Darwin’s Sexy Orchids:Darwin’s Sexy Orchids: Case Study Case Study OnOn TheThe Origin of Species by Origin of Species by

Means of Natural Means of Natural SelectionSelection

Harvey Brenneise, Harvey Brenneise, Associate Dean for Research Associate Dean for Research ServicesServices

University of Southern Mississippi (Hattiesburg) &University of Southern Mississippi (Hattiesburg) &Research Associate,Research Associate,

Rancho Santa Ana Botanic Garden (CA)Rancho Santa Ana Botanic Garden (CA)

Charles Robert Darwin (1809-Charles Robert Darwin (1809-82)82)

Childhood and youthChildhood and youth EducationEducation Voyage of the Voyage of the BeagleBeagle MarriageMarriage Move to the countryMove to the country ExperimentsExperiments

Life (cont.)Life (cont.) PublicationsPublications Scientific friendsScientific friends Publication of Publication of Origin of SpeciesOrigin of Species Publication of Publication of OrchidsOrchids Life after OrchidsLife after Orchids

Darwin as a childDarwin as a child

““By the time I went to … school By the time I went to … school my taste for natural history, and my taste for natural history, and more especially for collection, was more especially for collection, was well developed. . . . The passion well developed. . . . The passion for collecting, which leads a man for collecting, which leads a man to be a systematic naturalist, a to be a systematic naturalist, a virtuoso or a miser, was very virtuoso or a miser, was very strong in me, and was clearly strong in me, and was clearly innate, as none of my sisters or innate, as none of my sisters or brother ever had this taste.”brother ever had this taste.”

University of Edinburgh (1825-University of Edinburgh (1825-27)27)

Cambridge (1828-31)Cambridge (1828-31)

Rev. John Stevens Henslow Rev. John Stevens Henslow (1796-1861)(1796-1861)

Darwin as a young manDarwin as a young man

Orchis morio Orchis morio L. (1753)L. (1753)

Voyage of the Voyage of the H.M.S. BeagleH.M.S. Beagle(1831-36)(1831-36)

Capt. Robert FitzRoy (1805-Capt. Robert FitzRoy (1805-65)65)

Darwin’s room on boardDarwin’s room on board

Voyage of the Voyage of the Beagle, Beagle, 1831-1831-3636

OrchidaceaeOrchidaceae taxa collected: taxa collected:by date of collectionby date of collection

Epidendrum difformeEpidendrum difforme Jacq. [1760] (Brazil, Jacq. [1760] (Brazil, May/June 1832) May/June 1832)

Codonorchis lessoniiCodonorchis lessonii Lindl. [1840] (Tierra Lindl. [1840] (Tierra del Fuego, Jan./Feb. 1833)del Fuego, Jan./Feb. 1833)

Chloraea gaudichaudiiChloraea gaudichaudii Brongn. [1834] Brongn. [1834] (Argentina, Dec. 1833 or Jan. 1834)(Argentina, Dec. 1833 or Jan. 1834)

Chloraea magellanicaChloraea magellanica Hook. f. [1846] Hook. f. [1846] (Chile, Jan. 30, 1834) (Chile, Jan. 30, 1834) IsolectotypeIsolectotype

Bipinnula fimbriataBipinnula fimbriata I.M. Johnst. [1929] I.M. Johnst. [1929] (Chile, Aug. 1834)(Chile, Aug. 1834)

Epidendrum spicatumEpidendrum spicatum Hook. f. [1847] Hook. f. [1847] (Galapagos, Oct. 1835) (Galapagos, Oct. 1835) Holotype Holotype [Endemic][Endemic]

““Started about ½ after Started about ½ after six and passed over six and passed over scorching plains—cactuses scorching plains—cactuses and other succulent plants: and other succulent plants: on the stunted and on the stunted and decaying trees beautiful decaying trees beautiful parasites—orchids with a parasites—orchids with a delicious smell.” delicious smell.”

Epidendrum difforme Epidendrum difforme (May/June (May/June 1832)1832)

Codonorchis lessoniiCodonorchis lessonii (Jan./Feb. (Jan./Feb. 1833)1833)

Darwin’s field notesDarwin’s field notes

Chloraea gaudichaudii Chloraea gaudichaudii (Dec. 1833-Jan. 1834)(Dec. 1833-Jan. 1834)

Jan. 30, 1834Jan. 30, 1834

Bipinnula fimbriata Bipinnula fimbriata (Aug. (Aug. 1834)1834)

OrchidaceaeOrchidaceae in the Galapagos in the GalapagosWiggins and Porter, Wiggins and Porter, Flora of the Galapagos IslandsFlora of the Galapagos Islands

(1971)(1971) Cranichis schlimiiCranichis schlimii Rchb. f. (1876) Rchb. f. (1876) Epidendrum spicatumEpidendrum spicatum Hook. f. (1847) Hook. f. (1847)

EndemicEndemic Erythrodes weberianaErythrodes weberiana Garay (1970) Garay (1970)

Endemic Endemic (Isabella)(Isabella) Govenia utriculataGovenia utriculata (Sw.) Lindl. (1839) (Sw.) Lindl. (1839) Habenaria alataHabenaria alata Hook. (1826) Hook. (1826) Habenaria monorrhizaHabenaria monorrhiza (Sw.) Rchb. f. (1885) (Sw.) Rchb. f. (1885) Ionopsis utriculariodesIonopsis utriculariodes (Sw) Lindl. (1826) (Sw) Lindl. (1826) Liparis nervosaLiparis nervosa (Thunb.) Lindl. (1830) (Thunb.) Lindl. (1830) Ponthieva maculataPonthieva maculata Lindl. (1845) Lindl. (1845) Presscottia oliganthaPresscottia oligantha (Sw.) Lindl. 1840 (Sw.) Lindl. 1840 Tropidia polystachyaTropidia polystachya (Sw.) Ames (1908) (Sw.) Ames (1908)

