Flower Phenology and Sexual Maturation: Partial Protandrous Behavior in Three Species of Orchids

Total Page:16

File Type:pdf, Size:1020Kb

Flower Phenology and Sexual Maturation: Partial Protandrous Behavior in Three Species of Orchids Caribbean Journal of Science, Vol. 42, No. 1, 75-80, 2006 Copyright 2006 College of Arts and Sciences University of Puerto Rico, Mayagu¨ez Flower Phenology and Sexual Maturation: Partial Protandrous Behavior in Three Species of Orchids RAYMOND L. TREMBLAY*, GRIZEL POMALES-HERNÁNDEZ AND MARÍA DE LOURDES MÉNDEZ-CINTRÓN Department of Biology, University of Puerto Rico—Humacao, 100 carr. 908, Humacao, P.R., 00791 *Corresponding author: [email protected] ABSTRACT.—Plants have theoretically multiple alternatives for preventing self pollination and conse- quently the effect of inbreeding, such as sequential flowering, dichogamy and self–incompatibility to name a few. We investigated the reproductive biology of three sequentially flowering (acropetal) endemic orchids from Puerto Rico. Since sequential flowering is present in the studied species and very rarely (1.0%) is there more than one flower open simultaneously on an inflorescence, we hypothesized that the orchids should be self-compatible and show no effect of protandry (dichogamy). We performed hand self—and cross- pollinations and evaluated whether the species are self-compatible and whether the receptivity to pollination success (fruit set) is influenced by the age of flowers (protandry). We define protandry as pertaining to a hermaphroditic organism that assumes a functional male condition prior to shifting to a functional female state. We found that all three species are self-incompatible. Furthermore, flower age is important for pre- dicting the likelihood of fruits set. Older flowers (6+ days) are significantly more likely to produce fruits (functional protandry). The multiple mechanisms for preventing self-pollination (sequential flowering, di- chogamy and self-incompatibility) that are noted for these species suggest that the historical evolutionary processes for preventing inbreeding may be complex. We hypothesized that because multiple mechanisms are present for preventing self-pollination inbreeding depression is likely to be high. KEYWORDS.—Lepanthes, Puerto Rico, reproductive success, dichogamy, self-incompatible INTRODUCTION togyny) and solitary flowers (Delaporta and Calderón-Urrea 1993; Walker-Larson Low fruit set in many plant species is a and Harder 2000). consequence of pollinator limitation (Trem- In the Orchidaceae, pollinator limitation blay et al. 2005), or self-incompatibility, is frequently invoked as the cause of low which are perceived as mechanisms to pro- fruit set, but in self-compatible species low mote cross-pollination (Marshall and Fol- fruit set may additionally be attributed to som 1991). Promoting cross-pollination is incompatible self—and geitonogamous pol- believed to increase fitness by increasing linations, resulting in flower or fruit abor- genetic diversity, while self-pollination de- tion (Delaporta and Calderón-Urrea 1993; creases genetic diversity and can result in Walker-Larson and Harder 2000; Tremblay inbreeding depression (Lande and Schem- et al. 2005). In self-compatible orchids a ske 1985; Dudash 1990; Jarne and Charles- number of mechanisms for preventing self- worth 1993). Consequently, many species pollination (Johnson and Edwards 2000) probably evolved complex mechanisms in have been detected, including: sequential order to prevent self-pollination, including flowering (Psychilis spp., Ackerman (1989); self-incompatibility through sporophytic or Malaxis massonii (Ridl.) Kuntze, Aragón gametophytic processes, and through pre- and Ackerman (2001)), dichogamy/pro- visitation mechanisms, including sequen- tandry (Goodyera oblongifolia Raf., Acker- tial flowering, dichogamy (protandry, pro- man (1975); Listera cordata, Ackerman and Mesler (1979); Spiranthes, Catling (1983); ms. received October 2, 2004; accepted August 31, Sipes and Tepedino (1995); Prescottia 2005 stachyodes (Sw.) Lindl., Singer and Sazima 75 76 R. L. TREMBLAY ET AL. (2001a); Erythrodes arietina (Rchb. F. & manipulation (apomixis). No fruits were Warm.) Ames, Singer and Sazima (2001b); set unless flowers were hand pollinated. Sauroglossum elatum Lindl., Singer (2002); Mesadenella cuspidata (Lindl.) Garay, Singer Determination of the breeding system (2002); Notylia nemorosa Barb. Rodr., Singer and Koehler (2002); Warford (1992); Man- Cross pollination vs. self pollination.