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  1. 1. Pollination BiologyPollination Biology of Orchidsof Orchids Mark Whitten Florida Museum of Natural History University of Florida Ernst Haeckel, 1904 Kunstformen der Natur
  2. 2. Orchid DiversityOrchid Diversity Orchids are the largest family of flowering plantsOrchids are the largest family of flowering plants----~~ 10% of angiosperms10% of angiosperms ~25,000 species described so far (Kew Checklist of~25,000 species described so far (Kew Checklist of Orchids, RBG Kew)Orchids, RBG Kew) New species being described weeklyNew species being described weekly Highest diversity in poorlyHighest diversity in poorly--explored tropicsexplored tropics
  3. 3. Number of NewOrchid Species Described Each Year 0 100 200 300 400 500 600 1978 1980 1982 1984 1986 1989 1991 1993 1995 1997 1999 2001 Source: Cribb, P. 2005. Just how many orchids are there? Proceedings of the 18th WOC
  4. 4. Orchids are HYPERDIVERSE inOrchids are HYPERDIVERSE in montane tropicsmontane tropics Telipogon spp. Of Ecuador: C.H. Dodson
  5. 5. Why are orchids so diverse?Why are orchids so diverse? Inhabit all continents except poles;Inhabit all continents except poles; geologically old family (90geologically old family (90--100 MY ?)100 MY ?) Orchids exploit many different animals asOrchids exploit many different animals as pollinatorspollinators Pollination relationships often highly speciesPollination relationships often highly species-- specificspecific----reproductive isolationreproductive isolation Have obligateHave obligate mycorrhizalmycorrhizal associationsassociations Predominately epiphytic (new niches?)Predominately epiphytic (new niches?)
  6. 6. Habitat diversity fosters biologicalHabitat diversity fosters biological diversitydiversity Teagueia spp. : Lou Jost 21 new species in 2 years in 80 sq. miles Ecuador: Pastaza valley
  7. 7. Precision pollinationPrecision pollination Dichaea trulla
  8. 8. Reproductive IsolationReproductive Isolation How can sympatric,How can sympatric, interfertileinterfertile speciesspecies live together withoutlive together without hybridizing?hybridizing? Highly specificHighly specific pollinators preventpollinators prevent hybridization, andhybridization, and avoid wasting pollenavoid wasting pollen Precise pollenPrecise pollen placement mightplacement might preventprevent hybridizationhybridization
  9. 9. Aspasia epidendroides Floral morphology 3 sepals 2 lateral petals + lip Column (fused male + female parts) Ovary + pedicel
  10. 10. Anther cap Stigma Viscidium Pollinia
  11. 11. Pollinarium Pollinia Stipe Viscidium Pollinated stigma
  12. 12. Pollination rewardsPollination rewards Nectar (insects, birds)Nectar (insects, birds) PollenPollen----No!No! Floral fragrances (maleFloral fragrances (male euglossineeuglossine bees)bees) Resins (used by bees for nest building)Resins (used by bees for nest building) Oils (used byOils (used by anthophoridanthophorid bees to feedbees to feed larvae)larvae)
  13. 13. Pollination syndromesPollination syndromes A bit simplistic, but useful and oftenA bit simplistic, but useful and often predictivepredictive Based upon reward offered and uponBased upon reward offered and upon the behavior and morphology ofthe behavior and morphology of pollinatorspollinators Floral morphology, color, fragrance mayFloral morphology, color, fragrance may allow prediction of probable pollinatorsallow prediction of probable pollinators
  14. 14. Dendrobium cuthbertii Bird pollination -Nectar reward present or absent -Present in Neotropical and Paleotropical orchids Masdevallia rosea Fernandezia subbiflora Maxillaria sophronitis
  15. 15. Masdevallia rosea
  16. 16. Fernandezia subbiflora
  17. 17. Hexisea bidentata
  18. 18. Pollination of Disa chrysostachya by Sunbirds in South Africa S.D. Johnson and M. Brown. 2004. Transfer of pollinaria on bird’s feet: A new pollination system in orchids. Plant Systematics and Evolution Pollinaria are attached to bird’s feet!
  19. 19. Epidendrums can be taxonomically confusing…..