Epidendrum spicatum Epidendrum spicatum (Oct. (Oct. 1835)1835)

Trans. Linn. Soc. London 20(2): Trans. Linn. Soc. London 20(2): 180. 1847 [1851 publ. 11 Dec 180. 1847 [1851 publ. 11 Dec

1847]1847]

What next?What next? First love, Fanny Owens, had become First love, Fanny Owens, had become

engaged while on the voyage.engaged while on the voyage. What to do? The practical thing.What to do? The practical thing.

Reasons to get marriedReasons to get married Children (if it please God) Children (if it please God) Constant companion (& friend in old age) Constant companion (& friend in old age)

who will feel interested in one, object to be who will feel interested in one, object to be beloved & played with, better than a dog beloved & played with, better than a dog anyhowanyhow

Home, & someone to take care of houseHome, & someone to take care of house Charms of music & female chit-chatCharms of music & female chit-chat These things good for one's healthThese things good for one's health Forced to visit & receive relations Forced to visit & receive relations but terrible but terrible

loss of time.loss of time. — —

Reasons not to stay singleReasons not to stay single My God, it is intolerable to think of My God, it is intolerable to think of

spending ones whole life, like a neuter spending ones whole life, like a neuter bee, working, working, & nothing after all. bee, working, working, & nothing after all. — No, no won't do. — No, no won't do.

Imagine living all one's day solitarily in Imagine living all one's day solitarily in smoky dirty London Housesmoky dirty London House

Only picture to yourself a nice soft wife on Only picture to yourself a nice soft wife on a sofa with good fire, & books & music a sofa with good fire, & books & music perhaps — Compare this vision with the perhaps — Compare this vision with the dingy reality of Grt. Marlbro' St.dingy reality of Grt. Marlbro' St.

No children, (no second life), no one to No children, (no second life), no one to care for one in old age.— What is the use care for one in old age.— What is the use of working 'in' without sympathy from near of working 'in' without sympathy from near & dear friends—who are near & dear & dear friends—who are near & dear friends to the old, except relativesfriends to the old, except relatives

Reasons not to get marriedReasons not to get married Freedom to go where one liked — choice of Freedom to go where one liked — choice of

Society & Society & little of itlittle of it Conversation of clever men at clubsConversation of clever men at clubs Not forced to visit relatives, & to bend in every Not forced to visit relatives, & to bend in every

trifletrifle to have the expense & anxiety of children — to have the expense & anxiety of children —

perhaps quarellingperhaps quarelling Loss of time.Loss of time. — cannot read in the Evenings — cannot read in the Evenings Fatness & idleness — Anxiety & responsibility Fatness & idleness — Anxiety & responsibility Less money for books &c — if many children Less money for books &c — if many children

forced to gain one's bread. — (But then it is forced to gain one's bread. — (But then it is very bad for ones health to work too much)very bad for ones health to work too much)

Perhaps my wife wont like London; then the Perhaps my wife wont like London; then the sentence is banishment & degradation into sentence is banishment & degradation into indolent,indolent, idle fool — idle fool —

SummarySummary

The Governor says soon for otherwise The Governor says soon for otherwise bad if one has children — one's bad if one has children — one's character is more flexible —one's character is more flexible —one's feelings more lively & if one does not feelings more lively & if one does not marry soon, one misses so much good marry soon, one misses so much good pure happiness. —pure happiness. —But then if I married tomorrow: there But then if I married tomorrow: there would be an infinity of trouble & expense would be an infinity of trouble & expense in getting & furnishing a house, —in getting & furnishing a house, —fighting about no Society —morning calls fighting about no Society —morning calls — awkwardness —loss of time every — awkwardness —loss of time every day. (without one's wife was an angel, & day. (without one's wife was an angel, & made one keep industrious). — made one keep industrious). —

When? Jan. 29, 1839When? Jan. 29, 1839Then how should I manage all my business Then how should I manage all my business

if I were obliged to go every day walking with if I were obliged to go every day walking with one’s my wife. — Eheu!! I never should know one’s my wife. — Eheu!! I never should know French, — or see the Continent — or go to French, — or see the Continent — or go to America, or go up in a Balloon, or take solitary America, or go up in a Balloon, or take solitary trip in Wales — poor slave. — you will be worse trip in Wales — poor slave. — you will be worse than a negro — And then horrid poverty, (without than a negro — And then horrid poverty, (without one's wife was better than an angel & had one's wife was better than an angel & had money) — money) —

Never mind my boy — Cheer up — One Never mind my boy — Cheer up — One cannot live this solitary life, with groggy old age, cannot live this solitary life, with groggy old age, friendless & cold, & childless staring one in ones friendless & cold, & childless staring one in ones face, already beginning to wrinkle. — Never mind, face, already beginning to wrinkle. — Never mind, trust to chance —keep a sharp look out — There trust to chance —keep a sharp look out — There is many a happy slave — is many a happy slave —

Emma Wedgwood Darwin Emma Wedgwood Darwin (1808-96)(1808-96)

Emma a paragon of virtueEmma a paragon of virtue LoyalLoyal Cared for him when he was sickCared for him when he was sick Read to him at nightRead to him at night Fundamentalist religious viewsFundamentalist religious views Little sense of humorLittle sense of humor Angelic!Angelic!