— niella cypripedioides Salazar, T. Franke, Flowers of the three species of Lepanthes Zapfack & Beenken, Salazar et al. (2002)), studied do not show any evidence of self- temporal variation in pollinarium size (Bul- pollination, but to determine if autogamous bophyllum weddelii (Lindl.) Rchb. f., B. ipane- pollinations produce viable fruits, we per- mense Hoehne and B. involutum Borba, formed a series of hand self-pollinations. Semir and F. Barros, Borba and Semir We self-pollinated 3-day old flowers of L. (1999a); Trigonidium obtusum Lindl., Singer rupestris (n = 33) and L. woodburyana (n = (2002)), and change in position of lip and 50), and 6-day old flowers for L. rubripetala column after pollinarium removal or after a (n = 14). few days if pollinarium is not removed Determination of protandry.—To test if (Sosa and Rodríguez-Angulo 2000). protandry is present in Lepanthes we made We investigated whether or not self- cross-pollinations at different time intervals incompatibility is present in three sequen- of the flower age from one to eight days. tially flowering tropical species of Lepanthes We compared the percentage of fruit pro- (Orhcidaceae). In addition we determined duction per pollination for two age treat- whether their flowers are protandrous. ments (1-5 days and 6+ days) and among Protandry is a dichogamous mechanism; it the three species. We used contingency is defined as the maturation of the male tables to evaluate the null hypothesis that (androecium) structure before the matura- the pattern was random among days and tion of the female (gymnoecium) structures species. (Kearns and Inouye 1993). RESULTS MATERIALS AND METHODS Flower development and fruit phenology From the time we noticed flower bud set, We determined the breeding system of these took 11 to 15 days to open in all three three endemic species of Lepanthes from species. Open flowers lasted up to 10 days Puerto Rico: L. woodburyana Stimson (53 in L. rupestris and L. woodburyana, and 11 individuals), L. rupestris Stimson (56 indi- days in L. rubripetala when not manipu- viduals), and L. rubripetala Stimson (35 in- lated. However, flowers manipulated dividuals). Each species was grown in a TM through pollinia removal or pollination re- Wardian case (Orchidarium Case )inthe duced the life span of open flowers. In gen- laboratory at the same temperature and hu- eral, removal of pollinia caused flowers to midity (18-24°C, 45-100% humidity). close and wither within 1 to 2 days, more- The phenology of flower buds, flowers over, pollination (deposition of pollinia (including anthesis), and fruits for each onto the stigma) resulted in flower closure plant was recorded. The age of flower the next day (Table 1). Mean life span (until buds, flowers and viable fruits along each dehiscence) of fruits was estimated at 31 inflorescence was recorded daily. We con- and 39 days for L. rubripetala and L. rupes- sidered a fruit viable as soon as we noticed tris, respectively. None of the fruits that the expansion of the ovary. As a control we were set aborted (abscission of fruit in early segregated four specimens of Lepanthes stages of development). rupestris and L. rubripetala and L. wood- buryana to determine the effect of flower Hand self and cross-pollinations longevity on non-manipulated flowers, the effect of pollinia removal on flower longev- Hand self pollination did not produce ity and the ability to self-fertilize without any fruits for any of the three species of PROTANDRY IN ORCHIDS 77 TABLE 1. Flower