindrica Kaulf., P. -
Riqueza De Samambaias E Licófitas De Uma Mata De Galeria Na Região Central De Mato Grosso Do Sul, Brasil
Biotemas, 26 (1): 7-15, março de 2013 doi: 10.5007/2175-7925.2013v26n1p77 ISSNe 2175-7925 Riqueza de samambaias e licófitas de uma mata de galeria na região central de Mato Grosso do Sul, Brasil Carlos Rodrigo Lehn 1* Elton Luis Monteiro de Assis 2 1 Instituto Federal de Mato Grosso do Sul, Campus Coxim, Rua Pereira Gomes ,355 CEP 79400-000, Coxim – MS, Brasil 2 PPG em Botânica Tropical, Jardim Botânico do Rio de Janeiro Rua Pacheco Leão, 2040, Bairro Horto, CEP 22460-036, Rio de Janeiro – RJ, Brasil * Autor para correspondência [email protected] Submetido em 22/03/2012 Aceito para publicação em 14/10/2012 Resumo Neste trabalho é apresentado o levantamento florístico das samambaias e licófitas ocorrentes em uma mata de galeria, na região central de Mato Grosso do Sul, Brasil. Foram registradas na área de estudo 29 espécies e duas variedades. Dryopteridaceae e Pteridaceae foram as famílias mais ricas (oito e cinco espécies, respectivamente) e Elaphoglossum e Blechnum foram os gêneros mais ricos (três espécies cada). Preferencialmente, as espécies listadas ocorrem no interior da mata (68%), ocupam o substrato terrícola (77,4%), são hemicriptófitas (77,4%) e rosuladas (64,5%). Observamos quatro espécies ainda não citadas para Mato Grosso do Sul, sendo essas Blechnum lanceola L., Elaphoglossum pachydermum (Fée) T. Moore, Lindsaea lancea (L.) Bedd var lancea e ainda Mickelia nicotianifolia (Sw.) R. C. Moran et al., que possui seu limite sul de distribuição no Brasil, na área de estudo. Palavras-chave: Centro-Oeste; Cerrado; Lycophyta; Monilophyta; Pteridófitas Abstract Richness of ferns and lycophytes in a gallery forest in the central region of Mato Grosso do Sul, Brazil. -
Curriculum Vitae Alejandra Vasco
Alejandra Vasco CV 1 CURRICULUM VITAE ALEJANDRA VASCO Botanical Research Institute of Texas (BRIT) 1700 University Dr., Fort Worth, TX 76107-3400 USA Phone: (1) 817 773 0803 E-mail: [email protected] www.researchgate.net/profile/AlejandraVasco I EDUCATION Ph.D. in Biology (2010). City University of New York (CUNY)/ The New York Botanical Garden. Dissertation Title: Systematics and Phylogeny of Elaphoglossum section Lepidoglossa (Dryopteridaceae). Advisor: Dr. Robbin C. Moran M.Phil. in Biology (2007). City University of New York (CUNY)/ The New York Botanical Garden. Advisor: Dr. Robbin C. Moran B. S. in Biology (2002). University of Antioquia, Biology Department. Medellín, Colombia. Advisor: Dr. Ricardo Callejas II PROFESSIONAL APPOINTMENTS Research Botanist. Aug 2017–present. Botanical Research Institute of Texas, Fort Worth, Texas, USA. Adjoint Professor 2017–2020. University of Texas Arlington, Department of Biology, Arlington, Texas, USA. Associate member of the Graduate Faculty 2017–2023. Texas Christian University, Fort Worth, Texas, USA. Assistant Professor. 2015–2017. Department of Botany, Biology Institute, National Autonomous University of Mexico, Mexico City. Postdoctoral Research Associate. 2014–2015. Biology Institute, National Autonomous University of Mexico, Mexico City. “Leaf anatomy in the Elaphoglossum petiolatum group”. Supervisor: Dr. Teresa Terrazas. Postdoctoral Research Associate. 2010–2013. The New York Botanical Garden. “Leaf development and phylogeny of the fern group Elaphoglossum section Squamipedia (Dryopteridaceae)”. Supervisors: Drs. Barbara Ambrose and Robbin Moran. Research assistant. 2007–2009. The New York Botanical Garden. “Monographic revision and phylogeny of the fern genus Megalastrum (Dryopteridaceae)”. Supervisor: Dr. Robbin Moran. Herbarium Curatorial Assistant. 2004–2007. The New York Botanical Garden New York, USA. -
Bolbitis Lianhuachihensis (Dryopteridaceae), a New Species from Taiwan
A peer-reviewed open-access journal PhytoKeys 131: 69–81Bolbitis (2019) lianhuachihensis (Dryopteridaceae), a new species from Taiwan 69 doi: 10.3897/phytokeys.131.36548 RESEARCH ARTICLE http://phytokeys.pensoft.net Launched to accelerate biodiversity research Bolbitis lianhuachihensis (Dryopteridaceae), a new species from Taiwan Yi-Shan Chao1, Yu-Fang Huang2, Shi-Yong Dong3, Yao-Moan Huang4, Ho-Yih Liu2 1 Department of Biomedical Science & Environmental Biology, Kaohsiung Medical University, 100, Shih- Chuan 1st Rd., Kaohsiung, 80708, Taiwan 2 Department of Biological Sciences, National Sun Yat-sen Univer- sity, Kaohsiung 80424, Taiwan 3 South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650, China 