Move to Kent (Down House)--Move to Kent (Down House)--18421842

Darwin’s scientific Darwin’s scientific “buddies”“buddies”

Joseph Dalton Hooker (1817-Joseph Dalton Hooker (1817-1911)1911)

Darwin, Hooker, Charles Darwin, Hooker, Charles LyellLyell

Asa Gray (1810-88)Asa Gray (1810-88)

Alfred Russel Wallace (1823-Alfred Russel Wallace (1823-1913)1913)

Experiments (“little Experiments (“little triumphs”)triumphs”)

““Are you going to beat Dr. Hooker?”Are you going to beat Dr. Hooker?” Experiment BookExperiment Book (1856-) (1856-) Weed Garden (experimental theorist)Weed Garden (experimental theorist) Gardeners’ ChronicleGardeners’ Chronicle ““I am like a gambler, & love a wild I am like a gambler, & love a wild

experiment.”experiment.” Field studies and practical scientific Field studies and practical scientific

investigations both an amusement and investigations both an amusement and part of his research program part of his research program

Books--to Books--to OriginOrigin The zoology of the voyage of H.M.S. The zoology of the voyage of H.M.S.

BeagleBeagle (1838-43) (1838-43) The narrative of the voyages of H.M. The narrative of the voyages of H.M.

Ships Adventure and BeagleShips Adventure and Beagle [Journal [Journal of Researches] (3 vols.) (1839, 1845, of Researches] (3 vols.) (1839, 1845, 1860, 1880, 1890, 1905)1860, 1880, 1890, 1905)

Geology of the voyage of the Beagle, Geology of the voyage of the Beagle, (1842-46,1876-76, 1889-90)(1842-46,1876-76, 1889-90)

Books—to Books—to Origin (cont.)Origin (cont.) A monograph of the sub-class A monograph of the sub-class

Cirripedia, with figures of all the Cirripedia, with figures of all the species species (2 vols.) (1851-54)(2 vols.) (1851-54)

A monograph on the fossil Lepadidae, A monograph on the fossil Lepadidae, or, pedunculated cirripedes of Great or, pedunculated cirripedes of Great BritainBritain. (2 vols.) (1851-54). (2 vols.) (1851-54)

On the origin of speciesOn the origin of species (1859, 1860 (1859, 1860 [1[1stst American], 1860, 1861, 1866, American], 1860, 1861, 1866, 1869, 1872, 1876) 1869, 1872, 1876)

What next?What next? Variation among domestic plants Variation among domestic plants

and animals?and animals? DroseraDrosera (sundew)? (sundew)? Dimorphism among primulas?Dimorphism among primulas?

Carl Linnaeus (Carl von Linné) in 1746 Carl Linnaeus (Carl von Linné) in 1746 had portrayed a flower as a marriage had portrayed a flower as a marriage

bed of 9 gentlemen and 1 lady.bed of 9 gentlemen and 1 lady.

Christian Konrad Sprengel (1750-1816), Christian Konrad Sprengel (1750-1816), Das Das entdeckte Geheimnis der Natur im Bau und entdeckte Geheimnis der Natur im Bau und

in der Befruchtung der Blumen in der Befruchtung der Blumen [floral [floral ecology]ecology]

Orchids!Orchids! Summer 1860: “Orchis Bank”Summer 1860: “Orchis Bank” Appeal to Appeal to Gardeners’ Chronicle Gardeners’ Chronicle and repliesand replies 1861: All rest of year Orchid Book (diary)1861: All rest of year Orchid Book (diary) To Hooker: "I am intensely interested on To Hooker: "I am intensely interested on

subject, just as at a game of chess.” In subject, just as at a game of chess.” In September, he "dissected with the September, he "dissected with the greatest interest.” greatest interest.”

"The contrivances for insect fertilisation in "The contrivances for insect fertilisation in Orchids are multiform & truly wonderful & Orchids are multiform & truly wonderful & beautiful.”beautiful.”

Darwin’s periodical articles on Darwin’s periodical articles on orchids prior to publication oforchids prior to publication of

OrchidsOrchids ““Fertilisation of British orchids by insect Fertilisation of British orchids by insect

agency.” agency.” Gardeners' ChronicleGardeners' Chronicle no. 23 (9 no. 23 (9 June 1860): 528 and no. 6 (9 Feb. 1861): June 1860): 528 and no. 6 (9 Feb. 1861): 122. 122.

““Orchids, Fertilization of.” Orchids, Fertilization of.” Gardeners' Gardeners' ChronicleChronicle no. 37 (14 Sept. 1861): 831. no. 37 (14 Sept. 1861): 831.

““On the three remarkable sexual forms of On the three remarkable sexual forms of Catasetum tridentatumCatasetum tridentatum, an orchid in the , an orchid in the possession of the Linnean Society.” [Read possession of the Linnean Society.” [Read 3 April] 3 April] Proceedings of the Linnean Society Proceedings of the Linnean Society of London. Botanyof London. Botany 6: 151-157. 1862 6: 151-157. 1862

Wrote to Asa Gray prior to publication: “It Wrote to Asa Gray prior to publication: “It really seems to me incredibly monstrous to really seems to me incredibly monstrous to look at an orchid as created as we now see look at an orchid as created as we now see it. Every part reveals modification on it. Every part reveals modification on modification.” modification.”

Publication was delayed because of illness, Publication was delayed because of illness, but Darwin looked at it as “a hobby-horse, but Darwin looked at it as “a hobby-horse, which has given me great pleasure to which has given me great pleasure to ride.” ride.”