phenology of Lepanthes flowers. Time (in days with s.e., sample size in parentheses). Characteristics L. rubripetala L. rupestris Development of flower bud to anthesis 14.6 ± 0.3 (104) 11.7 ± 0.2 (150) Flower lifespan without manipulation 6.9 ± 0.5 (32) 5.8 ± 0.4 (36) Flower lifespan with pollinia removal only 1.2 ± 0.1 (11) 1.4 ± 0.1 (46) Flower lifespan when flower is pollinated 1.0 ± 0.0 (55) 1.0 ± 0.0 (61) Fruits life span 31.2 ± 2.2 (23) 39.3 ± 1.3 (34) orchids. On the other hand, fruit set by the flowers of Habenaria orbiculata (Pursch) cross-pollinations (irrespective of flower Torr. last about 12.7 days in the field. How- age) was 23.2% for L. rubripetala, 23.1% for ever, flowers of species such as Dendrobium L. rupestris and 41.1% for L. woodburyana. crumenatum Sw. and Sobralia macrantha Lindl. last only one day, while an extreme The effect of flower age on fruit set case are the flowers of Grammatophyllum multiflorum Lindl. which last nine months The number of hand cross-pollination in greenhouses. In Lepanthes flower longev- varied among species and was dependent ity is in the range of 6 – 8 days. When flow- on flower production. Three hundred sev- ers are manipulated (pollinaria removal enty-three pollinations were performed in and pollinaria deposited) flower age is L. rupestris, 142 in L. rubripetala and 192 in L. drastically reduce to one day. This phe- woodburyana. The pattern of fruit set and nomenon of shorten flower life span when flower age was consistent among species. flowers are manipulated is well known in Older flowers (6+ days) had significantly orchids. Given the small size of the flowers, higher fruit set in all three species as com- a reduction in the life span of open flowers pared from the expected proportion (Table may not simply be due to flower manipu- 2). Under the null hypothesis, the expected lation but also to actual physical damage to number of fruits (observed in brackets) delicate sexual structures. were all significantly lower than observed The effect of protandry has been ob- fruit set, 12 (17) for L. rubripetala, 22 (31) for served in orchids. In Goodyera oblongifolia L. rupestris, 38 (56) for L. woodburyana. pollen deposition is prevented by the prox- imity of the column to the lip in young DISCUSSION flowers. As flowers age the column be- comes exposed and thus receptive to pollen Flower longevity is often short (a few deposition (Ackerman 1975).
Recommended publications
  • Table of Contents
    Appendix C Botanical Resources Table of Contents Purpose Of This Appendix ............................................................................................................. Below Tables C-1. Federal and State Status, Current and Proposed Forest Service Status, and Global Distribution of the TEPCS Plant Species on the Sawtooth National Forest ........................... C-1 C-2. Habit, Lifeform, Population Trend, and Habitat Grouping of the TEPCS Plant Species for the Sawtooth National Forest ............................................................................... C-3 C-3. Rare Communities, Federal and State Status, Rarity Class, Threats, Trends, and Research Natural Area Distribution for the Sawtooth National Forest ................................... C-5 C-4. Plant Species of Cultural Importance for the Sawtooth National Forest ................................... C-6 PURPOSE OF THIS APPENDIX This appendix is designed to provide detailed information about habitat, lifeform, status, distribution, and habitat grouping for the Threatened, Proposed, Candidate, and Sensitive (current and proposed) plant species found on the Sawtooth National Forest. The detailed information is provided to enable managers to more efficiently direct the implementation of Botanical Resources goals, objectives, standards, and guidelines. Additionally, this appendix provides detailed information about the rare plant communities located on the Sawtooth National Forest and should provide additional support of Forest-wide objectives. Species of cultural