  20. 20. Butterfly pollinationButterfly pollination Epidendrum hybrid swarm, Hollin road, Ecuador ….But especially confusing when they hybridize
  21. 21. Comparettia macroplectron
  22. 22. HawkmothHawkmoth pollinationpollination Amesiella philippenensis
  23. 23. Angraecum rutenbergianum
  24. 24. Aerangis sp.
  25. 25. Dendrophylax fawcettii
  26. 26. Angraecum sequipedale & Xanthopan morganii
  27. 27. Epidendrum lacustre Ecuador
  28. 28. Disa draconis and tanglewing fly Johnson, S.D. and K. E. Steiner. 1997. Long tongued fly pollination and the evolution of spur length in Disa draconis. Evolution 51:45-53.
  29. 29. Pollen deceitPollen deceit Calopogon pulchellus
  30. 30. Pseudopollen Reward or Deceit? Maxillaria with pseudopollen on lip
  31. 31. MaleMale EuglossineEuglossine beebee pollinationpollination Ca. 250 species inCa. 250 species in NeotropicsNeotropics OnlyOnly malesmales collect floral fragrancecollect floral fragrance chemicals and store them in specialchemicals and store them in special hindhind tibialtibial organsorgans Probably use compounds in matingProbably use compounds in mating behavior (sexual selection)behavior (sexual selection)
  32. 32. Long tongues
  33. 33. Female Eufriesea Pollen Basket Stinger
  34. 34. Eulaema meriana (male; no stinger!)
  35. 35. Euglossa dodsoni and Eulaema meriana
  36. 36. Tarsal brush
  37. 37. Hind tibia Cross-section
  38. 38. Eulaema male Courtship display on tree trunk
  39. 39. Clowesia warsczewiczii and Eulaema meriana
  40. 40. Catasetum expansum
  41. 41. Catasetum pollinarium
  42. 42. Catasetum Female flower Cross section
  43. 43. Cycnoches lehmannii & Eulaema cingulata
  44. 44. Cycnoches Male flower
  45. 45. Cycnoches densiflorum Female/hermaphrodite flowers
  46. 46. Stanhopea costaricensis
  47. 47. Eulaema meriana with Stanhopea pollinarium
  48. 48. Sievekingia butcheri
  49. 49. Paphinia neudeckeri
  50. 50. Gongora tricolor With Euglossa cyanura Photo: Kerry Dressler
  51. 51. Gongora hirtzii
  52. 52. Gongora aff. quinquenervis with crystals of methyl-p-methoxycinnamate
  53. 53. Mormodes cartonii Mormodes cartonii
  54. 54. Mormodes pollinarium
  55. 55. Euglossa viridissima: Invading south Florida! Pollinating cultivated Lycaste in Ft. Lauderdale garden Photo: Bob Pemberton
  56. 56. Oil reward flowersOil reward flowers Oils (Oils (triacyltriacyl glyceridesglycerides) produced on) produced on surface of flowers, often in specializedsurface of flowers, often in specialized glandsglands---- elaiophoreselaiophores.. Oils are collected by femaleOils are collected by female anthophoridanthophorid bees; used to provision nest (food forbees; used to provision nest (food for larvae)larvae) Many orchids mimicMany orchids mimic MalpighiaceaeMalpighiaceae
  57. 57. Cyrtochilum With oil gland (elaiophore)
  58. 58. Sigmatostalix sp.
  60. 60. Ornithocephalus sp. Ornithocephalus sp.
  61. 61. Wax or resin rewardWax or resin reward Collected and used by female bees forCollected and used by female bees for nest constructionnest construction Found inFound in MaxillariaMaxillaria andand CymbidiumCymbidium;; poorly studiedpoorly studied
  62. 62. Resin reward: collected by female bees for nest construction Maxillaria notylioglossa Waxy resin crystals
  63. 63. Maxillaria proboscidea RESIN
  64. 64. Resin deceit?Resin deceit? Many Maxillaria species appear to produce resin, but “resin” is simply a shiny callus on lip. Deceit of resin-collecting female bees?