British orchids examined by Darwin (2ndBritish orchids examined by Darwin (2nd ed.) ed.)

[15 genera by Darwin’s count][15 genera by Darwin’s count]illustrated in illustrated in boldbold

Aceras anthropophora, A. longibracteata Aceras anthropophora, A. longibracteata [France][France] Caladenia dimorphaCaladenia dimorpha Cephalanthera ensifolia, Cephalanthera ensifolia, C. grandifloraC. grandiflora (left out Cypripedium) (left out Cypripedium) Epipactus latifoliaEpipactus latifolia, E., microphylla, , E., microphylla, E. palustrisE. palustris, E. , E.

purpurata, E. rubiginosa, E. viridiflorapurpurata, E. rubiginosa, E. viridiflora Epipogium gmeliniEpipogium gmelini Goodyera discolor, G. pubescens, G. repensGoodyera discolor, G. pubescens, G. repens Gymnadenia albida, Gymnadenia albida, G. conopsea,G. conopsea, G. odoratissima, G. odoratissima,

G. tridentataG. tridentata Habenaria bifolia,Habenaria bifolia, H. chlorantha H. chlorantha Herminium monorchisHerminium monorchis Listera cordata, Listera cordata, L. ovataL. ovata Malaxis paludosaMalaxis paludosa

More “British” orchidsMore “British” orchids Neotinea intacta Neotinea intacta [Italy][Italy] Neottia nidus-avisNeottia nidus-avis Nigritella angustifolia Nigritella angustifolia [alpine][alpine] Ophrys apifera,Ophrys apifera, O. arachnitesO. arachnites, O. aranifera, , O. aranifera, O. O.

musciferamuscifera, O. scolopas, O. scolopas Orchis fusca, O. hircina, O. latifolia, O. maculata, Orchis fusca, O. hircina, O. latifolia, O. maculata,

O. masculaO. mascula, O. morio, , O. morio, O. pyramidalisO. pyramidalis, O. , O. ustulataustulata

Peristylus viridisPeristylus viridis Platanthera chlorantha, P. dilatata, P. flava, P. Platanthera chlorantha, P. dilatata, P. flava, P.

hookeri, P. hyperboreahookeri, P. hyperborea Ptorostylis longifoliaPtorostylis longifolia [Australia][Australia] Pogonia ophioglossoidesPogonia ophioglossoides Serapias cordigera Serapias cordigera [France][France] Spiranthes australis, Spiranthes australis, S. autumnalisS. autumnalis, S. cernua, S. , S. cernua, S.

gracilisgracilis

British specimens (1British specimens (1stst ed.) donated ed.) donated by:by:

Mr. Bateman, Mr. Bateman, Goodyera discolor Goodyera discolor (foreign)(foreign) Dr. Battersby (Torquay), Dr. Battersby (Torquay), Spiranthesis autumnalisSpiranthesis autumnalis F. Bond (South Kent) F. Bond (South Kent) HadenaHadena dentina, H. plusiadentina, H. plusia; ;

Orchis fuscaOrchis fusca, moths with pollinia attached, moths with pollinia attached Rev. G. Gordon (Elgin) Rev. G. Gordon (Elgin) Goodyera repensGoodyera repens Mr. Malden (South Kent) Mr. Malden (South Kent) Orchis fuscaOrchis fusca A. G. More (Bembridge, Isle of Wight), A. G. More (Bembridge, Isle of Wight), Epipactis Epipactis

palustrispalustris, , Ophrys apiferaOphrys apifera (sent report of field (sent report of field work), work), Spiranthesis autumnalisSpiranthesis autumnalis

G. Chichester Oxenden (Broome Park, South G. Chichester Oxenden (Broome Park, South Kent) Kent) Neottia nidus-avisNeottia nidus-avis, , Ophyris araniferaOphyris aranifera, , Orchis ustulataOrchis ustulata

Mr. Wallis (Hartfield, Sussex) Mr. Wallis (Hartfield, Sussex) Malaxis paludosaMalaxis paludosa Professor Westwood (bees with pollinia attached)Professor Westwood (bees with pollinia attached)

Orchis masculaOrchis mascula L. (1755) L. (1755)

O. MasculaO. Mascula pollinium pollinium

““A poet might imagine, that A poet might imagine, that whilst the pollinia are borne whilst the pollinia are borne from flower to flower through from flower to flower through the air, adhering to a moth's the air, adhering to a moth's body, they voluntarily and body, they voluntarily and eagerly place themselves, in eagerly place themselves, in each case, in that exact each case, in that exact position in which alone they position in which alone they can hope to gain their wish and can hope to gain their wish and perpetuate their race.”perpetuate their race.”