    [Show full text]
  • Orchidaceae Juss. Aspectos Morfológicos E Taxonômicos
    ORCHIDACEAE JUSS. ASPECTOS MORFOLÓGICOS E TAXONÔMICOS VINÍCIUS TRETTEL RODRIGUES ORIENTADOR: DR. FÁBIO DE BARROS SÃO PAULO 2011 1 A FAMÍLIA ORCHIDACEAE Orchidaceae é a maior família, em número de espécies, entre as monocotiledôneas. Pertence à ordem Asparagales (APG 2006), sendo constituída por aproximadamente 24.500 espécies distribuídas em cerca de 800 gêneros (Dressler 1993, 2005). São plantas herbáceas, perenes, terrícolas ou, mais comumente, epífitas (cerca de 73% das espécies). Apresenta distribuição cosmopolita, embora seja mais abundante e diversificada em florestas tropicais, especialmente da Ásia e das Américas. (Figura 1). Figura 1: Distribuição da família Orchidaceae no mundo; a esquerda encontra-se o número de gêneros e a direita o de espécies. Adaptado de Dressler (1993). Nos Neotrópicos a família é amplamente diversificada, sobretudo na região equatorial, com grande diversidade de espécies na Colômbia, Equador, Brasil e Peru, (Figura 2). O Brasil detém uma das maiores diversidades de orquídeas do continente americano e do mundo, com cerca de 2.419 espécies das quais 1.620 são endêmicas deste país (Barros et al. 2010). Todas as formações vegetais brasileiras acomodam orquídeas, mas elas são mais numerosas nas formações florestais úmidas, principalmente na Mata Atlântica com cerca de 1.257 espécies distribuídas em 176 gêneros; dentre estas 791 espécies são endêmicas deste domínio (Barros et al. 2009). 2 Figura 2: Número de espécies da família Orchidaceae na América do Sul. Adaptado de Dressler (1993). Apesar da alta representatividade da família, que segundo Sanford (1974) abrange 7% das Angiospermas, ainda há muito a se descobrir. Dressler (1981) enfatiza que maiores estudos sobre a família devem ser feitos especialmente em regiões tropicais.
    [Show full text]
  • Phylogenetic Relationships of Discyphus Scopulariae
    Phytotaxa 173 (2): 127–139 ISSN 1179-3155 (print edition) www.mapress.com/phytotaxa/ PHYTOTAXA Copyright © 2014 Magnolia Press Article ISSN 1179-3163 (online edition) http://dx.doi.org/10.11646/phytotaxa.173.2.3 Phylogenetic relationships of Discyphus scopulariae (Orchidaceae, Cranichideae) inferred from plastid and nuclear DNA sequences: evidence supporting recognition of a new subtribe, Discyphinae GERARDO A. SALAZAR1, CÁSSIO VAN DEN BERG2 & ALEX POPOVKIN3 1Departamento de Botánica, Instituto de Biología, Universidad Nacional Autónoma de México, Apartado Postal 70-367, 04510 México, Distrito Federal, México; E-mail: [email protected] 2Universidade Estadual de Feira de Santana, Departamento de Ciências Biológicas, Av. Transnordestina s.n., 44036-900, Feira de Santana, Bahia, Brazil 3Fazenda Rio do Negro, Entre Rios, Bahia, Brazil Abstract The monospecific genus Discyphus, previously considered a member of Spiranthinae (Orchidoideae: Cranichideae), displays both vegetative and floral morphological peculiarities that are out of place in that subtribe. These include a single, sessile, cordate leaf that clasps the base of the inflorescence and lies flat on the substrate, petals that are long-decurrent on the column, labellum margins free from sides of the column and a column provided with two separate, cup-shaped stigmatic areas. Because of its morphological uniqueness, the phylogenetic relationships of Discyphus have been considered obscure. In this study, we analyse nucleotide sequences of plastid and nuclear DNA under maximum parsimony
    [Show full text]
  • Floristic Composition of a Neotropical Inselberg from Espírito Santo State, Brazil: an Important Area for Conservation