  65. 65. Sexual deceptionSexual deception Based on floral fragrance mimicry ofBased on floral fragrance mimicry of sexual pheromones of pollinatorssexual pheromones of pollinators Usually visual, tactile mimicry, tooUsually visual, tactile mimicry, too Has evolved repeatedly in severalHas evolved repeatedly in several different orchid groupsdifferent orchid groups
  66. 66. Sexual deceit doesn’t have to be perfect--just enough to be convincing!
  67. 67. Chiloglottis formicifera Calochilus paludosa Australia has many pseudocopulatory orchids!
  68. 68. Ophrys Europe
  69. 69. Tolumnia henekenii Hispaniola
  70. 70. Cyrtidiorchis sp. South America Chrysocycnis schlimmii
  71. 71. Mormolyca ringens Mormolyca schweinfurthiana
  72. 72. Trigonidium grande
  73. 73. Telipogon sp. Sexual deceit— Pollination by male Tachinid flies
  74. 74. Telipogon sp.
  75. 75. Telipogon sp.
  76. 76. Trichoceros antennifer
  77. 77. PseudocopulatoryPseudocopulatory pollination ofpollination of LepanthesLepanthes gleicensteiniigleicensteinii Mario Blanco & Gabriel Barbosa, Ann. Bot. 95:763-772. 720+ spp.
  78. 78. Food or Brood site deceptionFood or Brood site deception Not well studied, but probably not rareNot well studied, but probably not rare Possible examples:Possible examples: DraculaDracula ---- pollinated bypollinated by DrosophilidDrosophilid flies; mimicry of yeast?flies; mimicry of yeast? PaphiopediumPaphiopedium ---- some pollinated bysome pollinated by syrphidsyrphid flies; mimicry of aphids (prey forflies; mimicry of aphids (prey for larvae)?larvae)?
  79. 79. Paphiopedilum glanduliferum
  80. 80. Dracula sp.
  81. 81. Dracula chestertonii Lip mimics mushroom --visual, chemical mimicry?
  82. 82. Lip of Dracula sp. at Finca Dracula, Panama
  83. 83. Fly pollination inFly pollination in PaleotropicalPaleotropical BulbophyllumBulbophyllum andand CirrhopetalumCirrhopetalum
  84. 84. ““Motel 6Motel 6”” syndromesyndrome Bees seek out flowers to sleep inBees seek out flowers to sleep in overnightovernight No obvious food rewardNo obvious food reward Chemical attractants?Chemical attractants? Confirmed only for European genusConfirmed only for European genus SerapiasSerapias
  85. 85. Serapias lingua
  86. 86. Trap mechanismsTrap mechanisms Found in association with variousFound in association with various reward/attractant syndromesreward/attractant syndromes Pollinators are lured into an area theyPollinators are lured into an area they cancan’’t escape, then forced to exit pastt escape, then forced to exit past the columnthe column Or, the pollinator may be forcibly thrownOr, the pollinator may be forcibly thrown against the columnagainst the column
  87. 87. Selenipedium antioquiense
  88. 88. Porroglossum eduardii Lip open Lip closed Active trap mechanisms Some pleurothallid flowers have sensitive, motile lips (similar to Venus Fly Trap) Insect landing on lip triggers lip to close, lifting insect into contact with column.
  89. 89. Coryanthes sp. Fragrance attractant; Liquid-filled bucket trap
  90. 90. Coryanthes Cross-section
  91. 91. Nectar DeceitNectar Deceit Relatively few orchids actually produceRelatively few orchids actually produce a nectar reward (energeticallya nectar reward (energetically expensive? Longexpensive? Long--distance pollendistance pollen movement?)movement?) Most have emptyMost have empty nectariesnectaries; deceive; deceive pollinatorspollinators Might be mimics of other nectarMight be mimics of other nectar-- producing nonproducing non--orchids in same habitatorchids in same habitat
  92. 92. Cochleanthes anatona Sepaline false nectary
  93. 93. Oil deceitOil deceit Many species ofMany species of OncidiumOncidium do not offerdo not offer an oil reward, but are thought to mimican oil reward, but are thought to mimic flowers of the nonflowers of the non--orchidaceousorchidaceous MalpighiaMalpighia (an oil(an oil--rewardreward eudicoteudicot) or) or CalceolariaCalceolaria.. Typical yellow/brownTypical yellow/brown ““oncidioidoncidioid”” flowersflowers occur in many unrelatedoccur in many unrelated cladesclades ofof subtribesubtribe OncidiinaeOncidiinae Taxonomic confusion!Taxonomic confusion!