Bilder ur Nordens FloraBilder ur Nordens Flora (1905) (1905)

Ophrys apiferaOphrys apifera Huds. (1762) Huds. (1762)

Flora von Deutchland, Osterreich under der SchweizFlora von Deutchland, Osterreich under der Schweiz (1885) (1885)

Atlas des plantes de France Atlas des plantes de France (1891)(1891)

““Foreign” orchids examined by DarwinForeign” orchids examined by Darwin22ndnd ed. ed. Illustrated inIllustrated in bold bold

Acropera loddigesii, A. luteolaAcropera loddigesii, A. luteola Aerides cornutum, A. odorata, A. virensAerides cornutum, A. odorata, A. virens Angraecum distichum, A. eburneum, A. Angraecum distichum, A. eburneum, A.

sesquipedalesesquipedale Bulbophyllum barbigerum, B. cocinum, B. Bulbophyllum barbigerum, B. cocinum, B.

cupreum, B. rhizophoraecupreum, B. rhizophorae Bonatea speciosaBonatea speciosa BrassiaBrassia Calanthe dominii, Calanthe dominii, C. masuca,C. masuca, C. C.

veratrifolia, C. vestitaveratrifolia, C. vestita Catasetum callosum, C. luridum, C. Catasetum callosum, C. luridum, C.

mentosum, C. planicips, mentosum, C. planicips, C. saccatum,C. saccatum, C. C. tabulare, C. tabulare, C. tridentatumtridentatum

More “foreign” orchidsMore “foreign” orchids CattleyaCattleya crispa crispa ChysisChysis Coelogyne cristataCoelogyne cristata Coryanthes fieldingii, C. macrantha, Coryanthes fieldingii, C. macrantha, C. C.

speciosaspeciosa, C. triloba, C. triloba Cycnoches egertonianum, Cycnoches egertonianum, C. ventricosumC. ventricosum Cymbidium giganteumCymbidium giganteum Cypripedium Cypripedium acaule, C. barbatum, C. acaule, C. barbatum, C.

calceolus, C. candidum, C. pubescens, C. calceolus, C. candidum, C. pubescens, C. purpuratumpurpuratum

Dendrobium bigbbum, D. cretaceum, Dendrobium bigbbum, D. cretaceum, D. D. chrysanthemumchrysanthemum, D. formosum, D. , D. formosum, D. speciosumspeciosum

Disa cornuta, D. grandiflora, D. macranthaDisa cornuta, D. grandiflora, D. macrantha

And more!And more! Epidendrum cochleatum, E. floribundum, E. Epidendrum cochleatum, E. floribundum, E.

glaucumglaucum Eulophia viridisEulophia viridis Evelyna carivataEvelyna carivata Galeandra funkiiGaleandra funkii Gongora atro-purpurea, G. maculata, G. Gongora atro-purpurea, G. maculata, G.

truncatatruncata Laelia cinnabarinaLaelia cinnabarina LeptotesLeptotes Lycaste skinneriiLycaste skinnerii Masdevallia fenestrataMasdevallia fenestrata

More!More! Maxillaria ornithorhynchaMaxillaria ornithorhyncha Microstylus rhediiMicrostylus rhedii Miltonia clowesiiMiltonia clowesii MonachanthusMonachanthus viridis viridis Mormodes ignea,Mormodes ignea, M. luxata M. luxata Myanthus Myanthus barbatusbarbatus OrnithocephalusOrnithocephalus Phaius grandifoliusPhaius grandifolius Phalaenopsis amabilis, P. grandifloraPhalaenopsis amabilis, P. grandiflora Pleurothallis ligulata, P. proliferaPleurothallis ligulata, P. prolifera

That’s all, folks!That’s all, folks!

Rodriguezia secunda, R. suaveolensRodriguezia secunda, R. suaveolens Sarcanthus parishii, S. teretifoliusSarcanthus parishii, S. teretifolius Selenipedium palmifoliumSelenipedium palmifolium Sobralia macranthaSobralia macrantha Stanhopea devoniensis, S. oculataStanhopea devoniensis, S. oculata Stelis racemifloraStelis racemiflora Thelmitra carnea, T. longifloraThelmitra carnea, T. longiflora Vanilla aromaticaVanilla aromatica Zygopetalum mackaiZygopetalum mackai

““Foreign” specimens donated for 1Foreign” specimens donated for 1stst ed. by: ed. by: [the kindness of many friends and [the kindness of many friends and

strangers], 43 “exotic” genera “well strangers], 43 “exotic” genera “well dispersed through the subfamilies of the dispersed through the subfamilies of the

vast Orchidean series”vast Orchidean series” Joseph Hooker, “has never become weary of Joseph Hooker, “has never become weary of

sending me specimens from the Royal sending me specimens from the Royal Gardens at Kew”Gardens at Kew”

James Veitch, jun., “many beautiful Orchids”James Veitch, jun., “many beautiful Orchids” R. Parker, “extremely valuable series of R. Parker, “extremely valuable series of

forms”forms” Lady Dorothy Nevill, “most kindly placed Lady Dorothy Nevill, “most kindly placed

her magnificent collection of Orchids at my her magnificent collection of Orchids at my disposal”disposal”

Donors (cont.)Donors (cont.) Mr. Rucker (West Hill, Wandsworth), “sent Mr. Rucker (West Hill, Wandsworth), “sent

me repeatedly large spikes of Catasetum, me repeatedly large spikes of Catasetum, a Mormodes of extreme value to me, and a Mormodes of extreme value to me, and some Dendrobiums”some Dendrobiums”

Mr. Bateman, “a number of interesting Mr. Bateman, “a number of interesting forms, including the wonderful forms, including the wonderful Angraecum Angraecum sesquipedale”sesquipedale”

Mr. Turnbull (Down), “free use of his hot-Mr. Turnbull (Down), “free use of his hot-houses” and “giving me some interesting houses” and “giving me some interesting orchids, and his gardener, Mr Horwood, for orchids, and his gardener, Mr Horwood, for his aid in some of my observations”his aid in some of my observations”

Dr. Lindley, “fresh and dried specimens”Dr. Lindley, “fresh and dried specimens”

Lady Dorothy Nevill (1826-Lady Dorothy Nevill (1826-1913)1913)

Relationship with DarwinRelationship with Darwin Liked to put signed portraits of Liked to put signed portraits of

famous scientists on her wallsfamous scientists on her walls Flirtatious correspondence “filled Flirtatious correspondence “filled

with double entendre, describing the with double entendre, describing the orchids' body parts and the methods orchids' body parts and the methods by which these flowers fertilize each by which these flowers fertilize each other."other."