    13 1 2043 the journal of biodiversity data 11 February 2017 Check List LISTS OF SPECIES Check List 13(1): 2043, 11 February 2017 doi: https://doi.org/10.15560/13.1.2043 ISSN 1809-127X © 2017 Check List and Authors Floristic composition of a Neotropical inselberg from Espírito Santo state, Brazil: an important area for conservation Dayvid Rodrigues Couto1, 6, Talitha Mayumi Francisco2, Vitor da Cunha Manhães1, Henrique Machado Dias4 & Miriam Cristina Alvarez Pereira5 1 Universidade Federal do Rio de Janeiro, Museu Nacional, Programa de Pós-Graduação em Botânica, Quinta da Boa Vista, CEP 20940-040, Rio de Janeiro, RJ, Brazil 2 Universidade Estadual do Norte Fluminense Darcy Ribeiro, Laboratório de Ciências Ambientais, Programa de Pós-Graduação em Ecologia e Recursos Naturais, Av. Alberto Lamego, 2000, CEP 29013-600, Campos dos Goytacazes, RJ, Brazil 4 Universidade Federal do Espírito Santo (CCA/UFES), Centro de Ciências Agrárias, Departamento de Ciências Florestais e da Madeira, Av. Governador Lindemberg, 316, CEP 28550-000, Jerônimo Monteiro, ES, Brazil 5 Universidade Federal do Espírito Santo (CCA/UFES), Centro de Ciências Agrárias, Alto Guararema, s/no, CEP 29500-000, Alegre, ES, Brazil 6 Corresponding author. E-mail: [email protected] Abstract: Our study on granitic and gneissic rock outcrops environmental filters (e.g., total or partial absence of soil, on Pedra dos Pontões in Espírito Santo state contributes to low water retention, nutrient scarcity, difficulty in affixing the knowledge of the vascular flora of inselbergs in south- roots, exposure to wind and heat) that allow these areas eastern Brazil. We registered 211 species distributed among to support a highly specialized flora with sometimes high 51 families and 130 genera.
    [Show full text]
  • Orchid Historical Biogeography, Diversification, Antarctica and The
    Journal of Biogeography (J. Biogeogr.) (2016) ORIGINAL Orchid historical biogeography, ARTICLE diversification, Antarctica and the paradox of orchid dispersal Thomas J. Givnish1*, Daniel Spalink1, Mercedes Ames1, Stephanie P. Lyon1, Steven J. Hunter1, Alejandro Zuluaga1,2, Alfonso Doucette1, Giovanny Giraldo Caro1, James McDaniel1, Mark A. Clements3, Mary T. K. Arroyo4, Lorena Endara5, Ricardo Kriebel1, Norris H. Williams5 and Kenneth M. Cameron1 1Department of Botany, University of ABSTRACT Wisconsin-Madison, Madison, WI 53706, Aim Orchidaceae is the most species-rich angiosperm family and has one of USA, 2Departamento de Biologıa, the broadest distributions. Until now, the lack of a well-resolved phylogeny has Universidad del Valle, Cali, Colombia, 3Centre for Australian National Biodiversity prevented analyses of orchid historical biogeography. In this study, we use such Research, Canberra, ACT 2601, Australia, a phylogeny to estimate the geographical spread of orchids, evaluate the impor- 4Institute of Ecology and Biodiversity, tance of different regions in their diversification and assess the role of long-dis- Facultad de Ciencias, Universidad de Chile, tance dispersal (LDD) in generating orchid diversity. 5 Santiago, Chile, Department of Biology, Location Global. University of Florida, Gainesville, FL 32611, USA Methods Analyses use a phylogeny including species representing all five orchid subfamilies and almost all tribes and subtribes, calibrated against 17 angiosperm fossils. We estimated historical biogeography and assessed the
    [Show full text]
  • SPRING Windows to 2018 Wildlife Your Support at Work in Idaho’S Landscapes
    SPRING Windows to 2018 Wildlife Your support at work in Idaho’s landscapes Western toad Michael Lucid, IDFG Pale jumping slug Grizzly bear Michael Lucid, IDFG Helping Six Wildlife Species IN THIS ISSUE Spring Beat Climate Change Odds wildlife events by Michael Lucid, Regional Wildlife Diversity Biologist, Idaho Department of Fish and Game-Panhandle Region with Evan DeHamer*, Regional Habitat Biologist, Idaho Department of Fish and Game- Panhandle Region and Victoria Osborn*, Digital Media and Video Producer, Idaho Department of Fish and Game 4 Bumble bees An invasive frog in the wetlands of North Idaho is triggering climate related habitat of Idaho work on one of Idaho Fish and Game’s Wildlife Management Areas. In recent years warm temperature loving American bullfrogs have been making their way north along the Kootenai River Valley toward the Boundary-Smith Creek Wildlife Management