4 Taiwan Forestry Research Institute, 53 Nan-Hai Rd., Taipei, 10066, Taiwan Corresponding author: Yi-Shan Chao ([email protected]); Ho-Yih Liu ([email protected]) Academic editor: Thais Almeida | Received 27 May 2019 | Accepted 31 July 2019 | Published 5 September 2019 Citation: Chao Y-S, Huang Y-F, Dong S-Y, Huang Y-M, Liu H-Y (2019) Bolbitis lianhuachihensis (Dryopteridaceae), a new species from Taiwan. PhytoKeys 131: 69–81. https://doi.org/10.3897/phytokeys.131.36548 Abstract A new species of Bolbitis, B. lianhuachihensis sp. nov., was found in central Taiwan. It most resembles B. virens var. compacta and B. hainanensis. A phylogenetic tree of Taiwanese and other Asian species of Bolbitis species supports the recognition of the new species. Morphologically, the combination of anasto- mosing venation and fewer sterile pinnae are critical characters to discriminate B. lianhuachihensis from other Taiwanese Bolbitis species. Bolbitis lianhuachihensis can be further distinguished from B. virens var. -
Elaphoglossum
Dissertation zur Erlangung des Doktorgrades der Naturwissenschaften (Dr. rer. nat.) an der Fakultät für Biologie der Ludwig-Maximilians-Universität München EVOLUTION AND CLASSIFICATION OF ELAPHOGLOSSUM AND ASPLENIUM FERNS ON CUBA, AND DISCOVERY OF A MIOCENE ELAPHOGLOSSUM IN DOMINICAN AMBER Josmaily Lóriga Piñeiro München, 15. Januar 2018 ii A mi familia y mis amigos, donde quiera que estén To my family and my friends, wherever they are iii iv PREFACE Statutory declaration Erklärung Diese Dissertation wurde im Sinne von §12 der Promotionsordnung von Prof. Dr. Jochen Heinrichs betreut. Ich erkläre hiermit, dass die Dissertation nicht einer anderen Prüfungskommission vorgelegt worden ist und dass ich mich nicht anderweitig einer Doktorprüfung ohne Erfolg unterzogen habe. Eidesstattliche Erklärung Ich versichere hiermit an Eides statt, dass die vorgelegte Dissertation von mir selbstständig und ohne unerlaubte Hilfe angefertigt wurde. Josmaily Lóriga, 15. Januar 2018 (Unterschrift) 1. Gutachter: Prof. Dr. Susanne S. Renner 2. Gutachter: Prof. Dr. Reinhard Agerer Tag der Abgabe: 15. Januar 2018 Tag der Disputation: 28. Februar 2018 v vi Declaration of contribution In this thesis, I present the results from my doctoral research, carried out in Munich from March 2013 to January 2018 (maternity leave between February and December 2017) under the guidance of Prof. Dr. Jochen Heinrichs. My thesis resulted in three published manuscripts presented in Chapters 2 to 4. I generated all data and conducted all analyses myself with some exceptions: Chapter 2, Alejandra Vasco produced Fig. 4; Chapter 3, Ledis Regalado conducted the study of gametophytes and generated the drawings of Fig. 1; and Chapter 4, Kathrin Feldberg contributed to the divergence time estimates and Alexander R. -
Leaf Water Relations in Epiphytic Ferns Are Driven by Drought Avoidance
Leaf water relations in epiphytic ferns are driven by drought avoidance rather than tolerance mechanisms Courtney Campany1, Jarmila Pittermann2, Alex Baer2, Helen Holmlund3, Eric Schuettpelz4, Klaus Mehltreter5, and James (Eddie) Watkins6 1Shepherd University 2University of California Santa Cruz 3Pepperdine University 4Smithsonian Institution 5Instituto de Ecologia 6Colgate University February 4, 2021 Abstract Opportunistic diversification has allowed ferns to radiate into epiphytic niches in angiosperm dominated landscapes. However, our understanding of how ecophysiological function allowed establishment in the canopy and the potential transitionary role of the hemi-epiphytic life form remain unclear. Here, we surveyed 39 fern species in Costa Rican tropical forests to explore epiphytic trait divergence in a phylogenetic context. We examined leaf responses to water deficits in terrestrial, hemi-epiphytic, and epiphytic ferns and related these findings to functional traits that regulate leaf water status. Epiphytic ferns had reduced xylem area (-63%), shorter stipe lengths (-56%), thicker laminae (+41%), and reduced stomatal density (-46%) compared to terrestrial ferns. Epiphytic ferns exhibited similar turgor loss points, higher osmotic potential at saturation, and lower tissue capacitance after turgor loss than terrestrial ferns. Overall, hemi-epiphytic ferns exhibited traits that share characteristics of both terrestrial and epiphytic species. Our findings clearly demonstrate the prevalence of water conservatism in both epiphytic and hemi-epiphytic