Wisdom from Lady Dorothy Wisdom from Lady Dorothy Nevill:Nevill:

““The real art of The real art of conversation is not only to conversation is not only to say the right thing at the say the right thing at the right place, but to leave right place, but to leave unsaid the wrong thing at the unsaid the wrong thing at the tempting moment." tempting moment."

John Lindley (1799-1865)John Lindley (1799-1865)

Jas. Bateman (1811-97)Jas. Bateman (1811-97)

James Veitch, Jr. (1815-69)James Veitch, Jr. (1815-69)

Catasetum saccatumCatasetum saccatum Lindl. Lindl. (1840)(1840)

Darwin calledDarwin called Catasetum Catasetum “the “the most remarkable of all Orchids", and most remarkable of all Orchids", and showed how in these flowers "as showed how in these flowers "as throughout nature, pre-existing throughout nature, pre-existing structures and capacities [had been] structures and capacities [had been] utilised for new purposes". utilised for new purposes".

Catasetum tridendatumCatasetum tridendatum showed showed its “truly marvelous” mechanism, by its “truly marvelous” mechanism, by which it shot out a pollinium at any which it shot out a pollinium at any insect touching a part of the flower insect touching a part of the flower with “sticky gland always foremost.” with “sticky gland always foremost.”

Darwin imitated the action of Darwin imitated the action of an insect touching the flower’s an insect touching the flower’s “antenna” using a whalebone “antenna” using a whalebone spring. spring.

"I touched the antennæ of "I touched the antennæ of C. C. callosumcallosum whilst holding the flower whilst holding the flower at about a yard's distance from at about a yard's distance from the window, and the pollinium hit the window, and the pollinium hit the pane of glass, and adhered to the pane of glass, and adhered to the smooth vertical surface by its the smooth vertical surface by its adhesive disc.”adhesive disc.”

Sertum OrchidaceumSertum Orchidaceum (1838)(1838)

CatasetumCatasetum experiments experiments Fall onto a table from a height of 2-3 in.Fall onto a table from a height of 2-3 in. Cut off with a crash with scissorsCut off with a crash with scissors Deep pricks of the column and stigmatic Deep pricks of the column and stigmatic

chamberchamber A blow hard enough to knock off the A blow hard enough to knock off the

anther (an accident)anther (an accident) Press hard on pedicel and rostellumPress hard on pedicel and rostellum Nothing works except “violence” to the Nothing works except “violence” to the

antennae (not including stream of air, cold antennae (not including stream of air, cold water or human hair)water or human hair)

Curtis’Curtis’’ v. 131 (1905) as ’ v. 131 (1905) as C. C. christyanumchristyanum

Catasetum macrocarpum Catasetum macrocarpum [=tridentatum[=tridentatum]: ]:

3 species in 3 genera or one taxon?3 species in 3 genera or one taxon?

Robert Hermann Schomburgk’s Robert Hermann Schomburgk’s problem: 3 genera on the same plant!problem: 3 genera on the same plant!

Lindley’s response: “Such cases shake Lindley’s response: “Such cases shake to the foundation all our ideas of the to the foundation all our ideas of the stability of genera and species.” stability of genera and species.”

Robert Hermann Robert Hermann SchomburgkSchomburgk

Catasetum macrocarpumCatasetum macrocarpum Rich. ex Kunth Rich. ex Kunth (1822) (1822)

IPNIIPNI Monachanthus viridis Monachanthus viridis Lindl. (1832) Lindl. (1832)

==Catasetum trifidumCatasetum trifidum Hook. (1833) Hook. (1833) Monachanthus viridis Monachanthus viridis Schomb. (1831) Schomb. (1831)

==Catasetum barbatumCatasetum barbatum Lindl. (1844) Lindl. (1844) Monachanthus viridis Monachanthus viridis Lindl.Lindl. (1836 publ. (1836 publ.

1835) =1835) =C. macrocarpumC. macrocarpum FEMALE FEMALE C. tridentatumC. tridentatum Hook. (1823) Hook. (1823) ==C. C. MacrocarpumMacrocarpum MALE MALE

C. tridentatumC. tridentatum, var. viridiflorum (1834) , var. viridiflorum (1834) FEMALE?FEMALE?

Myanthus barbatus Myanthus barbatus Lindl. (1836 publ. Lindl. (1836 publ. 1835) =1835) =C. barbatum C. barbatum BISEXUAL?BISEXUAL?

Catasetum macrocarpum Catasetum macrocarpum Rich. ex Kunth Rich. ex Kunth (1822)(1822)

CurtisCurtis v. 52 (1825) as v. 52 (1825) as C. C. tridentatumtridentatum

CurtisCurtis v. 61 (1834) as v. 61 (1834) as C. tridentatumC. tridentatum

Edwards’Edwards’ v. 21 (1836) as v. 21 (1836) as Monachanthus Monachanthus viridisviridis

Angraecum sesquipedaleAngraecum sesquipedale Thou. Thou. (1822)(1822)

Thouars.Thouars. Histoire particulière des plantes orchidées Histoire particulière des plantes orchidées recueillies sur les trois îles australes d'Afrique recueillies sur les trois îles australes d'Afrique

(Paris, 1822)(Paris, 1822)

Orchid AlbumOrchid Album

Box arrived from Bateman on 25 Box arrived from Bateman on 25 January 1862January 1862

To Hooker: “I have just received such To Hooker: “I have just received such a Box full from Mr Bateman with the a Box full from Mr Bateman with the Astounding Astounding Angraecum sesquipedalianAngraecum sesquipedalian with a nectary a foot long—Good with a nectary a foot long—Good Heavens what insect can suck it”? Heavens what insect can suck it”?