Area (BSCWMA) near the US-Canadian border. Bullfrogs threaten native amphibians and pose an immediate threat to the last natural colony of native Northern leopard frogs in the northwestern portion of that species range. This bullfrog invasion has spurred emergency 7 international bullfrog control actions and inspired Idaho Fish and Game and partners to obtain funding to manage the WMA to benefit several climate sensitive Species of Greatest Conservation Need (SGCN) including native amphibians, bumble bees, slugs, and bears. Multiple conservation actions will be implemented in the Forested Lowland Ecosystem as directed by Idaho’s State Wildlife Action Plan. Idaho’s Orchids 9 2 Windows to Wildlife Spring 2018 Fish and Game and the Yellowstone to Yukon The globally changing climate brings local habitat Conservation Initiative (Y2Y) have partnered to secure shifts that SGCN’s may not be able to adapt to three grants totaling $549,624 to conduct climate without human intervention.
    [Show full text]
  • Phylogenetic Relationships of Discyphus
    Phytotaxa 173 (2): 127–139 ISSN 1179-3155 (print edition) www.mapress.com/phytotaxa/ PHYTOTAXA Copyright © 2014 Magnolia Press Article ISSN 1179-3163 (online edition) http://dx.doi.org/10.11646/phytotaxa.173.2.3 Phylogenetic relationships of Discyphus scopulariae (Orchidaceae, Cranichideae) inferred from plastid and nuclear DNA sequences: evidence supporting recognition of a new subtribe, Discyphinae GERARDO A. SALAZAR1, CÁSSIO VAN DEN BERG2 & ALEX POPOVKIN3 1Departamento de Botánica, Instituto de Biología, Universidad Nacional Autónoma de México, Apartado Postal 70-367, 04510 México, Distrito Federal, México; E-mail: [email protected] 2Universidade Estadual de Feira de Santana, Departamento de Ciências Biológicas, Av. Transnordestina s.n., 44036-900, Feira de Santana, Bahia, Brazil 3Fazenda Rio do Negro, Entre Rios, Bahia, Brazil Abstract The monospecific genus Discyphus, previously considered a member of Spiranthinae (Orchidoideae: Cranichideae), displays both vegetative and floral morphological peculiarities that are out of place in that subtribe. These include a single, sessile, cordate leaf that clasps the base of the inflorescence and lies flat on the substrate, petals that are long-decurrent on the column, labellum margins free from sides of the column and a column provided with two separate, cup-shaped stigmatic areas. Because of its morphological uniqueness, the phylogenetic relationships of Discyphus have been considered obscure. In this study, we analyse nucleotide sequences of plastid and nuclear DNA under maximum parsimony
    [Show full text]
  • Orchidaceae, Spiranthinae) and a New Genus from the Espinhaço Range, Southeastern Brazil Authors: Gerardo A
    Polyphyly of Mesadenus (Orchidaceae, Spiranthinae) and a New Genus from the Espinhaço Range, Southeastern Brazil Authors: Gerardo A. Salazar, João A. N. Batista, Thiago E. C. Meneguzzo, Lidia I. Cabrera, Coyolxauhqui Figueroa, et. al. Source: Systematic Botany, 44(2) : 282-296 Published By: The American Society of Plant Taxonomists URL: https://doi.org/10.1600/036364419X15562054132974 BioOne Complete (complete.BioOne.org) is a full-text database of 200 subscribed and open-access titles in the biological, ecological, and environmental sciences published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Complete website, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/terms-of-use. Usage of BioOne Complete content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Downloaded From: https://bioone.org/journals/Systematic-Botany on 29 Jul 2019 Terms of Use: https://bioone.org/terms-of-use Access provided by Universidad Autonoma de Ciudad Juarez Systematic Botany (2019), 44(2): pp. 282–296 © Copyright 2019 by the American Society of Plant Taxonomists DOI 10.1600/036364419X15562054132974 Date of publication May 30, 2019 Polyphyly of Mesadenus (Orchidaceae, Spiranthinae) and a New Genus from the Espinhaço Range, Southeastern Brazil Gerardo A. Salazar,1,6 Jo~ao A.