  94. 94. Malpighia spp.--- Oil reward flower Model for many yellow “Oncidiums”?
  95. 95. OTHER MODELS ??? CALCEOLARIA & OTOGLOSSUM HARLINGII Calceolaria is an oil reward flower; sympatric with “Oncidiums” at higher elevations in the Andes
  96. 96. Chelyorchis ampliata Tolumnia urophylla Tolumnia guianensis Orchids that are good at fooling pollinators are also good at fooling orchid taxonomists!
  97. 97. Rhizanthella gardneri Termite pollination
  98. 98. PlatantheraPlatanthera: Using Molecular Phylogenetics to: Using Molecular Phylogenetics to Study Adaptation to Different PollinatorsStudy Adaptation to Different Pollinators Work ofWork of HapemanHapeman and Inoue (1997)and Inoue (1997) Used DNA sequencing to produce aUsed DNA sequencing to produce a phylogenetic tree; usedphylogenetic tree; used characterscharacters independent of floral traitsindependent of floral traits Then, mapped floral characters ontoThen, mapped floral characters onto tree, allowing evolution of traits to betree, allowing evolution of traits to be tracedtraced
  99. 99. Floral Diversity in Platanthera
  100. 100. PlatantheraPlatanthera: Take: Take--home messageshome messages PollinationPollination--related floral traits (color,related floral traits (color, size, spur length, etc.) undergo rapidsize, spur length, etc.) undergo rapid and frequent evolutionary changeand frequent evolutionary change Floral traits areFloral traits are ““homoplasioushomoplasious””=evolutionarily labile=evolutionarily labile Orchid classifications shouldOrchid classifications should NOTNOT bebe based primarily on floral characters!based primarily on floral characters!
  101. 101. Emphasis on gross floral characters can result in incorrect classifications! Dendrophylax fawcetti “Campylocentrum” filiformis Dendrophylax filiformis
  102. 102. CoevolutionCoevolution?? Orchids are highly adapted to utilizeOrchids are highly adapted to utilize specific pollinators, butspecific pollinators, but…….. Do orchids affect the evolution of theirDo orchids affect the evolution of their animal pollinators?animal pollinators? ““Orchid and bee marching petal inOrchid and bee marching petal in tarsus down the aisle oftarsus down the aisle of coevolutioncoevolution””??
  103. 103. CoevolutionCoevolution?? NoNo---- no evidence that orchids affect theno evidence that orchids affect the evolution of morphology or behavior ofevolution of morphology or behavior of their pollinatorstheir pollinators Most orchids are rareMost orchids are rare----not the primarynot the primary source of nectar or other rewards forsource of nectar or other rewards for their pollinatorstheir pollinators ProbablyProbably ““parasitesparasites”” on otheron other plant/pollinator systemsplant/pollinator systems
  104. 104. Is Pollinator SpecificityIs Pollinator Specificity OverOver--estimated?estimated? How many observations are needed toHow many observations are needed to document pollinator set for an orchid?document pollinator set for an orchid? What about seasonal, geographicWhat about seasonal, geographic variation?variation? For most species, we donFor most species, we don’’t havet have enough observations to adequatelyenough observations to adequately measure pollinator specificitymeasure pollinator specificity
  105. 105. How many species? How many polllinators, fungal partners? Conservation— Ex situ, in situ?
  106. 106. What can you do to conserveWhat can you do to conserve orchids?orchids? $45 buys 1 acre of montane rainforest– makes a great birthday or holiday gift Ecominga Foundation: purchasing land to conserve Ecuador’s orchid-rich habitats: www.loujost.com
  107. 107. AcknowledgementsAcknowledgements National Science FoundationNational Science Foundation American Orchid SocietyAmerican Orchid Society Marie Selby Botanical GardenMarie Selby Botanical Garden Atlanta Botanical GardenAtlanta Botanical Garden Many commercial & private growersMany commercial & private growers