ExperimentsExperiments Bristles and needlesBristles and needles Cylinder to which the pollinia Cylinder to which the pollinia

attached themselvesattached themselves Conjectured that there was a moth Conjectured that there was a moth

with a long proboscis that could get with a long proboscis that could get the nectar from the bottom of the the nectar from the bottom of the nectarynectary

““The astonishing length of the nectary The astonishing length of the nectary may have been acquired by successive may have been acquired by successive modifications. As certain moths of modifications. As certain moths of Madagascar became larger through natural Madagascar became larger through natural selection . . . those individual plants of the selection . . . those individual plants of the Angræcum which had the longest nectaries . Angræcum which had the longest nectaries . . . and which, consequently, compelled the . . and which, consequently, compelled the moths to insert their proboscises up to the moths to insert their proboscises up to the very base, would be fertilised. very base, would be fertilised.

“These plants would yield most seed, “These plants would yield most seed, and the seedlings would generally inherit and the seedlings would generally inherit longer nectaries; and so it would be in longer nectaries; and so it would be in successive generations of the plant and successive generations of the plant and moth. Thus it would appear that there has moth. Thus it would appear that there has been a race in gaining length between the been a race in gaining length between the nectary of the Angræcum and the proboscis nectary of the Angræcum and the proboscis of certain moths.”of certain moths.”

Warner, Warner, Select orchidaceous plantsSelect orchidaceous plants v. 1 v. 1 (1862-65)(1862-65)

John Day scrapbook 039_452_632John Day scrapbook 039_452_632

John Day scrapbook 040-632-44John Day scrapbook 040-632-44

Xanthopan [Macrosila] morgani Xanthopan [Macrosila] morgani Walker Walker (1856)(1856)

subsp.subsp. praedicta praedicta Rothschild & Jordan (1903)Rothschild & Jordan (1903)

Reception to Reception to OrchidsOrchids Slow sales with general publicSlow sales with general public Botanists generally favorable (dialog Botanists generally favorable (dialog

began that resulted in expanded 2began that resulted in expanded 2ndnd ed.) ed.) To Lyell: “Entomologists are enough to To Lyell: “Entomologists are enough to

keep [evolution] back for half a century.” keep [evolution] back for half a century.” George Campbell, Duke of Argyll, George Campbell, Duke of Argyll, The The

reign of lawreign of law Wallace, Wallace, Creation by law Creation by law [measured [measured

Macrosila [Xanthopan] morganiMacrosila [Xanthopan] morgani in the in the British Museum (from South Central Africa) British Museum (from South Central Africa) and found the proboscis to be 7 ½ inches and found the proboscis to be 7 ½ inches long.]long.]

Present entomologists ask: Present entomologists ask: “What, then, pollinates “What, then, pollinates A. A. longicalcarlongicalcar, with a nectary , with a nectary

10 cm. longer than 10 cm. longer than A. A. sesquipedale?”sesquipedale?”

Darwin regarded these theological Darwin regarded these theological views as irritating views as irritating misunderstandings, but wrote to misunderstandings, but wrote to Asa Gray describing his approach describing his approach as a "flank movement on the as a "flank movement on the enemy". By showing that the enemy". By showing that the "wonderful contrivances" of the "wonderful contrivances" of the orchid have discoverable orchid have discoverable evolutionary histories, Darwin was evolutionary histories, Darwin was countering claims by natural countering claims by natural theologians that the organisms theologians that the organisms were examples of the perfect were examples of the perfect work of the Creatorwork of the Creator..

Hermann Müller, Hermann Müller, The fertilisation of flowersThe fertilisation of flowers (1883) with Darwin preface(1883) with Darwin preface

Fritz MüllerFritz Müller

Conclusion of bookConclusion of bookHe had “shown that Orchids exhibit an almost He had “shown that Orchids exhibit an almost endless diversity of beautiful adaptations. endless diversity of beautiful adaptations. When this or that part has been spoken of as When this or that part has been spoken of as contrived for some special purpose, it must contrived for some special purpose, it must not be supposed that it was originally always not be supposed that it was originally always formed for this sole purpose. formed for this sole purpose. The regular The regular course of events seems to be, that a course of events seems to be, that a part which originally served for one part which originally served for one purpose, by slow changes becomes purpose, by slow changes becomes adapted for widely different purposesadapted for widely different purposes.”.”

"In my examination of Orchids, hardly "In my examination of Orchids, hardly any fact has so much struck me as the any fact has so much struck me as the endless diversity of structureendless diversity of structure,—the ,—the prodigality of resources,—for prodigality of resources,—for gaining the gaining the very same endvery same end, namely, the fertilisation of , namely, the fertilisation of one flower by the pollen of another.”one flower by the pollen of another.”

I “found the study of orchids eminently I “found the study of orchids eminently useful in showing me how nearly useful in showing me how nearly all parts all parts of the flower are coadapted for of the flower are coadapted for fertilisation by insects, & therefore the fertilisation by insects, & therefore the result of n. selectionresult of n. selection,—even most trifling ,—even most trifling details of structure.”details of structure.”