    [Show full text]
  • Environmental Assessment
    United States Department of Agriculture Forest Service November 2007 Environmental Assessment Travel Plan Map Revision Elimination of Motorized Cross-Country Travel and Motorized Route Designation Sawtooth National Forest Fairfield Ranger District, Camas County, Idaho Ketchum Ranger District, Blaine County, Idaho Minidoka Ranger District, Twin Falls, Cassia, Power, and Oneida Counties, Idaho; Box Elder County, Utah For Information Contact: Kim Pierson—Team Leader Sawtooth National Forest, Supervisor’s Office, Twin Falls, Idaho (208) 737-3200 http://www.fs.fed.us/r4/sawtooth/projects/ The U.S. Department of Agriculture (USDA) prohibits discrimination in all its programs and activities on the basis of race, color, national origin, gender, religion, age, disability, political beliefs, sexual orientation, or marital or family status. (Not all prohibited bases apply to all programs.) Persons with disabilities who require alternative means for communication of program information (Braille, large print, audiotape, etc.) should contact USDA's TARGET Center at (202) 720-2600 (voice and TDD). To file a complaint of discrimination, write USDA, Director, Office of Civil Rights, Room 326-W, Whitten Building, 14th and Independence Avenue, SW, Washington, DC 20250-9410 or call (202) 720-5964 (voice and TDD). USDA is an equal opportunity provider and employer. Environmental Assessment Sawtooth National Forest Route Designation Table of Contents Acronyms, Abbreviations, and Initialisms .........................................................................................
    [Show full text]
  • New Species Discoveries in the Amazon 2014-15
    WORKINGWORKING TOGETHERTOGETHER TO TO SHARE SCIENTIFICSCIENTIFIC DISCOVERIESDISCOVERIES UPDATE AND COMPILATION OF THE LIST UNTOLD TREASURES: NEW SPECIES DISCOVERIES IN THE AMAZON 2014-15 WWF is one of the world’s largest and most experienced independent conservation organisations, WWF Living Amazon Initiative Instituto de Desenvolvimento Sustentável with over five million supporters and a global network active in more than 100 countries. WWF’s Mamirauá (Mamirauá Institute of Leader mission is to stop the degradation of the planet’s natural environment and to build a future Sustainable Development) Sandra Charity in which humans live in harmony with nature, by conserving the world’s biological diversity, General director ensuring that the use of renewable natural resources is sustainable, and promoting the reduction Communication coordinator Helder Lima de Queiroz of pollution and wasteful consumption. Denise Oliveira Administrative director Consultant in communication WWF-Brazil is a Brazilian NGO, part of an international network, and committed to the Joyce de Souza conservation of nature within a Brazilian social and economic context, seeking to strengthen Mariana Gutiérrez the environmental movement and to engage society in nature conservation. In August 2016, the Technical scientific director organization celebrated 20 years of conservation work in the country. WWF Amazon regional coordination João Valsecchi do Amaral Management and development director The Instituto de Desenvolvimento Sustentável Mamirauá (IDSM – Mamirauá Coordinator Isabel Soares de Sousa Institute for Sustainable Development) was established in April 1999. It is a civil society Tarsicio Granizo organization that is supported and supervised by the Ministry of Science, Technology, Innovation, and Communications, and is one of Brazil’s major research centres.