Books—from Books—from OrchidsOrchids on on On the various contrivances by which On the various contrivances by which

British and foreign orchids are fertilised British and foreign orchids are fertilised by insectsby insects (1862, 1877, 1882). (1862, 1877, 1882).

On the movements and habits of climbing On the movements and habits of climbing plantsplants (1865, 1875, 1876, 1882). (1865, 1875, 1876, 1882).

The variation of animals and plants under The variation of animals and plants under domesticationdomestication (2 vols.) (1868, 1878). (2 vols.) (1868, 1878).

The descent of man, and selection in The descent of man, and selection in relation to sex relation to sex (1871,(1871, 1874, 1882).1874, 1882).

The expression of the emotions in man The expression of the emotions in man and animals and animals (1872, 1873, 1890). (1872, 1873, 1890).

Insectivorous plants Insectivorous plants (1875, 1888).(1875, 1888).

Books—from Books—from OrchidsOrchids on on (cont.)(cont.) The effects of cross and self fertilisation in The effects of cross and self fertilisation in

the vegetable kingdom the vegetable kingdom (1876, 1877, 1878). (1876, 1877, 1878). – ““A complement to the 'Fertilisation of Orchids,' because it A complement to the 'Fertilisation of Orchids,' because it

shows how important are the results of cross-fertilisation shows how important are the results of cross-fertilisation which are ensured by the mechanisms described in that which are ensured by the mechanisms described in that book.”book.”

The different forms of flowers on plants of the The different forms of flowers on plants of the same species same species (1877, 1884).(1877, 1884).

The power of movement in plantsThe power of movement in plants (1880, (1880, 1881).1881).

The formation of vegetable mould, through The formation of vegetable mould, through the action of worms the action of worms (1881, 1882).(1881, 1882).

The life and letters of Charles Darwin, The life and letters of Charles Darwin, including an autobiographical chapter including an autobiographical chapter (1887).(1887).

Articles on orchids post Articles on orchids post publication of 1publication of 1stst ed. of ed. of OrchidsOrchids ““Fertilisation of Orchids.” Fertilisation of Orchids.” Journal of Journal of

Horticulture Horticulture (31 March 1863): 237.(31 March 1863): 237. ““Appearance of a plant in a singular Appearance of a plant in a singular

place.” place.” Gardeners' ChronicleGardeners' Chronicle no. 33 (15 no. 33 (15 Aug. 1863): 773 [Aug. 1863): 773 [Epipactis latifoliaEpipactis latifolia].].

““Fertilisation of Cypripediums.” Fertilisation of Cypripediums.” Gardeners' Gardeners' ChronicleChronicle no. 14 (6 April 1867): 350. no. 14 (6 April 1867): 350.

““Notes on the fertilization of orchids. Notes on the fertilization of orchids. Annals and Magazine of Natural HistoryAnnals and Magazine of Natural History (Ser. 4) 4 (Sept. 1869): 141-159. (Ser. 4) 4 (Sept. 1869): 141-159.

Darwin’s life after Darwin’s life after OrchidsOrchids

Darwin’s contributions to Darwin’s contributions to botanybotany

Understanding of and ability to Understanding of and ability to demonstrate that the demonstrate that the flower is a flower is a product of evolution product of evolution

Complex ecological relationships Complex ecological relationships resulted in the resulted in the coevolution of orchids coevolution of orchids and insectsand insects

Pollination researchPollination research and reproductive and reproductive ecology (ecology (floral ecologyfloral ecology))

Greenhouse extension project Greenhouse extension project (1862-63)(1862-63)

Asked Hooker for plants: "I long to Asked Hooker for plants: "I long to stock it, just like a school-boy.”stock it, just like a school-boy.”

Sent his butler with a cart to Kew to Sent his butler with a cart to Kew to pick up 160 plantspick up 160 plants

Apologized for depleting the Apologized for depleting the “national collection”“national collection”

To Hooker: "You cannot imagine To Hooker: "You cannot imagine what pleasure your plants give me ... what pleasure your plants give me ... Henrietta & I go & gloat over them." Henrietta & I go & gloat over them."

Select BibliographySelect Bibliography Allan, Mae. Allan, Mae. Darwin and his flowers: the key to natural Darwin and his flowers: the key to natural

selectionselection (New York: Taplinger, 1977) (New York: Taplinger, 1977) The complete works of Charles Darwin onlineThe complete works of Charles Darwin online ( (

http://darwin-online.org.uk/)) Darwin Correspondence Project (Darwin Correspondence Project (

http://www.darwinproject.ac.uk/)) Darwin’s garden: an evolutionary adventureDarwin’s garden: an evolutionary adventure (New York: (New York:

New York Botanical Garden, 2008).New York Botanical Garden, 2008). Pollan, Michael. “Love & lies {orchids}.” Pollan, Michael. “Love & lies {orchids}.” National National

GeographicGeographic 216:3 (Sept. 2009), p. 100-121. 216:3 (Sept. 2009), p. 100-121. Porter, Duncan M. (various journal articles)Porter, Duncan M. (various journal articles) The works of Charles DarwinThe works of Charles Darwin (New York: New York (New York: New York

University Press, 1988). 29 vols.University Press, 1988). 29 vols. Yam, Tim Wing, et al. “’The orchids have been a splendid Yam, Tim Wing, et al. “’The orchids have been a splendid

sport’—an alternative look at Charles Darwin’s sport’—an alternative look at Charles Darwin’s contribution to orchid biology.” contribution to orchid biology.” Am. J. BotAm. J. Bot. . 2009;96:2128-2154.2009;96:2128-2154.

Questions and commentsQuestions and comments

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