    [Show full text]
  • Orchid Pollination: Recent Developments from Brazil
    LANKESTERIANA 7: 111-114. 2003. ORCHID POLLINATION: RECENT DEVELOPMENTS FROM BRAZIL RODRIGO B. SINGER Depto Botânica, IB, Unicamp Campinas, SP, Brasil, 13083-970 Because of their remarkable flower morphology, visits. The reasons why Euglossini males collect these orchids are particularly interesting objects of study. compounds are still a matter of speculation (see Most orchid species hold their pollen in discrete Singer & Koehler 2003 for a brief discussion of the packs, so-called pollinia (singular; pollinium). These theories). Yet, it is possible that these compounds are pollinia, together with other pollinium stalks, form somehow used during courtship. In the last six years, more or less complex translatory units, so-called pol- a number of contributions on orchid pollination have linaria (singular: pollinarium). During their visits to been made by Brazilian or Brazilian-based the flowers, orchid pollinators dislodge these polli- researchers. The purpose of this contribution is to naria and deposit their pollen content during succes- give a brief review of these papers and to communi- sive flower visits. Pollinators may be attracted to the cate some – in my opinion - remarkable findings flowers in many ways. Many orchids offer flower which are part of ongoing projects. I have chosen to parts or secretions (nectar, trichomes, wax-like or describe the findings by taxonomic groups. resin-like compounds, aromatic compounds, oils, etc.) that can be gathered/foraged by their pollinators. Subfamily Orchidoideae (including Spiranthoideae, Yet, many orchids “deceive” their pollinators and sensu Dressler 1993). Studied species in this subfami- offer no flower reward. Many orchids are merely ly have been shown to be self-compatible, but polli- “food-frauds”, displaying a set of flower features nator-dependent.
    [Show full text]
  • Phylogenetic Relationships of Discyphus Scopulariae
    Phytotaxa 173 (2): 127–139 ISSN 1179-3155 (print edition) www.mapress.com/phytotaxa/ PHYTOTAXA Copyright © 2014 Magnolia Press Article ISSN 1179-3163 (online edition) http://dx.doi.org/10.11646/phytotaxa.173.2.3 Phylogenetic relationships of Discyphus scopulariae (Orchidaceae, Cranichideae) inferred from plastid and nuclear DNA sequences: evidence supporting recognition of a new subtribe, Discyphinae GERARDO A. SALAZAR1, CÁSSIO VAN DEN BERG2 & ALEX POPOVKIN3 1Departamento de Botánica, Instituto de Biología, Universidad Nacional Autónoma de México, Apartado Postal 70-367, 04510 México, Distrito Federal, México; E-mail: [email protected] 2Universidade Estadual de Feira de Santana, Departamento de Ciências Biológicas, Av. Transnordestina s.n., 44036-900, Feira de Santana, Bahia, Brazil 3Fazenda Rio do Negro, Entre Rios, Bahia, Brazil Abstract The monospecific genus Discyphus, previously considered a member of Spiranthinae (Orchidoideae: Cranichideae), displays both vegetative and floral morphological peculiarities that are out of place in that subtribe. These include a single, sessile, cordate leaf that clasps the base of the inflorescence and lies flat on the substrate, petals that are long-decurrent on the column, labellum margins free from sides of the column and a column provided with two separate, cup-shaped stigmatic areas. Because of its morphological uniqueness, the phylogenetic relationships of Discyphus have been considered obscure. In this study, we analyse nucleotide sequences of plastid and nuclear DNA under maximum parsimony
    [Show full text]